STUDIES ON THE GENUS
COENOGONIUM EHREHBERG

 

Them for Hm Dogma 0? pk. D.
MICHIGAN STATE UNIVERSITY
Flordeliz R. Uyenco
1963

THESIS

This is to certify that the

thesis entitled

STUDIES ON THE GENUS COENOGONIUM EHRENBERG

presented by

Flordeliz R. Uyenco

has been accepted towards fulfillment
of the requirements for

M degree in %

(fir-’5‘? fli 19,44 {’21.}, V “L 4

Major professor V

Date 6 me (44}

0-169

LIBRARY

Michigan State

University

 

ABSTRACT
STUDIES ON THE GENUS COENOGONIUM EHRENBERG
by Flordeliz R. Uyenco

The present work is primarily a study of the morphology
and taxonomy of the lichen genus Coenogonium Ehrenberg. This
study was undertaken for the purposes of establishing the
taxonomic bases on which this genus is placed and the characters
by which the species of Coenogonium are separated. It was
hoped that a detailed study of a homoiomerous lichen genus
where the algal symbiont is more conspicuous than the mycobiont
would contribute to the solution of the nomenclatural problems
recently raised by taxonomists as to whether lichen names
belong to the lichen thalli or to the fungal components.

Algal characters such as, the shape and sizes of
the cells and the shape of the thallus, have been used in
the past to identify the species of Coenogonium. This study
was made to determine the value of these characters in the
taxonomy of the genus. The specificity requirements of the
fungus are established in the study: i.e.. whether the

different fungal species are obligately symbiotic with one

Flordeliz R. Uyenco

algal species, or whether the lichen fungi may have different
species of algal symbionts. The disposition of partially
lichenized algae and of sterile thalli are also discussed

and the role of the lichen fungi in determining the types

of thalli is demonstrated.

This investigation is based upon a large number
of herbarium specimens obtained from different herbaria in
the United States, and on fresh material collected in
Florida and Panama. The value of cultural work to supplement
the classical taxonomic methods is considered in order to
establish the role of the phycobionts in the taxonomy of
the genus and to determine which species of Trentepohlia
grow in symbiosis with the fungus. In this study of the
genus Coenogonium, most of the emphasis has been placed
on the lichen fungus morphology; i.e., the structure of the
apothecia and of the spores.

A historical review of the taxonomical, morphological
and physiological studies was made and information has been
added to these from personal observations. The ecology and
distribution of the genus Coenogonium are presented along
with a distribution map of the species.

Keys to the species of the algal symbionts and of

the lichen genus Coenogonium have been written, the latter

Flordeliz R. Uyenco

based on some characters not previously considered by any

author. Six species of Trentepohlia and one species of

 

Physolinum are included in the key. Of the original forty-

 

eight species referred to as Coenogonium by earlier authors,

 

eighteen Species have been recommended for transfer to the

genus Trentepohlia as valid algal species or as synonyms;

 

four species are considered as belonging to the Fungi
Imperfecti and two species are treated as synonyms of

Coenogonium Linkii. Eight species are considered doubtful:

 

fifteen species are recognized and accepted as valid and a
key has been made for these. One species is rejected as
having been based on two discordant elements.

Eleven plates containing forty—five figures, five

tables and eight maps supplement the written text.

STUDIES ON THE GENUS COENOGONIUM.EHRENBERG

 

BY
\(rn
(o-

Flordeliz RT Uyenco

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Botany and Plant Pathology

‘

1963

g5 3951‘s
2 wag-ai-

ACKNOWLEDGEMENTS

It is my privilege here to express my sincere
appreciation and gratitude for the valuable assistance
and criticisms offered by Dr. Henry A. Imshaug under whose
guidance this work was carried out. It is equally a great
pleasure to thank him for the use of his library and lichen
collections from the West Indies.

I also wish to thank the members of the Guidance
Committee for their interest and cooperation during the
progress of this work and for taking the time and trouble
to concern themselves with the present problem.

I am especially indebted to the Agency for Inter-
national Development, U. S. Department of State, under
whose training program this investigation was financed and
carried out.

Grateful appreciations are extended to the Botany
Department, University of Florida and to Miss Carol C.
Beck, Botanist of the Highlands Hammock State Park, Florida,
for providing me their valuable assistance and facilities

during my collections trips in those areas.

ii

I extend my special thanks to Dr. and Mrs. Frank C.
Craighead who unselfishly gave much of their time to accompany
me to the hammocks of the Everglades National Park, Florida.

Sincere thanks are due Mr. I. Brodo, Mr. D. Jackson,
Department of Botany and Plant Pathology, Michigan State
University, and Miss Anita Alejo for their assistance in
the preparation of the photographs and illustrations
included in this paper.

Lastly, it is my most important wish to acknowledge
the generosity of the curators and directors of the numerous
herbaria and libraries who have cooperated with us in supply-
ing their materials and important reference books without
which it would have been impossible to fulfill and_realize

the objectives of this work.

iii

TABLE OF CONTENTS

INTRODUCTION

Purpose of the Study
Historical Review

THE LICHEN THALLUS

Morphology of the Thallus

Lichen Substances
Ecology
Distribution

THE ALGAE

The Lichenized Algae
The Algae in Culture

.Materials and Methods

Observations
Results
Discussion'
Conclusions

Taxonomy

History

Morphology of the Phycobionts

Key to Species
Systematic List

Trentepohlia
Trentepohlia
Trentepohlia
Trentepohlia
Trentepohlia
Trentepohlia

abietina

arborum
aurea
elongata
odorata
umbrina

iv

Page

H

15

15
17
20
26

34

34
35

38
41
45
45
57

57

57
62
69
71

71
72
74
77
78
79

THE FUNGUS

Morphology

Thallus
Apothecia
Pycnidia

Taxonomy

DISCUSSION

SUMMARY

General
Lichenized Hypomycetes
Non-Lichenized Trentepohlia

Partially—Lichenized Trentepohlia

 

 

 

Lichenized Coenogonium
Key to Recognized Species

Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium
Coenogonium

BIBLIOGRAPHY .

MAPS

PLATES

acrocephalum

botryosum
complexum
curvulum
disjunctum
implexum
interplexum

interpositum

Leprieurii
Linkii
moniliforme
ornatum
pannosum
tomentosum
velutinum

Page
82
82

82
83
93

96

96
99
100
101
103
107

110
111
112
113
114
116
118
121
124
132
138
141
142
143
144

146
153
155
170

177

Table

LIST OF TABLES

 

Page

Distribution of Coenogonium species in the

different geographical areas . . . . . . . 32
Summary of phycobiont-mycobiont associations . 46
Comparison of cell sizes in lichenized and

non—lichenized phycobionts arranged

according to algal species . . . . . . . . 48
Comparison of cell sizes in lichenized and

nonélichenized phycobionts arranged

according to fungal species . . . . . . . . 50
Data of collections from Florida and Panama . . 55

vi

INTRODUCTION

Purpose of the Study

This study was made for the purpose of establishing
the characters by which Coenogonium species should be
determined. In the past, workers have used the characters
of both the alga and the thallus for the determination of the
lichen-fungus. It was hoped that a detailed study of a lichen-
fungus where the alga assumed a major role in lichen morphology
would contribute to the solution of the nomenclatural problems
recently raised by taxonomists as to whether lichen names
should belong to the lichen thalli or continue to belong
only to the fungal components.

Since the form of the thallus is mainly that of
the alga itself, it was also the objective of this investi-
gation to determine whether the different fungal species
are symbiotic with only one definite algal species, or
whether these lichen-fungi may form lichen thalli with various
species.

The species of Coenogonium have been distinguished

in large part on the shape and size of the algal cells.

It is possible, however, that the "species" named in such
a manner may actually include different lichen—fungi,
especially since many lichens were described in their
sterile forms.

It was further necessary to demonstrate whether the
fungus determines the shape of the thallus or if different
species of algae form different thalli. The extent of the
lichenization of the species of algae as well as environmental
conditions were studied to determine their role in the
different external forms of the lichen thallus. Only if the
thallus characters were determined by the fungus could they
be used in identification.

In addition to the basic questions concerning the
nature of the species in Coenogonium, it was hoped that
a workable key to the species of this genus could be
prepared. Therefore. descriptions and illustrations of all

the species examined were prepared.

Historical Review

The genus Coenogonium was founded by Ehrenberg in
1820 with the single species 9, Linkii which was collected
from the bark of trees at Santa-Catharina Island off the

coast of Brazil. Ehrenberg observed in the apothecia of this

genus the same structures found in the fructifications of

Peziza and other fungi: that is, the surface of the apothecium

is formed by elongated capsules that are stalked and con-

taining spores which are not bare, contrary to what

Acharius (1798) had supposed. Ehrenberg further observed in

this genus the same organization found in the other lichens.
Coenogonium has been shifted in the Plant Kingdom

from one group to another, and variously classified by

systematists as alga, fungus or lichen. A study of the

original descriptions of the forty-eight species belonging

to this genus mention Chroolepus as the gonidial (algal)

 

symbiont. Early authors described the thallus as consisting
of confervoid or cylindrical cells with yellowish-green
contents surrounded by colorless slender threads. Wainio

(1890) mentioned Trentepohlia species as constituting part

 

of the thallus of Coenogonium.

 

Much of the early investigations on the genus
Coenogonium involved principally the system of classification,
treating the algal and fungal symbionts as of equal taxonomic
importance. The genus was placed in a confervoid algae by
Agardh in 1824, a classification which was considered
erroneous by Fée (1824) on account of the organization of the

thallus, it being composed of cylindric fibers and attaching

itself on the substratum by its side, in a manner of certain
fungi. Fee, (op. cit.) after studying the asci, proposed

that the Family Corniculariae, previously used by Fries (1825),
include the genus Coenogonium. Fries (op. cit.) assigned

the genus Coenogonium to the Tribe Coenogoneae and the

related genera Collema, Leptogium and Lichina to the Tribe

 

Collemaceae and assembled the whole group under the Family
Byssacées. His classification based on the fungus characters
of these lichens greatly influenced nearly all subsequent
lichen systematists, the first of whom was Endlicher (1836)
who included the genus Coenogonium in the Tribe Collemaceae
established by Fries, disregarding the Tribe Coenogoneae
to which the genus originally belonged. Endlicher then
grouped the other genera in his new Tribe Lecidineae.
Montagne (1846) considered Coenogonium as a degenerate
lichen and therefore an anomalous genus and included it in
the Family Byssacées designated by Fries to unite closely the
lichens and the algae. Montagne, however, believed that the
Family Byssacées emphasized the characters of the fungus
too much and suggested that the Subfamily Collémacées be
established to define the lichen properties of these plants
more clearly. Montagne was the first to separate the plants

whose ascocarps are open at maturity from those whose ascocarps

remain closed and established the Orders Gymnocarpes.and
Angiocarpes to apply to these plants respectively. He

extended his analysis of Fries' classification system by
reorganizing the genera under the Family Byssacées into

three tribes; namely, Collémacées, Cénogoniées and Lichinées.
Montagne assigned the genera Collema, Leptggium, Stephanophorus_
and Omphalaria to the Tribe Collémacées: the genera Coenogonium,

Cilicia, Ephebe and Thermutis to the Tribe Cénogoniées; and the

 

genera Lichina and Paulia to the Tribe Lichinées. Payer
(1850) adopted Fries' classification in part, and included
the Tribes Collemaceae and Coenogonieae under the Family
Lichens next to the Family Pezizes.

The lecidioid characteristic of the apothecium of

the genus Coenogonium led Nylander (1858, 1862) to believe

 

in the close relationship between this plant and the Lichineos
and, following Payeris scheme, placed the genus in the Tribe
Lecideinei under the Series Placodei of the Family Lichenacei.
Nylander's classification system and the studies of
Schwendener (1860) on the nature of lichens had important
consequences on the taxonomy of the lichens.

De Bary (1866) considered Montagne's classification
as a concentration of these plants in the fungi stressing
the significance of the lichen characters that the genus

Coenogonium possessed.

Thwaites published in 1849 a new genus of minute
plants which he called Cystocoleus, to contain a form of

Byssus nigra Huds., also called Chroolepus ebeneus. He

 

described the thallus as growing by means of apical growth
and consisting of rows<ofattenuatecylindrical cells with
colorless cellulose walls and green—colored contents.

These cells are surrounded by a dense covering of dark brown
hyphae with rough walls and colorless contents. Hariot in
1890 recognized the composite nature of this plant and ex—

cluded Chroolepus ebeneus (=sttocoleus ebeneus) from the

 

 

genus Trentepohlia. He also noted that Persoon's Racodium
rupestre was a synonym. De Bary, likewise, regarded the

Cystocoleus of Thwaites as synonymous with Racodium

 

rupestre Pers., and correctly described it as associated

with Chroolepus. Glfick later in 1896, treated Thwaites'

 

material and called it Coenogonium germanicum, noting that

the thallus consisted of Chroolepus filaments. It is thus

 

evident that this species had been recognized primarily on
the basis of the algal cells or on the color of the sur-
rounding hyphae.

Karsten (1861), Schwendener (1862) and Nylander
(1862) contributed much to the studies on the morphology

of the genus and described several species, some of which

were based on the algal constituents of sterile thalli.
Flotow (1869) in recognizing the systems of Fries
and Nylander, divided the lichens into two series:
Lichenes Heteromerici and Lichenes Homoeomerici, both of
which, he believed, incorporated the principal characters
on which the Family Lichens of Payer and Family Byssacées of
Fries, as well as the Family Lichenacei of Nylander were
based.
Tuckerman, in 1872 and later in 1882, organized the

genus Coenogonium under the Series Gymnocarpi in which the

 

apothecia are normally open and the spores are exposed. He
established the Tribe Lecideacei under this Series to include
those with free, round, open and disk-like apothecia bordered
by a proper exciple. The Family Coenogoniei was placed under
this Tribe to separate those with horizontal thallus and
confervoid filaments. Wainio, in 1890, however, raised

the Family into the Tribe Coenogonieae and established the

Family Coenogoniaceae to include the single genus, Coenogonium.

 

Mfiller (1881), in discussing Coenogonium pannosum,

 

stated that the algal symbiont of this lichen belongs to

the Graphideen and that the genus Coenogonium is closely

 

related to the genus Biatorinopsis Mfill. Arg. on the basis

of their spore characters.

Shirley (1889) designated the Tribe Lecideinei in
classifying the genus Coenogonium and emphasized the absence
of the cortical tissue usually found in lichens.

Wainio (1890), Lindau (1895) and Clements (1897)
incorporated the genus Coenogonium in the fungi as evidenced
by the schemes of classification that they presented.
Wainio, in organizing the Class Ascophyta, assembled the
Discolichenes and the Discomycetes in one group called
Gymnocarpeae and the Pyrenolichenes and the Pyrenomycetes
in the Pyrenocarpeae. He followed Montagne's arrangement of
the tribes, restoring the genus Coenogonium into the Tribe
Coenogonieae. He justified his scheme of classification
on the homoiomerous structure of the thallus. The_two types

of spores of the genus Coenogonium became the basis later

 

in the establishment of two Sections of the genus under the
Tribe Coenogonieae. Clements in 1909 classified the genus
under Order Pezizales, separating those plants with Chroolepus
constituting their thalli as belonging to the Family Chryso—
trichaceae, and further dividing the genus into the Section
Holocoenis to include those with one-celled spores and the
Section Coenogonium to comprise those with one septate
espores. Wainio later in 1921, extended his analysis of the

Family Coenogoniaceae, still upon the system he established

in 1890, and divided the genus into two sections: Section
Coenobiatora, to include all the species having one-celled
spores, and the Section Coenobiatorina, for those with two-
celled or one-septate spores. Lindau (1895) followed the
classification of Tuckerman but proposed the addition of the
Series Gymnocarpaceae to include those whose hymenium do
not open with a pore but remain closed and spread out only
at maturity. He organized the sub-group Cyclocarpineae
under the Gymnocarpaceae and re-established the Family
Coenogoniaceae to include the single genus Coenogonium

and excluded the Tribe Coenogonieae in his classification
of the genus.

Clements (1897), in his study on the polyphyletic
disposition of the lichens, recognized the importance of the
spores and placed the genus Coenogonium in the Family
Chrysotrichaceae. He and Wainio later divided the genus
into the.two Sections as described above. This scheme was
adopted by all subsequent workers of this genus.

Reinke (1894) and Schneider (1905) had opposed the
classification systems organized by the earlier workers
arguing that these incorporated the lichens in question in
the system of the fungi.

In the lichen taxonomy of Zahlbruckner (1905, 1926)

10

which is now generally used, the Coenogoniaceae and related
families are placed among the Cyclocarpineae (a position
first used by Lindau) which, to a great extent, are arranged
according to their algal hosts. Zahlbruckner placed the

genera Coenogonium and Racodium together in the Family

 

Coenogoniaceae on the bases of their confervoid algal symbionts
and on the homoiomerous character of their thalli. These
genera are very similar in their appearance and habitat but

are distinguished by their algal symbionts. In Racodium,

 

the investing fungus lies in straight unbranching lines

along the Cladophora filaments, while in Coenogonium, the

 

dark fungal hyphae branch repeatedly, and wind around the

 

irregular bulging cells of the Trentepohlia filaments.
Zahlbruckner classified the Family Coenogoniaceae under the
Ascolichenes and included the Sections Holocoenis and
Coenobiatorina in distinguishing the different species in

the genus. According to Zahlbruckner, Coenogonium, Cystocoleus

 

and Racodium form an autonomous family, the Coenogoniaceae.
Santesson (1952) disagreed to this idea and suggested that
Racodium and Cystocoleus must be transferred to the lichenized
Fungi Imperfecti, while Coenogonium is very closely related

to Dimerella and should therefore be included in the

Gyalectaceae. Zahlbruckner constituted the Family Gyalectaceae

11

mainly on the algal characters of the lichens and included

in it genera characterized by the presence of Trentepohlia,

 

Phycopeltis, and Scytonema as phycobionts.. Santesson,

however, believes that some genera referred to the Gyalectaceae
by Zahlbruckner are probably more closely related to the

genera of other families than to those forming the natural

center of the Gyalectaceae; such as Gyalecta, Dimerella,

 

Microphiale and Ramonia. Santesson grouped the genera

Dimerella, Coenogonium, Gyalecta and Semigyalecta together

 

in the Family Gyalectaceae. He maintains that the anatomy
of the apothecia and the characteristic thin-walled asci
of these genera prove the very close relationship that exist

among them. The close relationship between Coenogonium and

 

Dimerella was reported as early as 1881 by Mfiller who
stated that Coenogonium is closely related with his genus
Biatorinopsis (=Dimerella) on the basis of their spore
characters. Malme (1937) concurred with the statement of
Mfiller and used spore characters to delimit the species of
this genus.

Smith (1906), in examining the British forms of
Byssus, Racodium and Cystocoleus, believed that these genera
were associated with Chroolepus and therefore included them

in the genus Coenogonium.

12

The remaining studies on Coenogonium have been
primarily systematic. Hooker (1867), Leighton (1869),
Krempelhuber (1876) among others, have added new species,
or listed and redescribed old ones from various tropical
and temperate regions throughout the world. Very few
fundamental studies on the structure and development of
the Coenogonium have been added.

The first description of the development of the
apothecium of Coenogonium was made by Karsten in 1861 who
described the fertilization apparatus of this plant as being
analogous to the archegonia of the higher cryptogams. His
work was severely criticized by Schwendener and Nylander who
considered Karsten's descriptions to be without foundation
since he did not examine the plants microscopically.

Fink (1915) adhered to the Class Ascolichenes of
Zahlbruckner and placed the Family Coenogoniaceae within his
new large group Lecanorales which only included lichens.
This arrangement, in reality, is the same as the group
Cyclocarpineae in the pure lichen system of Zahlbruckner.
Fink's classification is based largely on the algal hosts
and considered Coenogonium and Racodium as related to
those belonging to Gyalectaceae and Ephebaceae. Nannfeldt

(1932) later used Fink's Order Lecanorales in a somewhat

13

wider sense in his new system of the Ascomycetes where it is
included as a special group among the Ascohymeniales.

Dodge (1933) placed Family Coenogoniaceae between
Rocellaceae and Ephebaceae, both of which have Sgytonema
as the algal symbiont. He suggested, however, that the
Family Coenogoniaceae would be better dropped and the
species distributed among the fungi and algae. He considers
this family very aberrant among the lichens in several respects

and recognizes the relationship of Coenogonium to Patinella

 

and Orbilia. Santesson (1952), on the other hand, disagreed
with Dodge and finds, instead, that Coenogonium is very

closely related to Dimerella and Gyalecta, and must be

 

 

included in the Family Gyalectaceae. The diagnostic value
of the apothecium in deciding the speciation in Coenogonium
has been emphasized by Santesson and he proposed that sterile
specimens be considered as lichenized Trentopohlia.

A more exact knowledge of the composition of the

genus Coenogonium led to the various classifications in which

 

it has been placed by subsequent investigators. On account
of the presence of oil drops containing carotene in their
cells, two algal symbionts, Trentepohlia spp. Mart. and

Physolinum.mgnil§ Printz, were recognized. Chroolepus

 

was later used as a section of the genus Trentepohlia

 

14

under the Family Trentepohliaceae by Printz in 1939.

Collins (1928) in discussing the genus Trentepohlia,

 

indicated that the terrestrial habit of the species of
Trentepohlia makes them very accessible to lichen—forming
fungi, and in many cases, it is rather an exception to
find an alga that is completely free of fungi. Collins
described g. rigidula (Mfill. Arg.) Hariot as a species
always epithallic on lichens and which may be attached
either to rocks or to the bark of trees. De Toni (1889)

mentioned Trentepohlia villosa as associated with Coenogonium

 

confervoides Nyl. and Trentepohlia monilia de Wild. associated
with Coenogonium rigidulum Mfill. Arg.

After reviewing the literature on the genus
Coenogonium, the writer believes that a critical evaluation
of the morphological characteristics used in the classifi-
cation of Coenogonium is necessary. It can be seen that
in the Family Coenogoniaceae, the criteria in determining the
species have emphasized thallus morphology and algal measure-
ments, relegating the apothecium and the other fungal characters
to a minor role without the proper understanding of their

significance and value as a taxonomic feature.

THE LICHEN THALLUS

Morphology of the Thallus

The external form of the thallus of Coenogonium

 

Species is quite variable. Coenogonium Linkii, on which
Ehrenberg constituted the genus, is characterized by a loose,
spongy-byssoid thallus, its filiform and branched filaments
forming a more or less determinate round or nearly reniform
and flat tissue. The thallus is firmly attached to the
substratum by one margin from which it grows erect or
pendant. Occasionally, the Trentepohlia filaments composing
the thallus become interwoven into a more or less parallel
tissue, giving an appearance of a zonate, lamellate and
radially-organized thallus. In 9, Linkii, the algal filaments
are evidently bound together so that at a later stage, the
plant becomes composed of many imbricate thalli that are
thinner towards the periphery.

In addition to the common and typical shelf-like

forms of g. Linkii and g, Leprieurii, the thallus may be

 

cushion-shaped and round, with the filaments comprising

it firm, straight and closely aggregated: or it may be

15

16

crust—like, as in Q. epiphyllum; or densely—velvety, appearing

 

like a dark felt of short hairs, as in C, interplexum. In

 

 

several species of Coenogonium, the thallus may be partly or
wholly adnate to the surface, occasionally expanded and
effused and exhibiting a horizontal growth. Other species
may be prostrate, adhering to the substratum as very thin,
delicate and widely spreading filaments.

More rarely, the thallus of Coenogonium species
forms small, erect and soft turf-like, homoiomerous growths
on the surface of the substratum. In 9, moniliforme, in
which the algal symbiont is Physolinum monile (de Wildem.)
Printz, the effused thallus is made up of short, moniliform
filaments. In still other Coenogonium species, the

branching threads of Trentepohlia are loosely woven together

 

to form an adnate, irregularly tomentose and diffused
thallus.

The upper surface of the thallus of Coenogonium
is generally smooth, but it may also appear as if it were

trimmed and wrinkled, as has been observed in g. pannosum.

 

The color of the thallus varies from yellow to
golden-yellow, or green to yellow—green. It has been
found that the color of the thallus is of no significance

since the plant loses much of its color upon drying, as does

17

Trentepohlia. The filaments of the thallus of Coenogonium

 

do not imbibe water easily, and it is particularly this
characteristic that differentiates this genus from all the
other lichens and provides its habitat restrictions.

Since the thallus is homoiomerous, there are no
cortical, medullary or algal layers. The filaments of

the algae are merely surrounded by fungal hyphae.

Lichen Substances

Attempts by the writer to identify lichen substances
in the thallus and apothecia of the different species in the
genus Coenogonium by the use of the standard chemical
reactions were unsuccessful. Early investigators reported
color reactions of the hymenium of certain species to dilute
iodine solution, but the results are inconclusive.

Sato (1933) noted that the hymenium of Q. boninense
turned wine red in iodine. The same color reactions were
observed by Nylander (1862) in the hymenium of Q. implexum and
by Wainio (1896) in Q. Leprieurii and Q. subvirescens.
Zahlbruckner (1909) and Malme (1937) reported dark red
color reactions of Q. interpositum and Q, velutinum with
dilute iodine, while the hymenium of Q, Leprieurii turned

sordid red upon treatment with the same chemical. Tuckerman

18

(1862) reported that Coenogonium moniliforme gave a deep
green color reaction with iodine but did not react with
potassium hydroxide. Zahlbruckner (1909) further reported
the hymenium of g. curvulum turning at first blue but later
to yellow with iodine. Wainio (1921) indicated a dark blue
color in the hymenium of g. gpiphyllum when treated with
iodine and a wine red color reaction in g. subvirescens.
Coenogonium disjunctum and Q. Linkii showed no color reactions
with iodine, according to Nylander, and he added that g.
interplexum changed to dark violet when treated with the
same solution.

The absence of color reactions with paraphenyl-
enediamine and potassium hydroxide when applied to the
thallus and apothecia of the different Coenogonium species
indicate that these plants do not possess the depsides and
depsidones of more advanced lichens. It is believed that
the different color reactions of the hymenium with iodine
solution cannot serve as a taxonomic method in distinguishing
the species. Dughi (1942) stated that there are in reality
no negative reactions with iodine on lichen tissues provided
that the concentration of the iodine is sufficient in
every case. A positive and negative reaction to iodine

refer to the Special concentration of the chemical and the

19

degree of reaction is the same in all specimens of the

same "species" or "variety" and is independent of their age;
or from thallus to thallus and even in different parts of the
same thallus. In the present study, the different color
reactions of the apothecia to iodine can be ascribed to the
concentration of the iodine solution used. It can be in-
ferred, however, that Coenogonium, being a homoiomerous
lichen, does not possess an advanced type of thallus which
might explain the absence of lichen substances commonly

found in the more advanced forms.

Degelius (1954) found in his studies of the lichen
genus Collema that the constancy of the reaction of the
NOstoc alga to iodine varies from one species to the other.
He remarked that the positive reaction depends on the
presence of glycogen and has nothing to do with the fungus.

It is the opinion of the writer that the use of
chemical reactions in separating the species in many other
lichen genera cannot be used with the genus Coenogonium.

A polyhydroxy alcohol or sugar alcohol has been
reported in certain species of Trentepohlia. This 4—carbon
compound, known as erythritol, is usually present up to a
concentration of 1.4% in the cells (Lewin, 1962). This

compound contains a greater amount of photosynthetic energy

20

("reducing power") than do the corresponding sugars.

Several Species of Trentepohlia have been reported

 

positive for cellulose I, an important constituent of the
cell walls. No sterols have so far been detected in any
species, a condition that is considered anomalous Since this

plant reproduces sexually.

Ecology

The lack of complete ecological data on the
collectors' labels presented some difficulties in the explan-
ation of the ecological problems relative to the distribution
of this lichen genus. In many instances, the exact location
and substratum were not specifically defined. For example,
9, Leprieurii bore only such information on the herbarium

label as "On trees" or "On forest slope,‘ and very often,
only the country from which the material was collected was
given, the specific locality having been omitted. These
data would have been useful in interpreting the ecological
distribution of the Species of Coenogonium.

All altitudes and elevations have been converted
to meters and insofar as possible, the scientific names of

the tree Substrates were obtained.

An analysis of all the habitats reported for the

21

genus Coenogonium reveals that this is mainly a tropical

 

genus belonging almost exclusively to warm, humid regions.
It is widely distributed and occurs principally on the

bark of trees, often associated with mosses and hepatics.
Generally, damp localities which receive continuous precipi-
tation are suitable places for the growth of this lichen.

Fée (1824) mentioned Coenogonium as growing on the leaves of

 

cinchona and angostura trees in Brazil. Santesson (1952)
listed seven foliicolous Species, two of which he considered
as probably more or less obligately foliicolous lichens and
the others as only facultatively foliicolous.

Since climatological data for the habitats reported

 

for Coenogonium are not available for all the collections,
it is not possible to make a detailed discussion on the
habitat requirements of this genus, but generally speaking,

it can be noted that the species of Coenogonium range in

 

ecological distribution from high altitudes to sea level.
These plants do not exhibit great tolerance to intense
sunlight as evidenced by the fact that they occur abundantly
in shaded areas, on the lower portions of the trunks of old
trees and on the under Side of branches and leaves of living
trees. Species growing on leaves are characterized by

their delicate, veil—like thin thalli which are so densely

22

compressed on the under surface of the leaves that they are
distinguished from one another with difficulty.

Coenogonium is rarely found on the surface of damp

 

rocks. Two collections from Cuba and the Philippines were

identified as Q. Leprieurii and g, disjunctum, respectively.

 

 

The Species of Coenogonium grow abundantly in

 

tropical rain forests. In the drier parts of the tropical
regions, they are commonly found in the montane mist forests
where the increased elevations not only compensate for a
drier climate but also provide moisture in the form of mist.
Regions where dense forests prevail and where the trees are
Sheltered from wind and sunlight yield a variety of species

of Coenogonium. Many forms of this lichen inhabit trunks,

 

branches and twigs of living trees. 9, interplexum has

 

been collected from a dense wet limestone forest in Guatemala
at an altitude of 900-1500 meters. The forest slopes of
Cuba, with an altitude of 600—700 meters, yielded a

rich collection of C, Linkii and g. Leprieurii which were

 

found growing among mosses. An apparently similar slope

in China also yielded 9, Leprieurii at an elevation of 480

 

meters. It is interesting to note that g, Leprieurii

 

seems to be the most cosmopolitan Species in the group in

that it occurred in all levels of altitude, ranging from

23
10 meters, in Costa Rica, to 2000 meters, in the Philippines.
It grew on the banks of rivers in Brazil and the flood plains
of Rio Sandalo in Costa Rica. Coenogonium Leprieurii was
collected from exposed roots of a forest tree in a deep
ravine in Jamaica at altitudes of from 840 to 1524 meters.
It was likewise found on the bark of a palm tree and on
evergreen trees in British Honduras and Mexico, respectively.
A Spruce collection from the Amazon basin in Brazil showed a
good growth of this species on leaves.

The bases of trees in the swampy woods of Louisiana
provided good growth of g. interplexum, while Q. Linkii
dominated the growth on the bark of oak and other deciduous
trees in Florida. Coenogonium disjunctum occurred in the
summit forest of Panama and Costa Rica (about 1300 meters
elevation) and was also collected from the twigs and bark of
large trees in the wet forests of Mexico and Honduras.

Coenogonium curvulum, a species reported by
Santesson (1952) to be probably more or less obligately
foliicolous, was growing on bark in a collection from Puerto
Rico. The wooded ravine of Trinidad, Puerto Rico, the
Dominican Republic and Haiti yielded the greatest variety
in Species of Coenogonium.

A noteworthy comparison of the growths of g. Leprieurii

24

and_g. Linkii can be made on the collections from a dense
forest in Peru. Coenogonium Leprieurii occurred at 625—
1100 meters altitude but at higher elevations, 1067-1900
meters, 9. Linkii dominated the collections. This latter
species also exhibited an interesting growth response in
that it occurred at lower elevations (750—1000 meters)

in open woods, but required higher altitude for its growth
in dense forests. A similar Situation in the occurrence of

both C. Linkii and Q. Leprieurii was observed in the Dominican

 

Republic where the latter species occurred at 200-500 meters
elevation while g. Linkii grew at 900-1220 meters altitude.
This was likewise observed in collections from Haiti and
Puerto Rico where Q. Leprieurii tended to grow in lower
elevatiOns while 9. Linkii was found in the higher altitudes.
Both Species, however, grew in the same range of altitudes
in Cuba (600-700 meters) and in Colombia (1760-2300 meters).

Coenogonium was found frequently in the hammocks

 

of Florida. This habitat consists primarily of a community
of broad-leaved evergreens and large deciduous trees with
occasional pines. Such habitats are raised somewhat above
surrounding, usually wetter, areas made up of swamp forests
and marshes. These mesic habitats provide subtropical

conditions of high moisture throughout the year and little

25

variation in temperature.

Four different Species of Coenogonium were collected
from habitats in Florida ranging from solution Sinks to
dense and low hammocks. Many collections came from the
bark of trees, including Qggrcus virginiana, magnolias,
hollies and some tropical species such as Ficus aurea, Lyssoloma,
Bursera and the prominent palm (Sabal). Coenogonium was
found primarily on the lower portions of the trunks of these
trees, associated with mosses and hepatics. They would also
be found on the smaller branches if there was enough shade
and moisture.

Coenogonium moniliforme,also reported by Santesson

 

(1952) as a facultatively foliicolous species, was collected
from the bark of Lyonea and other deciduous trees in the
hammocks of Florida. A collection of the same species was
made from a dead branch (of a living tree?) in Guatemala.
An ecological distribution of this Species cannot be
accurately made due to the inadequacy of the collection.
This Species has an almost crustose thallus and hence is
easily overlooked in the field.

Coenogonium species are not commonly found associated
with other lichens. The spreading, horizontal, and sometimes

prostrate growth of this plant on the substratum probably

26

provides an unfavorable shade to other lichen thalli and
so hamper their development. Coenogonium grows in shadier
and wetter habitats than tolerated by most other lichens.
Few works have dealt on the ecology of the lichen
genus Coenogonium and much less is known about the cor—
relation between the ecological factors and the fruiting
stages of this plant. The correlation between such
ecological factors and the distribution of the Species
would provide an interesting study by future investigators

of this group of plants.

VDistribution

The only Coenogonium hitherto recorded from conti-
nental Europe was found by Hugo Glfick in Saxony and the Harz

in 1896. Glfick named it Q. germanicum. This plant grew

 

abundantly on a siliceous substratum and attained a con-
siderable size, in one instance, up to 1.5 meters. Santesson
(1952) pointed out that this plant is a lichenized Hyphomycete
with dark brown hyphae and is probably a synonym of Cystocoleus
nigg; (Huds.) Hariot or Racodium rupestre Pers.

A similar situation exists in the case of g, ebeneum
Smith, originally described to occur on rocks in Great

Britain and which Santesson (1952) considers to be synonymous

27

with Q. nigrum (Huds.) A. Zahlbr. It is also a Hyphomycete

and, therefore, cannot be referred to the genus Coenogonium.

 

Hale and Culberson (1960) reported four Species of

Coenogonium from the United States and Canada: .9. disjunctum,

 

 

 

g, interpositum, Q. Linkii and Q. moniliforme.

 

The present Study covers North America, Central
America, west Indies, South America, Australia, New Zealand,
Eastern Asia and West Africa. The material examined was
drawn from the collections available in the herbaria of
Michigan State University, University of Michigan, University
of Washington, University of Florida, Missouri Botanical
Garden, the United States National Herbarium and the New
York Botanical Garden. Some Specimens were collected
Specifically for this Study by the writer in Florida.

Living collections were also provided for this work by Dr.
G. W. Prescott from Panama, Dr. D. B. Ward from Florida

and Dr. A. J. Sharp from Mexico: the cooperation of these
botanists is gratefully acknowledged. Specimens were
placed in polyethylene bags and mailed in regular envelopes.
It was found that Specimens of Coenogonium remained viable
for at least a week or ten days after they were collected.

Since the identification of the species of Coenogonium

 

was based only on fertile specimens, the distribution of the

28

Species cannot be entirely accurate. The relative abundance
of each species could not be determined due to the lack of
collections in some of the regions, the unequal intensity of
the collecting and the sterile condition of much of the
material. It may be noted that some areas were thoroughly
collected but the materials were unidentifiable and could

not be represented on the distribution map. As illustrated
in the map, the West Indies have been more thoroughly covered
than any other area included in this study. Coenogonium

Leprieurii appears to be the most abundant species in this

 

region and it occurs in almost all the islands, commonly on
the bases of trees in dense thickets. Coenogonium Linkii,
however, is the most widely distributed Species, occurring
in all the tropical countries covered in the study.

The indicated distribution of the genus in North
America Shows clearly that it is entirely southern in
range, with the exception of one collection of g. interplexum
from southern Michigan. A sterile Specimen was seen from
BE§%: River, Jefferson County in New York State. These two
localities are certainly unusual for this lichen genus. Other

species of Coenogonium are found along the southern Atlantic

 

Coastal Plain and along the Gulf Coastal Plain to Texas.

Coenogonium Leprieurii has been mentioned as

29

capable of maintaining itself at all altitudes and thus

has a wider range of distribution than all the other species.
Although it has not been reported as occurring in the United
States, it cannot be safely said that it is absent in this
region as it may have never been encountered in a fertile
condition. According to the herbarium data, this species,
however, was the most abundant in any of the regions where
it has been reported. Coenogonium Linkii ranks second in
relative abundance and was particularly common in the West
Indies.

Coenogonium velutinum was identified from Jamaica

 

and the Juan Fernandez Islands in the Pacific Ocean off
the southern coast of South America, indicating a wide
geographical range for this species. Coenogonium interpositum

is as widely distributed as g, Leprieurii and Q. Linkii but

 

did not show the same relative abundance in the collections.
This may be partly due to the unequal intensity of the field
work or the absence of fruiting bodies in some of the material.

Although Q. botryosum is reported only from Mexico, it is

 

possible that it occurs also in the other geographical areas
for the same reasons stated above. If some Species of
Coenogonium have seasonal production of apothecia, the

collectors might not have collected the thalli at the

30

appropriate time and this might explain the paucity of fertile
material of certain Species.

Coenogonium acrocephalum, Q. tomentosum, Q. curvulum

 

 

and g, pannosum were identified only in collections from
Puerto Rico, Bahamas and Guadeloupe, all of which are in the

West Indies. g, acrocephalum and g, pannosum were first

 

collected from Brazil while Q. tomentosum was first reported

 

from New Zealand. Q, curvulum was found growing on leaves
in Java by Schiffner.
The tropical areas in Eastern Asia show a discon-

tinuous distribution of the genus Coenogonium, perhaps due

 

to the lack of collections. Coenogonium Leprieurii is
common to all the collections from Borneo, Ceylon, China
and the Philippines. The range of distribution of this
species can be gleaned from the fact that it also occurred
in collections from Australia and West Africa.

The presence of pycnidia was noted in some of the
collections but Since this reproductive structure cannot be
used to determine the Species, the collections could not be
included in the distribution map. The infrequent occurrence
of thalli with asexual fruiting stages may be related to

the environmental conditions. It is believed that Coenogonium

 

species do not display great versatility in their methods of

31

reproduction, a feature, perhaps, of a homoiomerous lichen.
Long distance dispersal of Coenogonium is believed to be
achieved principally by the sexual spores which are

hardly complemented by conidia.
An analysis of the table of distribution of the

genus Coenogonium (Table 1) Show no evidence of endemism.

 

Although the tables given for each country may be
incomplete, it presents the relative extent of field
work done in these areas. More material is needed from

Asia, Africa, Australia and New Zealand.

 

32

Table 1. Distribution of Coenogonium species in the different

geographical areas.

circles refer to literature records of original

X signs refer to material seen,

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

collection.
COENOGONIUM NORTH AMERICA CENTRAL AMERICA SOUTH AMERICA
Species fl
1 N
‘ o m
7‘ 'U C
I a m
m m .a
m . o m c 5
a c *4 H m m LI 0 m
m m m m m m m m -H H o m
E 'U -H 61 E u o m rd .4 .Q o m .c 5
rd r4 m -H m m m :3 t) E > -H E 13 O m
A H -H .c ,m u u to -a m -H N o m c c m
m o s o :x m m a x c .4 m .4 s m o H
Pi H o -H 1a) o s o o o o H o o s H m
«n a. .4 2+: 6 o :1: z cum on o m n a. a.
acrocephalum : E j i0
I i .
botryosum : X: 3 i
complexum X E X; X~ 0i 3 X
: ,‘ . j .'
curvulum g j g i
disjunctum X X 3X XI X. E
implexum X Xi X ; ~ ' X x;
f j q I .
interplexum X; X. X X X: X 0? X
': g 1 -.
interpositum ‘ XS 0 ; X F X;
, 3 _ I i '
Leprieurii ! X Xix X X g X x; 0 !
Linkii x; X x 5x x i x xix x?
i T l
1 I ‘ ‘n
moniliforme X} i X , - 3X5
ornatum : f i j X 5x:
I. , 1 :'
pannosum ’ ' g E O ' « ?
~ 1 E
tomentosum '! ' i
1 I ', t] _,
velutinum j 1 - 1 4;#i - X 1 3
Total No. ‘1 5: 3. 1 1; 5 3 2; 8 4 3 5 1 1 4’
J - ‘ i j; . f 4 t v__
of Spec1es ll 22 21

examined

 

 

 

33

 

moaum< 6mm:

 

pamamwN 3oz
w MHHmHuwsfi

 

mmcflsmflanem

 

m>mb

 

COHhmo

l

 

mango

 

owsuom

 

1

 

sausages

 

OUHm ouuosm

 

moamfimb

Huflmm

X

X

 

omsoamomsw

1; 5' 8 9; 2 1

 

momsouw

l

 

UflHQDQom CMUHQHEOQ

 

mflso

 

mmEmnmm

 

maosnoso>

3i

4g4 4

 

Emcflusm

 

 

snow

 

X X X X X X X

 

2

 

38

THE ALGAE

Trentepohlia occurs in all the species of Coenogonium

 

 

so far reported except in g, moniliforme where the algal

 

symbiont is Physolinum monil§g(de Wildem.) Printz.

 

The Lichenized Algae

The following descriptions refer to the lichenized

algae found in the lichen thallus.

Trentepohlia: The filaments are generally 14-36 H

 

in diameter with the main axis usually larger than the
branches. They are septate and sometimes constricted at the
septa. The cells are uniformly cylindrical or barrel-shaped

and 1-3 times as long as wide. In some species of Coenogonium,

 

the algal filaments are fasciculate—conglutinate in arrange-
ment and the threads are held together by very short funiculi.
Branching is less common with lichenized filaments as com-
pared to free—living plants whose branches may be dichotomous,
one—sided or more commonly, at right angles to the main
filaments. The apical cells of Trentopohlia appear to be

somewhat longer than the other vegetative cells of the

34

35

filaments and are free from the fungal hyphae.

Physolinum: The filaments are branched and consist
of torulose or chain-like cells which are globose or elliptic.
The globose cells measure about 18 u in diameter while the

elliptic cells are 10—15 x 22-32 H in size.

The Algae in Culture

Since it is not possible to identify the Species
of Trentopohlia from the lichenized filaments, it was
necessary to study the algae in culture. The culture studies
were also designed to:
(a) provide a solution to the problem of whether
each fungal species is symbiotic with only one algal species;
(b) understand the range of morphological variation; and,
(c) demonstrate indirectly the influence, if any, of the
fungus component on the alga in the symbiotic relationship.
Although it was not possible to obtain in culture

the algal symbionts of all the reported species of Coenogonium,

 

the knowledge and experience gained from the limited culture
study were useful in evaluating the importance of the algae

for the identification of the species of Coenogonium,

 

particularly in the case of sterile material that was named

by earlier authors solely on the basis of the vegetative

36

morphology of the algal symbiont and the lichen thallus.

It is obvious thatplants grow differently in culture
media, but it is necessary that one should determine the
characteristics which are least variable in relation to
the environmental conditions and how these characters behave

under laboratory conditions. Trentepohlia has been reported

 

by Collins (1928) to be a very variable genus. Hansgirg
(1888) described the extreme variability of this alga in

culture and claimed that polymorphism is commOn in Trentepohlia.

 

It is possible that the external environment is partly
responsible for the variations in the morphology of the alga
in nature. Moreover, one cannot overlook the fact that an
alga may exhibit more variable characters once it becomes
overgrown with the fungus hyphae.

Bornet (1873) gave an excellent description of

the algal symbiont of Q. confervoides and illustrated its

 

relation to the fungus constituent in detail.

Glfick (1896) in his report on Q. germanicum, stated

 

that the algal symbiont of this lichen is the green alga,
Trentepohlia germanicum. Skuja (1933), however, believed
that Glfick's description of the alga tallied very closely
with that of Trentepohlia aurea.

Another attempt at isolating and growing the algal

37

symbiont in Coenogonium has been that of Skuja on Q. niqrum

 

(1933). This plant, which was found growing on bark of
trees in Lettland, appeared as a black mass of felted
filaments. Skuja cultured the filaments of the thallus in
four types of media; namely, distilled water, tap water,
Benecke nutrient solution and Benecke agar media. Parts of

the bark of Picea, Betula and Corylus were introduced into

 

the nutrient solution and Benecke agar medium before these
were sterilized. Skuja observed the development of the

alga and identified it as Trentepohlia umbrina. He found

in his culture studies of the algal symbiont that differences
exist in the vegetative growth of the algae in the four

types of culture media. He noted that branches developed
rapidly in the filaments of the plants grown in distilled
water. Those cultured in tap water measured from 4 to 6 or

8 microns in width and from 14 to 37 microns in length.

The end cells were 50 microns long. In the nutrient solution,
the cells were 10 microns wide and were relatively shorter
than the cells of the filaments grown in the previous two
cultures. The filaments were greener, and the cells were
longer in the agar culture, while the articulate cells (?)
were thinner. The vegetative cells tended to become barrel—

shaped in appearance in agar medium. The haematochrome

38

pigmentation was more distinct in the middle portions of
the cells, while the light-green chromatophores concentrated
at the peripheral part of the cells. The cells grown in
agar medium were wider (up to 12 microns), rounder and had
thicker cell walls which were layered. Skuja observed akinetes
in the filaments, which led him to believe that the algal
synbiont he had cultured was Trentepohlia umbrina.

Karling (1934) made a similar study on the morphology
of g. Linkii collected from Honduras. He cultured the
filaments in Detmer's agar and observed the vegetative growth

and reproduction of the Trentepohlia. Karling presented very

 

good descriptions and illustrations of the structure and
development of the globular zoosporangium (the only type he
saw in his cultures), and the behavior of the zooSpores. The
structure and morphology of the alga under the cultural con-
ditions were, however, abnormal and Karling was unable to

identify the particular Species.

Materials and Methods

Specimens of Coenogonium which have been recently

 

collected from Panama by Dr. G. W. Prescott, Department of
Botany, Michigan State University, and from Florida by the

writer and by Dr. D. B. Ward, Department of Botany, University

39

of Florida, were used for culture studies in the present
investigation.

Pringsheim's soil—water biphasic system (1950) and
Bristol's inorganic medium (modified) (Starr, 1960) were
used for the purpose. Peat and loam soil were employed
separately to determine which soil type would best support
the normal growth of the alga.

For the soil-water experiments, three sets of culture
flasks were prepared: for the first set 72 grams of sandy
loam soil were placed in the bottom of each of 250 m1.
Erlenmeyer flasks, and then 100 ml. of Pyrex distilled water
was added. The pH of the mixture was adjusted to 6.2, using
NaHCO buffer solution. In another set, a small amount of

3

CaCO (about one-sixteenth of a teaspoon) was placed in the

3
bottom of the flasks before the soil and water were added.
The pH was set at 7.5. In the third set of culture flasks,
72 grams of peat-soil were used, adding 100 ml. of the
distilled water afterwards, setting the pH at 5.0. The
flasks were then plugged with cotton and steamed for one
hour on two consecutive days. The purpose of steaming the
culture flasks was to kill the micro-organisms in the soil,

such as protozoa, fungi and bacteria which might interfere

with the growth of the algae.

40

Another type of culture medium, Bristol's inorganic

medium with the following modified formula, was also tried.

NaNO3 . . . . . . . . . . . . .‘. .. 10.0 g.
Cac12 . . . . . . . . . . . . . . . 1.0 g.
MgSO4'7 H20 . . . . . . . . . . . . 3.0 g.
K2HP04 . . . . . . . . . . . . . . 3.0 g.
KH2P04 . . . . . . . . . . . . . . 7.0 9.
NaCl . . . . . . . . . . . . . . . 1.0 g.
Distilled water . . . . . . . . . . 940 m1.
Soil—Water (steamed) . . . . . . . 10 m1.
pH . . . . . . . . . . . . . . . . 7.5

To this solution was added 12 grams of agar and
subsequently autoclaved for twenty minutes at 15 pounds
pressure. The medium was then poured on sterile petri
dishes and allowed to solidify.

Individual filaments from the thallus were picked
by means of fine forceps and were washed carefully in
distilled water, using capillary pipettes for the purpose.
Equal amounts of the washed fragments were then introduced
into the flasks containing the three types of culture
solutions, .while the same amounts were plated by means of
a sterile inoculating needle on Bristol’s inorganic medium.

The flasks and plates containing the isolates were then

41

placed under continuous fluorescent light (approximately 350
foot candles). The cultures were kept in a lighted refriger-
ator maintained at a constant temperature of 22°C. The
flasks were placed at a distance of about five inches below

the source of light.

We

Two days after the start of the experiments, the
cultures were examined for vegetative growth and reproductive
structures, if any. Cultures using peat-soil—water medium
were filled with profuse growths of fungi, with very little
growth of the algae. Therefore, this set of the cultures
was subsequentlydfiscarded. The set of flasks containing a
small amount of CaCO3 exhibited a good growth of the algae
whilerx>appreciable change was noted in the third set of
flasks which contained plain loam soil. It seemed evident,
from these observations, that a slightly alkaline medium
supported a good algal growth and, hence, this type of
culture medium was utilized for the rest of the study.

The algae were then washed and transferred into
new culture flasks containing the small amount of CaCO3
and sandy loam soil and were kept at the same temperature

and exposed to constant illumination. The pH was maintained

at 7.5-8.

42

The algae were examined periodically to detect the
presence of reproductive structures which were needed for
the identification of the algae. It was noted that the
basic medium inhibited the growth of the fungus While the
vegetative growth of the algae progressed. Four days after
the transfer of the algae to the fresh culture solutions,
the fungus constituent appeared to loosen from the algal
threads and disintegrate, leaving the alga to grow
independently. The cell walls of the alga thus appeared
clear and almost devoid of the surrounding hyphae when seen
under the high—power objective of the compound microscope.
At the same time, branching tended to become more free.
This greater tendency to branch in all directions appears
to be associated to a certain extent with the absence of the
fungus hyphae around the cells. In the lichenized alga
in nature, it has been Shown that the fungus develops
strongly around new branches that are just beginning to
form. Karling (1934) believed that the fungus mycelia
inhibit the development of branches to a certain degree
and thus produce the characteristic appearance of the
algal filament in the lichen thallus.

After a period of one week, round or ellipsoid

reproductive structures were found attached either laterally

43

or terminally, to the cells of the filaments. These were
either sessile on the vegetative cells or attached to special
hooked stalk cells, singly or in a whorl. 'These structures
were later determined to be either sporangia or gametangia.
The algal filaments continued to produce new branches and the
apical cells were much elongated. The cells assumed a brighter
green color and the yellow and golden—red globules of haemato—
chrome decreased perceptibly.

Growth on the agar medium was not evident until
seven days after inoculation, at which time the algal filaments
started to Show vegetative growth by spreading out on the
agar plate. The difference in the rate of growth of the
algae in the two culture media may be partly due to the
thickness of the agar which presumably hampered the rapid
growth of the algal filaments. It was further observed
that the characteristic greenish-brown color of the
Trentepohlia filaments remained longer in the plants grown
on agar medium than on those in the soil—water medium.

By subjecting the cultures to alternate 16—hour
light and 8-hour dark periods, the liberation of zoo-
spores could be induced. Biflagellated zoospores and iso-
gametes (in various stages of fusion) were observed in most

of the cultures, both in the soil-water medium and in the

44

solid inorganic medium (Pl. 5, Figs. 3 and 4), although
vegetative growth in the latter medium seemed to be slower.
Reproductive structures and motile cells were observed almost
simultaneously in both types of media four to seven days
after the transfer of the algae from one culture medium to

a fresh one. The cells producing the motile zoospores or
gametes, as the case may be, were seen to be much larger

and longer than the other cells in the filament. Drawings
were made by means of a Zeiss Drawing Apparatus to Show the

reproductive structures and cells of each of the Coenogonium

 

species cultured. Measurements were taken of the cells

of the filaments bearing reproductive structures and these
were compared with those of the filaments supporting the
apothecia in the lichen thallus. The ratios of the width
to the length of these cells were also determined in order
to Show the variations in cell sizes between the algae
grown in culture and the algal symbionts as they grow

in nature. It is presumed that the cells bearing the
reproductive structures in culture would have reached

the "peak" of their normal growth and differentiation

and thus would serve as a good index for comparison with

the cells of the fertile threads of the lichen.

45

Results

In the present investigation, six Species of
Trentepohlia were isolated and successfully grown to
maturity in Pringsheim's Soil—water medium and in Bristol's
solid inorganic medium.

Trentepohlia arborum was identified as the symbiont

 

of three Species of Coenogonium: namely, 9. Linkii, g.

 

interplexum and g. interpositum. Trentepohlia aurea

 

 

was observed to be the algal symbiont of Coenogonium Linkii

 

and Q. interplexum. Two cultures of Coenogoneum Leprieurii

 

presented two different algal symbionts; nemely, Trentepohlia

 

umbrina and I. odorata. Trentepohlia elongata is symbiotic

 

with Q. Linkii and g. interplexum while 3. abietina was

 

identified as the algal symbiont of Coenogonium interplexum.

 

 

Discussion

 

Table 2 lists the species of lichen fungi and their
corresponding algal symbionts. It can be seen from this
table that there is no Specificity of algal host in the
lichen genus Coenogonium. Six different species of
Trentepohlia exist in symbiosis with the four species of
lichen fungi (Coenogonium) studied for their algal components.

Therefore, a fungus can maintain a symbiotic relationship with

46

 

 

 

 

 

 

 

 

 

 

 

N mCHHQED .B
d
N mumuoco .B .M
4 m
x N mummcoao .e a.
T...
O
N N mohsm .B W
S

N N N ESHOQHM .B

N mcsumflnm .8

Hfluswflumoq .0 HHNSHA .U Esuflmomuoucfl .O Esxmamumucfl .O
mucosnoowz

.mGOHUMHUommm uaoflnoomfilucowfloomsm.mo Numfiesm .N manna

47

any one of two or three different algal species. This fact
strengthens the previous assumption that one should not
rely solely on the algal component of the lichen in the

determination of the species of Coenogonium. The variable

 

size ratios obtained in the culture studies justify further
the opinion that algal cell measurements cannot be a primary
characteristic in the identification of the species of

Coenogonium, especially when dealing with sterile material.

 

The figures shown on Tables 3 and 4 to describe the
sizes of the algal cells represent the average measurements
of ten cells taken from each culture flask. The range in
size for length and width is also Shown. AS has been
previously stated, only those cells bearing reproductive
structures as well as those filaments supporting the apothecia
in the lichen thallus were measured for comparison of their
cell sizes.

Table 3 presents a comparison of cell measurement
data from lichenized algae and algae grown in culture.

An analysis of this table Shows the following differences
between species:

Trentepohlia arborum, as the symbiont of g, Linkii,

 

g. interpositum and Q. interplexum showed three patterns

 

of growth:

48

Table 3. Comparison of cell sizes in lichenized and non-
lichenized phycobionts arranged according to algal
Species.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Collection Trentepohlia Coenogonium LICHENIZED
Number Species Species Average Length x

Width (u)

18 T. abietina C. interplexum 50 x 15
17 T. arborum C. Linkii 38 x 17
65 T. arborum C. interplexum 38 x 16
66 T. arborum c. Linkii 38 x 14
67 T. arborum C. interpositum 38 x 16
78 T. arborum C. interpositum 49 x 15
109 T. arborum C. interplexum 41 x 16
64 T. aurea C. Linkii 52 x 15
114 T. aurea C. interplexum 36 x 16
116 T. aurea C. Linkii 35 x 14
117 T. aurea C. Linkii 36 x 17
76 T. elongata C. interplexum ' 35 x 16
77 T. elongata C. Linkii 35 x 15
107 T. elongata C. Linkii 34 x 13
4 T. odorata C. Leprieurii 19 x 5

 

2 T. umbrina C. Leprieurii 32 x 14

\\

\\

 

SOiL-WATER MEDILM

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

ALGA
RATIOS Min. Max. 66666 x W 191-, BAEIQ§F
Ave;_L Average Length X Width (H) Ay§__L
529" W Ave. W
3.3 - 32.6 - 47.6 x 14.4 x 18.6 2.4
39 X 16
2 2 32.6 46.5 X 12 9 - 20.9 ; 4
1 40 X 11 H
34.6 47.4 X 12.7 ~ 18.6
2.4 '.
41 x 16 1‘6
, 24.6 42.6 X 9.3 ~ 17.5 7
i 7 34 X 14 2’4
2 4 . 52.3 65.1 x 13 / 1 ‘7.4 -
” 58 x 15 j°9
3.3 . 40.5 69.2 x 10.9 _ 18.6 3 6
50 X 14 3
2,6 _ 35.2 58.5 x 13.2 — 17.2 2 9
44 X 15 .
3.5 ‘ 31.6 55.8 x 14.2 — 16.1 2 7
41 X 15 .
2_2 . 20.8 48.8 x 9.3 — 20.9 2 3
34 x 15 °
205 . 19.7 46.5 X 6.6 — 16.9 2 6
34 X 13 4
251 _ 30.6 46.5 X 13.3 — 16.7 2 4
38 X 16 .
3.1 _ 27.9 63.3 X 13.1 — 15.4 3 l
47 x 15 ‘
2‘3 . 30.2 46.5 X 10 ~ 18.6 2 5
37 x 15 .
2-6 . 37.2 65.1 X 14.5 — 23.5 2 8
53 x 19 .
4 : 1 14.4 23.4 X 3.1 - 5.6 4 .
20 x 5
2,3 . 23.9 48.8 x 11.3 - 18 2.3

 

 

 

34 X 15111

. 7 “a e:

 

 

 

 

AGAR MEDIUM

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Min Max. Length X W Béglgg
Ave. Length X Ave. Width (M) Ave. L

Ave. W
26.2 ~ 44.: :29.3 — 13.8 3.1 _ 1
22.6 — 32.: $614.5 - 17.6 2.4 : 1
32.5 - $3.: $513.8 — 16. 2.6 . l
30 — 42$: : 12.5 — 17.1 205 = 1
26.6 - $8.: $411.6 - 20. 2.1 : 1
53.8 — ::.i $411.9 - 16. 4.1 _ 1
32.3 - 33.: $614.5 — 16. 2.4 : 1
31.3 — 2%.: i614 — 17.6 2.6 : 1
20.3 - 32.; $19.2 — 13 2.4 _ 1
29 - 3753 : 12.1 — 18.5 294 : 1
29.1 — 32.: :412.9 — 16. 2.4 _ 1
38.6 ~ 25.: :016.9 — 23. 2_4 : 1
25.7 - 33.: :39.3 — 17.1 2.2 : 1
37.2 - :3 : 12.7 — 16.4 2.5 2 1
13.3 — 12.: z 3.8 — 7.7 2.3 : 1

No Growth

50

Table 4. Comparison of cell sizes in lichenized and non-
lichenized phycobionts arranged according to
fungal species.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Collection COENOGONIUM TRENTEPOHLIA
Number species species
18 C. interplexum T. abietina
65 #7 C. interplexum T. arborum
76 C. interplexum T. elongata
109 C. interplexum T. arborum
114 C. interplexum T. aurea
67 C. interpositum T. arborum
78 C. interpositum T. arborum
2 C. Leprieurii T. umbrina
4 C. Leprieurii T. odorata
17 C. Linkii T. arborum
64 C. Linkii T. aurea
66 C. Linkii T. arborum
77 C. Linkii T. elongata
107 C. Linkii T. elongata
116 C. Linkii T. aurea

 

117 C. Linkii T. aurea

 

LICHENIZED ALGA

51

SOIL‘WATER

MEDIUM

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Average Length X Ratios Min.—Max. L. X Min -MaX Ratios
W1dth (h) L/W Ave. Length X Width .0)

33—i—15 3.3 32.6 - :;.: $614.4 — 18. 254 1
EE_X 16 204 34.6 — ::.: $612.7 — 18. 2.6 1’
firx 16 232 27.9 — 23.: :513.1 — 15. 3 1 1
61 X 16 2.6 35.2 — 23.: $513.2 — 17 2‘9 1
:6 x 16 212 20.8 - 42.: :59.3 e 20. 2‘3 1
38 X 16 204 52.3 — 22.: $513.7 - 14., 3.9 1
::~X 15 3.3 40.5 - 23.: $410.9 1 18. 3.6 1
32 x 14 2.3 23.9 1 32.: $511.3 — 18 2.3 1
19 x 5 4 14.4 - 33.: E 3.1 — 5.6 4 E_
38 X 17 2_2 32.6 — 23.: $712.9 — 20.9 204 1
33—? 15 3.5 31.6 — 25.: :514.2 — 16.1 2.7 1
3? X 14 2.7 24.6 ~ 32.: :49.3 — 17. 204 1
35 X 15 203 30.2 — $3.: $510 — 18.6 2.5 1
34 x 13 2.6 37.2 — 23.; :914.5 - 23.5 2.8 1
35 x 14 2.5 19.7 — 42.: :36.6 — 16.9 2.6 1
36 X 17 2.1 30.6 — 42.: :613.3 — 16.7 204 1

 

 

 

 

 

GAR MED I UM

 

Min1mun~MaX1mum Length X W

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

. .. R'tiuc
Average Length X Width (u) a Cg
26.2 — 44.2 X 9.3 — 13.8
. * 1
37 X 12 3 1
32.5 - 44.8 X 13.8 — 16.7 2.6 _ 1
39 X 15
38.6 — 55.8 X 16.9 ~ 23.3 2
- 2.4.31
4/ X 20
32.3 — 44.9 X 14.5 — 16.6 2.4 :1
39 x 16
20.3 — 32.7 X 9.2 ~ 13
26 x 11 2.4-111_-1-_____ __
26.6 — 40.6 X 11.6 — 20.3
30 x 14 2"l ' 1
53.8 — 65.1 x 11.9 ~ 16.7
.. 2 1
58 X 14 4 1
No Growth ~~~~~~~~~~
13.3 — 16.1 X 3.8 _ 7.7
2., : 1
14 X 6 3
22.6 — 42.3 X 14.5 x 17.6 2.4 : 1
38 X 16
31.3 — 52.6 X 14 — 17.6
41X16 2‘6'1
3O - 42.6 X 13.5 — 17.1
38 X 15 2‘5 ' 1
25.7 — 33.9 x 9.3 - 17.1
29 X 13 2'2 ' 1
37.2 — 42 X 14.7 ~ 16.4 .
40 x 16 2°5 ' l
29 - 37.2 X 12.1 — 18.5 204 2 1
33 x 14
2.. - ’ . . — .
9 1 35 5 X 12 9 16 4 2.4 = 1

34 X 14

/

 

.651:

4 . . .
—c 1" “A o“ r
V .o a
.au wam ”h nu.
n... ..... L

‘
L..an

11'. C1:

9“.“
, ,H
.:

“~C

SN.

9.
rs

52

(a) increase in size of cells in both types of culture
media, with the same cell length to width ratios:

(b) increase in size of cells in soil-water medium
and a decrease in size of cells in agar medium; and,

(c) decrease in size of cells in soil—water medium
and no appreciable change in size in agar medium.

Trentepohlia aurea, as the algal symbiont of Q.
Linkii showed variations of growth in several ways:

(a) decrease in size of cells in both soil-water and
agar media, the cells in the former medium being smaller
than those in the latter, although their size ratios are
nearly the same;

(b) lichenized algal sizes are larger than those
in culture but the size ratio of the cells in the soil—
water medium is larger than those in the agar medium
and the lichenized algae: and,

(c) cells in the soil-water medium are larger than the
lichenized cells and those in the agar medium, the cells
in the latter exhibiting higher size ratio than the
lichenized cells, although the cells in the agar medium
are smaller.

Trentepohlia aurea as a symbiont of Q. interplexum

 

showed a decrease in the size of its cells in soil-water

53

medium and a further decrease in cell size and size ratio
in the agar medium.

Trentepohlia elongata, as the symbiont of Q. Linkii,
showed two variations in growth:

(a) increase in cell size and ratio in soil-water
medium and a decrease in size of cells in agar medium:
and,

(b) increased size of cells in soil-water medium
and in agar medium, the cells in the former being much
larger than those in agar medium.

As a symbiont of Q. interplexum, Trentepohlia
elongata cells showed an increase in length and width in
culture, the cells in the agar media displaying a lower
size ratio than in the other two.

The cells of Trentepohlia abietina decreased in
size in culture but have higher size ratio in agar medium.

Trentepohlia odorata gave a larger average length
in soil-water medium but the average width of the cells
in the agar medium were much increased, giving a size ratio
lower than in both the lichenized alga and those in the
soil-water medium.

Coenogonium interpositum is presented here with only

one symbiotic algal species, Trentepohlia arborum. It is

54

possible that more collections of this lichen would have
yielded more significant results.

Collection numbers 18, 67, 77 and 78 (Table 5)
present diverse growth patterns in comparison with one another.
For instance, No. 18 shows a sharp decrease in average length
and width measurements when grown in culture, while No. 67
shows a sharp increase in average cell size in soil-water
medium. There is noted a large decline in average cell
length in No. 77 when grown in agar medium; while a sharp
increase is observed in No. 78 — a distinct contrast to

No. 77. While both Nos. 67 and 78 are Trentepohlia arborum,

 

their growth responses differ with respect to the culture
media they are grown in. Here is an instance where two
lichens with the same algal and fungal components and
collected from two separate localities behave differently in
culture. Obviously, there are some limiting factors that
influence the growth of these four collection numbers in
culture and it is possible that nutritional deficiencies
in the medium play a role.

Collection No. 2 is not represented with any data
on growth in agar medium in the Tables. The alga grew
very poorly in this medium, being so much reduced in form

that the attempt to culture it further in agar medium had

55

 

 

 

 

 

 

 

 

.mEmcmm n

= = = = : “mug“ 0—H. “HHVFHHHHA 0U FHH
= = = = = mmusm .9 flflxcflq .0 baa
moma .wa .nmm .00 moan .thm
Hmcoflumz mmpmamum>m .xooafimm comm mmusm .9 Esxmamumusfl .0 waa
: = = : Edmonum .9 Esxmamumucfl .0 moa
mood .MH .Qmm .00 mpmn .xumm Hmsoflumz
mommamum>m .xoosfimm mxmcmmauumm mummcon .9 flflxcfln..0 boa
= = . . snuonum .e sauflmomumuan .0 ms
. . . . mummcoaw .a anuflg .0 up
moms .HH .nmm .oo mncmanmflm .mcnunmm .xumm
muMpm xUOEEmm nonnasmflm .Hflmn9 “woman mummcoam .9 demamumucn .0 on
= = = : = Esuonum .9 Esuflmomnmusu .0 no
.- : 3 = 3 EDHOQHM 0-H. flflxcfiq 0 U 00
= .00 mHOCHEwm .puowcmm Home xUOEEmm 30A. Edmonum .9 Esxmamumusfl .0 mo
mmma .m .Qmm .00 maocflfimm
buomsmm .xUOEemm Eouuom xomam women .9 Hflxsflq .0 so
= . = . mcfiumnnm .9 esxmamumucfl .0 ma
mmma .m .Qmm .00 mssoma<
maafl>mmcflm0 .xUOEEmm ummmonaaflz Edmonum .9 flflxcflq .0 pa
= = = . = mumuoco .9 anusmflummq .o w
mmoa .wm .own mEmcmm .UsmHmH oomHoHoo ounmm mcflnafis .9 Hausmfinmmq .0 N
o o
coauomaaoo sofluumaaoo no woman madnommucwu9 Esasom smoo 02
mo mumn mo mwaowmm mo mmflummm coauumaaoo
cm mbflnomh EOHM mCOfluomHHoo mo muma .m manm9

56

to loe given up. The writer was unable to determine the

limniting factors in this case. It showed significant

inczxease in growth in the soil-water medium although the

SiZe ratios of the lichenized alga and the filaments grown

ill culture remained the same.

Collection numbers 67, 76, 78 and 107 grow better

iri culture and apparently, no limiting factor is present

9

heaz:e.

The extent of the lichenization of the algae may be

thought of as one of the factors that slows down or hastens
tlie vegetative growth of the thallus. As has been described

earlier, one of the first signs of growth of the algae in

tflae culture media was the shedding off of the hyphal elements

*Enveloping them. There appears to be an inverse relation-

ship between the amount of hyphal threads around the alga

and the rate of growth of the alga.

It might be well to mention at this point that the

 

Egentopohlig species presented here have variable growth
requirements and these do not necessarily correspond to

the different taxonomic forms, nor do these have significant

relation to size measurements.

a.“
.6

n

I «
our

t

 

a:
51‘

 

57

Conclusions

 

1. One species of the algal genus Trentepohlia

 

can be associated with more than one fungus species of

the genus Coenogonium in the development of a lichen thallus.

 

2. One species of the fungus genus Coenogonium can be

 

associated with more than one species of the algal genus
Trentepohlia in the formation of a lichen thallus.

3. The shelf—form of lichen thallus is a character-
istic of the fungal component since both shelf-form (g.

Linkii) and non-shelf—form of thalli (g. interpgsitum)

 

have identical algal species.
4. Cell sizes of the alga cannot be used in the
classification of the lichen nor in the identification

of the species of Coenogonium as evidenced by the polymorphic

 

tendencies shown by both the lichenized and the unlichenized

algae.

Taxonomy

History

The species of Trentepohlia are largely polymorphic
plants and many of them have been described on characters of

little permanence. The genus was founded in 1817 by Martius

58

and was based on Byssus aurea Linn. Agardh (1824), however,

 

included 3. aurea and the unrelated genus Chantransia in his

 

new genus Chroolepus. More species were added to this list

by subsequent workers who classified Chroolepus in various

 

ways in the Class Chlorophyceae (green algae).

 

Kfitzing, in 1843, placed the genus Chroolepus, along

with the genera Cladophora and Stigeoclonium, in the Family

 

 

Confervinae based on the organization and shape of their
cells. Rabenhorst (1864) added nine species and seven varieties
to the list of Agardh, rejecting two of Kfitzing's previously

described species. He classified Chroolepus under the Order

 

Nematophyceae and Family Chroolepidae. Gobi published an

interesting memoir on the genus Chroolepus in 1871 and

 

a year later, Nordstedt described a new species from Brazil
which he named Trentepohlia pleiocarpa. He was the first to

recognize the priority of the name Trentepohlia over Chroolepus

 

 

and this led to the subsequent transfer of the various species

named by the earlier authors into the genus Trentepohlia.

 

Wolle (1887) described six species and two varieties
which he classified in the Family Confervaceae under the
Order Confervoideae. De Wildeman (1888) compiled all the
species of Trentepohlia previously described into a new

listing and included 16 new Species and one variety. He

59

organized these under the Order Confervineae and Family
Confervinees, placing this after the Family Ulotrichiacees.

The remaining studies on Trentepohlia were mostly

 

concerned with the description of new species or the re-
description of old ones. The use of the vegetative habit
of the thallus as the main character in the classification
of the genus seemed to dominate the systems of the later
authors.

Hariot (1889-90), brought together and analyzed

the scattered descriptions of the Chroolepus and Trentepohlia

 

 

species and presented a good listing of these under the

name Trentepohlia. His work comprised enumerations and

 

descriptions of the different species but were not fully
illustrated and he failed to include keys to the species.
Hariot based his classification on the cells of the thallus,
their relative dimensions according to the two axes of the
filaments, the shape of the cells, whether cylindrical or
torulose: the type of extremity of the filaments and of the
branches, and the differentiation of the filaments into the
creeping and the erect threads, together with explanations
on how these arise. He did not consider the structure of
the sporangia as a good taxonomic character in view of

what he thought was its great variability. Hariot further

60

divided the genus Trentepohlia into two groups:

 

(1) species with cylindrical cells in long filaments;
and,

(2) species with torulose or moniliform cells in
a pulverulent growth layer.

Collins (1932) placed the genus Trentepohlia in

 

the Family Trentepohliaceae under the Order Ulotrichales
on account of the uninucleate cells and the presence of a
single parietal laminate chloroplast in the cells. He

listed ten species and five varieties of Trentepohlia from

 

the United States of which one species and two varieties
occur in Florida; namely, 2. odorata (Wiggers) Wittrock,
.I- odorata var. umbrina (Kfitz.) Hariot and 1. odorata var.
ggercina Collins.

Printz (1927, 1939) considered the heterotrichous
organization of the thallus of Trentepohlia as the main
character in the classification of the genus and included
it in the Order Chaetophorales. He placed the genus in
the Family Trentepohliaceae mainly on the fact that the
zoospores and gametes are formed in special cells differing
more or less in shape and structure from the vegetative
cells. He stated that these characters are important dis-

tinguishing features of the genus to justify its separation

61

from the Order Ulotrichales. Printz (1939) has published
7 what is considered to be the first comprehensive monograph
of the Family Trentepohliaceae, complete with keys and
illustrations. He placed the genera Trentepohlia and Physolinum
in the same family and distinguished them by their reproductive
structures and vegetative habit. Printz divided the genus
into two sections and characterized each thus:

(1) Section Chroolepus - plants consisting mostly of
free, more or less cylindric or torulose, irregularly branched,
erect filaments, attached only at the base and not forming a
distinct prostrate tissue: and,

(2) Section Heterothallus - plants consisting of
distinct basal threads forming a pseudoparenchymatous tissue
attached to the substratum on its underside; and, loose,
unbranched or sparingly branched, upright threads.

The genera included in this study are placed in the

Order Chaetophorales of Class Chlorophyceae (green algae)
on the basis of the heterotrichous character of the filaments.
The Family Trentepohliaceae, to which these plants belong,
is characterized by the high degree of adaptability to a
terrestrial existence, their sub-aerial habit,the presence
of the haematochrome pigment in their cells and the absence

of pyrenoids in the chloroplasts.

62

Some species of Trentepghlia and Physolinum occur

 

 

as symbionts in certain genera of lichens, in addition to
Coenogonium. Ahmadjian (1958) lists Trentepohlia as the

algal symbiont of Cystocoleus niger (a9, nigrum), Chaenotheca

 

Aphaeocephala var. hispidula, Qpegrapha atra, Graphis scripta

and Arthonia radiata f. astroidea. He also listed the genus

 

as occurring in the following lichen families, among others:

Cypheliaceae, Arthoniaceae, Chiodectonaceae, Dirinaceae,

Rocellaceae, Lecanoactidaceae, Thelotremaceae and Gyalectaceae.
Physolinum monilggoccurs as the algal symbiont of

.Q. moniforme.

 

General Morphology of the Phycobionts

Trentepohlia and Physolinum are sub-aerial plants

 

with decumbent filaments producing erect filaments which

are simple or branched, parallel or irregularly arranged,

and whose cells may be cylindrical or spherical. In the
majority of the species, the Trentepohlia appear as orange—
yellow tresses or cushions, although in a few species, like

.2. umbrina (Kfitz.) Born., and Physolinum monilia(de Wildem.)
Printz, they may occur merely as fine crusts. In many sepcies
of Trentepohlia, the thallus shows differentiation into

prostrate and erect threads but in g. umbrina the filamentous

tn

63

habit is poorly developed. Species belonging to the

Section Heterothallus have upright filaments with few or

no branches and are gradually attenuated. The basal threads
form a pseudoparenchymatous tissue. In most cases, the
plants of both Sections Chroolepus and Esterothallus have
well-branched filaments, the branches being alternate and
arising either from the top of the parent-cell or sub-termin-
ally or rarely from the middle. The growth of the filament
is apical.

The vegetative cells vary greatly in length and
diameter, and it is very difficult to identify the species
on this character alone. As has been shown in the culture
studies, measurements of cells in different parts of the
lichen thallus and under different cultural conditions
showed a wide range of variation.

The cylindrical or barrel-shaped cells have thick
cellulose walls which are sometimes plainly stratified,
the successive layers being parallel or divergent. In
forms showing divergent strata in their cell walls, the
free ends of the apical cell are covered by charaCteristic
caps of pectose (Pl. 3, Fig. 5), which represents a secretion
and serve to restricttranspiration from the exposed tips.

In 2. aurea, the caps are commonly pushed aside during the

64

grcwth of the apical cells and the remains of such displaced
CE1}95 may be found attached to the lateral walls of the older
3€E1gments: in such cases the apical cell forms new caps
(Fritsch, 1956).
The characteristic yellowish to red color of the
‘laiving plant is due to the presence in the cells of orange-
lted haematochrome globules of various sizes dissolved in
:Eat (Geitler, 1923), especially around the discoid and band-
shaped chloroplasts. In shaded habitats, the pigment may
be masked by the chlorophyll so that the growths appear green.
Haematochrome globules accumulate in large quantities in the
cells of filaments exposed to well-lighted situations and
during dry periods when growth is slowed down, greatly
diminishing in size and numbers when the cells are shaded
or when placed in nutritive mineral solutions.
The septa separating the cells are often provided With
a single large pit which is occupied by a protoplasmic strand.
A distinct characteristic of this genus is the absence of
pyrenoids in the cells. Older cells are commonly multi-
nucleated.
Irentepohlia and Physolingm_are widely distributed
and occur attached to or forming matted layers on rocks,

tree—trunks, wood—work, and on leaves and bark of trees.

65

They range from arctic or high mountain regions to the
damp tropical or sub-tropical climates where they grow
P13<Dfusely as epiphytes on leaves (Schmidle, 1897). In
tléezmperate climates, they are found wherever the necessary
degree of moisture exists. They are thus most common in wet
In£1untainous regions and are usually comparatively rare

iqn the lowlands. It is possible that the original range

<>f these plants was in tropical regions, and that from

there, they either spread to the damp mountainous areas

of the temperate regions or they survived in these regions

after certain climatic changes.

Plants of Trentepohlia and Physolinum can withstand
long periods of drying without appreciable change (Howland,
1929), making them suitably adapted to terrestrial existence.
Oltmanns (1922) has reported that the highly refractive
yellow and golden-red globules in the cells of Trentepohlia
act as light filters which enable the plants to withstand
the intense illumination of the tropics and thus compete
successfully with other algae. A curious adaptation to the
terrestrial habit of Trentepohlia is found in the fact that
the mature sporangia are frequently shed entire from the
filaments and the zoospores are released only when they fall

into the water, or are wet with dew or rain.

66

vegetative reproduction by fragmentation is probably
C’ii frequent occurrence. Howland (1929) reports that propa—
EIEition by swarmers in T. aggea only takes place at certain
times of the year. The zoospores are formed in special
ellipsoid or ovoid sporangia and are of three kinds:

(1) sessile sporangia which may be terminal, lateral,
:intercalary, or rarely axillary (Pl. 3, Fig. 2). They are
formed merely by the enlargement of a vegetative cell. The
zoospores are liberated without the detachment of the
sporangium from the main cell. Karsten (1861) reported that
such sporangia were soon transformed into globular forms
when thalli were grown in water, and that the globular forms
occurred most abundantly in moist places.

(2) stalked sporangia which appear only as terminal or
lateral appendages and are formed by the cutting off of a
tubular outgrowth from a somewhat enlarged supporting cell
which may give rise to several such sporangia (Pl. 3, Fig. l).
The apical part of the outgrowth swells up to form the sporan-
gium which is often bent in a characteristic sickle-shaped
manner. The stalked sporangia become detached and dispersed
by wind, the liberation of the biciliate zoospores taking
place on subsequent wetting.

(3) funnel-sporangia which are produced terminally from

67

0163 apex of a cylindrical cell and become detached at

mali2nrity. Oltmanns (1922) believes that this type of
S'EZ'CDrangia is but a modification of the stalked sporangia.
It: is not, presently known what kinds of reproductive cells
are formed by the funnel-sporangia.

The sessile sporangia produce biflagellated zoospores
Vflnich have been observed in certain cases to behave as iso-
‘gametes (Wille, 1889). However, they can also germinate
‘Without fusion or may give rise to aplanospores (Meyer, 1911).
This type of sporangium is often regarded as a gametangium
(Pl. 5, Fig. 1). These structures are commonly produced on
the basal parts of the plant, in which case, sexual fusion
readily takes place.

The stalked sporangia produce a considerable number
of bi- or quadriflagellated zoospores liberated through a
terminal or lateral opening. The zoospores appear to germinate
directly (Karsten, 1891). The usual production of the stalked
sporangia on the erect threads facilitates wind dispersal.

The zoospores vary in size and shape but are generally
flattened and ovoid. The characteristic light green color
is more or less diffused in the cells. The two flagella
are borne posteriorly and appear to be of equal length and

approximately twice the length of the zoospore. They range

68

fr(Dru 6 x 12 u in size to slightly larger after their release
f37<>nlthe sporangia. Gametes are smaller, measuring about

4 )< 8 u in size (P1. 5, Fig. 2). Sexual fusion has been
Olbaserved in culture, the process taking place especially
after the cultures have been placed briefly in the dark
{1?i9s. 3 and 4).

In many cases, stalked and sessile sporangia occur
(Dn the same plant, the sessile sporangia being produced from
the cells of the prostrate filaments while the stalked
sporangia are developed on the upright threads. This has
been reported in T. umbrina (Fritsch, 1935), a fact which the
writer considers as questionable since Trentepohlia umbrina
has a thallus where there is no distinction between the
prostrate and erect systems.

Physolinum monilfi;(de Wildem.) Printz grows and
develops branches by a process similar to budding and the
sporangia produce only aplanospores (Pl. 4, Fig. 5). The
vegetative cells are chain—like and moniliform and these bear
the sporangia directly. The chromatophore in the vegetative
cell is long, narrow and ribbon-shaped, one or several in
each cell. Intercalary cell division is not known to occur
in this plant and hence, growth is apical. This species is

the algal symbiont of Coenggonium moniliforme Tuck.

69

Key to Species

 

.7

The sh: species of Trentepohlia found symbiotic with

C3<Denogonium in this study belong to the Section Chroolepus.

 

JEIi the preparation of the key below, the system of classifi—

c:ation by Printz (1939) was adopted because of his use of more
J:eliable characters, such as the type of sporangia and the
shape of the cells. This key is for the identification of

the species of Trentepohlia and Physolinum which are known

 

 

to be found as phycobionts in Coenogonium based on culture

-studies.

1. Cells globose to elliptic, 10-18 x 22—32 H: filaments
chain-like, branched; reproduce by means of aplano-

spores; algal symbiont of Q, moniliforme

 

Physolinum monile‘

 

1. Cells cylindrical or barrel-shaped, 14-36 u in diameter,
1-3 times as long as wide; filaments variously branched:
zoosporangia and gametangia produced, sessile or on

stalk cells: algal symbiont of g, interplexum, Q.

 

interpositum, Q. Linkii and Q. Leprieurii . . . . . . . 2
2. Cells cylindric . . . . . . . . . . . . . . . . . . 3
2. Cells elliptic . . . . . . . . . . . . . . . . . . 6

3. Cells mostly between 5-10 u wide, occasionally slightly

70

thicker or thinner; sporangia ovoid, lateral or
terminal . . . . . . . . . . . . . Trentepohlia abietina
Cells more than 10 u wide . .0. . . . .-. . . . . . . . 4
4. Cells ll—l7 u wide, 1-3 times longer than

wide; cell walls stratified, smooth or rugose:

sporangia sickle-shaped . . . . . . . . . . . . . . 5
4. Cells up to 20 u wide, 3-4 times longer than

wide: branches often one—sided: sporangia

single, ovoid or elliptic attached directly

on vegetative cells . . . . . . . . . . ..T. elongata
Cells 14-17 H wide; sporangia in groups of 2-8,
attached to swollen end cells of a branch . . . . .

‘3. arborum

Cells thinner, ll-l7 u wide; sporangia single,
attached on one—celled sub-sporangial cells

arising eccentrically from a not swollen base

332:2;
6. Cells;t swollen, elliptic, often single or

in short, easily breaking threads; cells

14-15 u wide . . . . . . . . . . . . . . .2. umbrina

6. Cells ovoid-elliptic or nearly cylindrical,
united into long threads, loose and richly
branched; cell wall slightly thickened; cells

5-6 u wide . . . . . . . . . . . . . . . .2. odorata

71

Systematic List

 

Trentepohlia abietina (Flot.) Hansg.

 

:F’Jrodomus Alg. 1886. 86. Chroolepus abietinum Flotow

 

3.11 Kfitz. Phyc. germ. 228. 1845.

Synonyms: Trentepohlia velutina (Kfitz.) Hansgirg.

 

IProdr. der Alg. von Bohmen. I. 2. 1888.

'2. abietina var. cupressicola Hansgirg. Ibid.

 

Exsicatti: Wittrock and Nordstedt, Alg. 917 and

1418. 1882.

Thallus reddish or nearly golden-yellow, thin, caes—
pitose, becoming ashy when dry: cells of prostrate threads
more or less torulose; cells of erect filaments cylindrical,
12-16 H wide, l—3 times longer than wide; terminal cells
sometimes considerably longer; cell walls consisting of
divergent layers, difficult to recognize in living specimens;
cell wall at end of tip cell fairly wide; calyptra poorly
developed; gametangia terminal or lateral; zoosporangia
globose or ovoid, 12-25 u in diameter (Pl. 4, Fig. 2).

This species resembles very closely the fragile
forms 0f.E-.§E£§§ except for the more delicate and flexuous
filaments of the former; the gametangia are smaller and are

found anywhere on either the basal or erect filaments.

72

Trentepohlia abietina is generally found in places
V9114th high humidity, particularly in shady forests, on the
EDEark of coniferous trees and occasionally on deciduous trees

(JPrintz, 1939). Symbiotic with g. interplexum. It is here

 

Iteeported for the first time from Florida.
Phycobiont studied: FLORIDA: On bark of gingg.
glabra, Solution sink, Millhopper's Hammock, 8 miles
east of Gainesville, Alachua County, Uyenco 18, 6 February

1963 (MSC).

Trentepohlia arborum (A9.) Hariot

Jour. Bot. France. 20:345. Figs. 8 & 9. 1890.

Conferva arborum Ag. Systema Algarum 21. 1824. —

 

Orig. coll.: Marianas Isl., Gaudichaud (no date).

 

Synonyms: Tretepohlia pleiocarpa Nordst. Bot.
Not. 131. 1882.

Trentepohlia bisporangiata_Karsten. Ann. Jard.
Bot. Buitenzorh. 10:1-66. 1891.

Trentepohlia Wainioi Hariot, Journ. Bot. France.

 

3:381. 1889.

Trentepohlia Kurzii (Zeller) de Toni, Sylloge
Algarum. I. 261. 1889.

Trentepohlia polycarpa de Wildem. p.p. Bull. Soc.

Bot. Belg. 26:71-80. 1888.

73

Chroolepus flavum var. tahitense Grun. Abhandl.

zool.-bot. Vereins. 1861.

Exsicatti: Wittr. and Nordst., Alg- Exsicc. 409

and 516. 1882.

Filaments long and branching at right angles,
branches somewhat thinner than the main filament: end
cells of the sterile branches often attenuated and more
or less cuspidate: cells 14-17 u wide, 1—3 times longer than
wide; wall colorless or slightly yellowish, more or less firm,
convergent stratified, smooth or rugose; gametangia sessile,
usually terminal, sometimes 3 or more attached laterally by
a short branch; zoosporangia ovoid or globose, rarely single,
2-7 arranged in a characteristic manner around the large
swollen.apical cells by means of their bent stalk cells;
zoosporangia ll-24 11 wide, opening through a pore which
lies opposite their point of attachment to the stalk cells:

gametangia lateral, sessile, globose to elliptic, 19 x 22 u.
in size. (Pl. 3, Fig. l).
The vegetative filaments of this species resemble

veryr<closely T. aurea and is difficult to distinguish from

theellatter, except for the characteristic arrangement of the

zoosporangia in _T_. arborum.

74

This species is common in the tropics growing on
rocks and tree trunks (Printz, 1939). Symbiotic with g.

Linkii, Q. interplexum, and g, interpositum. It is reported

 

here for the first time from Florida.
Phycobionts studied:p FLORIDA: On bark of garya

tomentosa, in limestone sink, Millhopper's Hammock, 8 miles

 

northwest of Gainesville, Alachua Co., Uyenco l7, 6 February
1963 (MSC); on bark of citrus, Low Hammock near Sanford,
Seminole Co., Uyenco 65, 9 February 1963 (MSC): on bark of
‘Qgercus virginiana, growing among liverworts, Low Hammock

near Sanford, Seminole Co., Uyenco 66, 9 February 1963 (MSC):
on bark of sweet gum, Low Hammock near Sanford, Seminole

Co., Uyenco 61, 9 February 1963 (MSC); on bark of sweet gum,
Lieder Trail, Highlands Hammock State Park, Sebring, Highlands
Co., Uyenco 18, 11 February 1963 (MSC); on bark of Lyssoloma,
Rattlesnake Hammock, Everglades National Park, Dade Co.,

_gyenco 109, 13 February 1963 (MSC).

Trentepohlia aurea (Linn.) Mart.

Flora Crypt. Erlang. 351. 1817. Byssus aurea Linn.
Spec. P1. 2:1347. 1759. Lichen aureus (Linn.) Ach.
Lich. Suec. Prodr. 11. 1798. Conferva aurea (Linn.)

Dillwu Brit. Conf. London. pl. 35. 1809. Ectocarpus

75

aureus (Linn.) Lyngb. Hydrophyt. Dan. 164. pl. 44.
1819. Chroolepus aureum (Linn.) Wolle. Freshw. Alg.

U.S. 121. pl. 115. f. 1-21. 1887.

Synonyms: Amphiconium petraeum Nees. Syst. Pilze
69. 1817. in adnot.

Trentepohlia velutina Kutz. Phyc. gen. 1843.

Trentepohlia Tuckermanniana Mont. Sylloge gener.
spec. pl. crypt. 1856.

Trentepohlia villosa de Toni. Sylloge Algarum 235.
1889.

Trentepohlia polymorpha Deck. Script. Bot. Hort.
Petropolit. IV. 1893.

Trentepohlia germanica Gluck. Flora 82:268. 1896.

Trentepohlia Gobii Meyer Bull. Soc. Nat. Moscou,
S. Biol. 45. 1963.

Exsicatti: Wittr. and Nordst., Alg. Exsic. 40, 918,
1228, 1419. 1882.

Thallus golden—yellow to orange-red, yellowish to green-
ish‘gray when dry forming.thick, extensive felt-like growths on
the shady.sides of tree trunks: cells of basal filaments

cylindrical or somewhat swollen, erect filaments cylindrical,

. 76

parallel or irregular, richly branched; branches obtuse, some-
times tapering; cells 11-17 H wide, 1.5 — 3 times longer than
wide; cell walls smooth or somewhat scaly or divergent strati—
fied; pectic calyptra of the tip cell not developed in a
capitate form, appearing like a cylindrical stump, often
pushed to the side during the growth of the cell; gametangia
round or elliptic, terminal or lateral, borne directly on

the vegetative cells of either the prostrate or the erect
threads; zoosporangia oval, 11-17.5 u in diameter, mostly
single, rarely in two's, each attached to a subsporangial cell
by means of special curved stalk cells; base not swollen

(Pl. 3 Figs. 3 and 5).

This is an extremely variable species and distinguished
by the cylindrical, obtuse filaments of moderate diameter

and by the solitary or seriate gametangia.

Trentepohlia aurea is widespread in cold, temperate
climates and in tropical regions, growing mostly on old wood,
stems, rocks and limestone, often associated with mosses and
hepatics (Printz, 1939). Symbiotic with Q. Linkii and g.
interplexum. This species is reported here for the first

time from Florida.

77

Phycobionts studied: FLORIDA: On bark of sweet gum,

 

Black Bottom Hammock, 8 miles west of Sanford, Seminole

County, Uyenco 64, 9 February 1963 (MSC); on bark of Lyssoloma,
Redd Hammock, Everglades National Park, Dade Co., Uyenco 114,
14 February 1963 (MSC); on bark of Figgg, Redd Hammock,
Everglades National Park, Dade Co., Uyenco 116, 14 February
1963 (MSC): on bark of Dipholis salicifolia, Fire Road near
Gate 3, Redd Hammock, Long Pine Key, Everglades National Park,

Dade Co., Uyenco 117, 14 February 1963 (MSC).

 

Trentepohlia elongata (Zeller) De Toni

Sylloge Algarum I. 5:243. 1889. Chroolepus elongatum
Zeller. Hedwigia 12:190. 1873.

Exsicatti: Wittr. and Nordstedt, Alg. Exs. 1421.

1882.

Filaments extended, attenuated,, branched, obtuse;
branches often one—sided, short or long, frequently
spreading, distant or close to one another; threads
nearly tufted-verticillate, branches of approximately
the same width as the main filaments: cells 13-20 u wide,
29-53 u long; sporangia 12-36 H in diameter, sessile or
stalked, single (Pl. 3, Fig 2; Pl. 4, Figs. 1, 3 and 4).

Common on tree trunks and native to East Asia (Printz,

78

1939). Symbiotic with Q. interplexum and g. Linkii. This

species is being reported for the first time from Florida.

Phycobionts studied: FLORIDA: On bark of oak tree,
Lieder Trail, Highlands Hammock State Park, Sebring, Highlands
Co., Uyenco 16, 11 February 1963 (MSC); on twigs of sweet
gum, Lieder Trail, Highlands Hammock State Park, Sebring,
Highlands Co., Uyenco 21, 11 February 1963 (MSC): on twigs
and bark of Lyssoloma, Rattlesnake Hammock, Everglades

National Park, Dade Co., Uyenco 107, 13 February 1963 (MSC).

Trentepohlia odorata (Wiggers) Wittrock

Scand. Vaext. 4:16. 1880. Chroolepus odoratum Wiggers
Primitiae Florae Holsatiae. 122. 1780.

Synonyms: Chroolepus irregulare Kfitz. Spec. Alg.
427. 1849.

Chroolepus betulinum Rabenh. Krypt. Flor. v Sachs.
225. 1863.

Chroolepus quercinum Rabenh. Fl. Eur. Algar. III.
372. 1868.

Trentepohlia lichenicola De Toni, Sylloge Algarum.
I. 235-249. 1889.

Trentepohlia sinense De Toni Ibid.

79

Thallus felt—like, reddish—yellow to orangeered,
yellowish to gray—green when dry, forming a more or less
dense, pulverulent or tomentose growth on the substratum;
threads not distinguished into prostrate or erect filaments:
branched: cells more or less rounded, ovoid—elliptic to nearly
cylindrical, 10 u in diameter, l-l.5 times longer than wide:
cell wall either thin or thick and stratified, layers oblique
or concentric: calyptra narrow; gametangia nearly spherical
to elliptic, lateral, terminal or intercalary: zoosporangia
similar in shape, 13 u in diameter (Pl. 5, Fig. 5).

This species occurs mostly on the bark of deciduous
trees, vary rarely on coniferous trees. Cosmopolitan but never

abundant (Printz, 1939). Symbiotic with g. Leprieurii.

 

Phycobiont studied: PANAMA: On bark of palm tree,
_Barro Colorado Island, Canal Zone, Prescott 4, 26 December

1962 (MSC).

Trentepohlia umbrina (Kfitz.) Born.

in Wille, Algol. Mittheil 26. 1878. Chroolepus umbrinum Kfitz.
Phyc. gener. 283. 1843. Protococcus umbrinus Kfitz. Phyc.
gener. 169. pl. 7, f. 2. 1843. Trentepohlia odorata var.
umbrina (Kfitz.) Hariot Jour. Bot. France. 36:317. 1890.
Synonyms: Protococcus salicis Ag. Systema Algarum

88. 1824.

“li— Fumu tab... . .nD'OU‘JIRHl'E.IIII_1II|:I-lilvlu
A.

_.._

80

Chroolepus irregulare Kfitz. Species Algarum. 427.
1849.

Protococcus crustaceus Kfitz. Sp. Alg. 203, n. 19. 1849.

Lepraria Jolithus Engl. Bot. pl. 2471. (sec. Kfitz.,
1849).

Pleurococcus seriatus Wood Fresh. Alg. U.S. 78.
pl. 10, f. 2. (sec. Wolle, 123. 1887).

Trentepohlia Bleischii var. piceae Wille Bot. Not.
6. 1878.

Trentepohlia polymorpha Deck. Script. Bot. Hort.
Petropolit. IV. 1893.

Exsicatti: Wittr. and Nordst. Alg. 42, 221, 915,

916 and 1423. 1882.

Thallus thin, appearing as a fine crust on the sub—
:stratum; reddish—brown in color, fading when dried; filaments
generally poorly developed and very indistinct, no distinction
between prostrate and erect portions: cells of filaments
elliptic, irregularly connected, easily separating from one
another into groups of cells or breaking at the thick septa
into single cells: 14-15 u wide, 2—2.5 times longer than
wide; wall thick, concentric stratified; gametangia similar

to the vegetative cells, but with a short neck; zoosporangia

81

globose, single, 15 u in diameter, zoospores quadriflagellated.
(Pl. 3, Fig. 6).

This species is very variable especially as to the
size of the cells. Printz reported the range as 7-35 u
thick and 1-2 times longer. The specimen studied here
has the characteristic elliptic cells which break easily
into small fragments.

very common in temperate regions, occurring on the
bark of deciduous trees, rarely on conifers and on rocks
(Printz, 1939). Symbiotic with g. Leprieurii.

Phycdbiont studied: PANAMA: 0n bark of palm and
Eiggg, Barro Colorado Island, Canal Zone, Prescott‘g, 26

December 1962 (MSC).

THE FUNGUS

Nbrphology

Thallus

The fungal elements which surround the algal filaments
consist of a network of thin, delicate, hyaline or colorless
threads which are septate. These branching hyphae range in
size from 1.5 to 2.5 u in diameter, although in some Species

of Coenogonium, particularly in g. Linkii, the hyphae may

 

reach up to 3.5 u in width. These threads grow around the

 

 

Trentepohlia and Physolinum filaments in a longitudinal
direction, anastomosing and Spreading moderately or irregular-
ly. The hyphae may partially cover the algal filaments or
they may completely envelop the algae, especially when the
former are pressed together. In this case, they appear like
a dense, compact, cylindrical coat around the algae.

Fungal filaments appear to be denser and more
articulated near the base of the thallus, and do not become
intracellular at all.‘ Obviously, the fungus hyphae do not
have any injurious effect on the algal cells. The apical
cells of the algal symbionts are usually free of hyphal

82

83

elements, especially during a period of rapid growth when the
fungal hyphae appear to lag behind. However, if the growth

of the alga is very slow, the hyphae may extend up to and
around the tip of the algal cells and the young branches. The
number of hyphae around each algal filament varies. They are
particularly abundant around the branches and may often be

several cells in thickness.

Apothecia

The apothecia are found attached to the filaments of
the thallus, scattered here and there, singly or in groups.
In the shelf—like forms of the genus, they can be found on
either the upper or the lower surfaces of the thallus.

The apothecia vary in size from 0.2 mm- (C. curvulum)
to 1.5 mm. in diameter as in g. implexum.

The external features of the apothecium are quite
variable. The fructification is generally round in shape
and may be biatorine or lecidioid in structure. It may be
sessile or provided with a short stalk and has been described
variously as substipitate, sub-pedicillate or subsessile”

(Pl. 1, Fig. 2). In many species of Coenogonium, the young

 

apothecium is at first stalked appearing like a top or goblet-

shaped, gradually becoming subsessile or sessile, as the

84
case may be, while the disk widens at maturity.

The disk of the apothecium may be flat to slightly
concave or concave, and is oftentimes convex (Pl. 1, Figs.
1-6). Generally, the disk of the young apothecium starts
as concave in shape, later becoming convex as development

proceeds. In 9. velutinum, the disk may appear pruinose,

 

a characteristic that distinguishes this species from
the other members of the genus.

The color of the apothecium has been variously
described as carneous-yellow or carneous-orange to yellowish-
orange or saffron yellow. It has been found that the color
of the apothecium may change upon drying and the original
color is not fully retained. However, the orange-carneous
color seems to be the general appearance of the mature
fructification in the living state. The apothecium is always,
however, whitish in color during the early stages of its.
development, turning into the specific color upon reaching
maturity. In 9, Linkii, the apothecia of the living lichen
is pale-yellow and changes to red when dried. The apothecia
of g, pannosum is whitish-orange when young and become
flesh-colored when fully developed (Mfiller, 1881). Dodge

(1933) separated g, Leprieurii from the other species of

 

Coenogonium on the color of the disk. The use of this

85

differentiating character, however, was refuted by Santesson
(1937), who stated that the apothecia of probably all species

of Coenogonium get the same color and appearance if they are

 

repeatedly moistened and dried.

In many species of Coenogonium, the margin of the

 

apothecium may be present, but rarely, it is absent.

The young apothecium of g. acrocephalum has a thin, unrolled

 

margin which thickens as the apothec1um develops, later
becoming thinner again and finally disappearing at the
maturity of the apothecium. The margin is oftentimes entire
or smooth, rarely crenulate or subdenticulate. In C.

moniliforme, the margin of the apothecium becomes fringed

 

after it is fully developed, while in g. curvulum, the margin
is uneven; or it may even be seen wearing away, as in g.
ornatum. This margin is not always concolorous with the

apothecial disk as observed in many species of Coenogonium.

 

Generally, the margin is paler in color than that of the
disk, or it may be fleshy to whitish-carneous and sometimes
white. It was observed to turn paler very frequently when
the dried material is wet with water.
The apothecium has a well—developed proper exciple.
It is largely euparaplectenchymatous and is easily distinguished

from the other tissues of the apothecium (Pl. 1, Fig. 6).

86

The hypothecium is pallid or colorless and is formed from
thin, delicate hyphae that are woven closely and irregularly,
sometimes becoming conglutinate. In g. velutinum, the
hypothecium is yellow in color and paraplectenchymatous, a
characteristic that has been observed likewise in g. curvulum,
where the cells are globose to angular in shape. The hypo-

thecium in g. interpositum and g. Leprieurii is composed of

 

paraplectenchymatous tissue, the cells of which are angular
and thin-walled. In 9. Linkii, the hypothecium is very thin,
often becoming evanescent as the apothecium matures.

There are neither algal nor medullary layers in the
apothecium of Coenogonium.

The hymenium consists of the asci bearing the spores
and the paraphyses. The hymenium ranges from 40-75 u in
thickness and is generally colorless. In 9. velutinum the
hymenium is light yellow in color and reaches from 100-110 u
in thickness. In fresh Specimens, the hymenium has a
gelatinous texture. The asci are thin-walled and 8-spored.
They are generally clavate but can occur in various other
Shapes; such as, sub—elongate, sub-cylindric or cylindric-
clavate to fusiform-cylindric with round apices, and some-
times as sub-clavate in Shape. Asci vary in size from 4-8 u

in width but may attain a width of 20 u as in g. Leprieurii.

 

87
The anatomy of the apothecium, the characteristic

thin-walled asci and the structure of the spores of

the genus Coenogonium are very similar to those among the

 

members of the family Gyalectaceae.

The paraphyses occur as filiform Slender threads
between the asci. They are free and loosely arranged and
not at all conglutinate. These sterile structures are
either septate or aseptate and are never observed to form
branches. Their apiceS are variable and may be clavate or
capitulate (capitate), round or globose to sub-globose-
clavate, occasionally appearing as obovoid or truncate. The
filament is commonly from 1-l.5 u in width although it may
become as wide as 3.5—4.5 u, as in g. Linkii. The apex
is generally about 3 times wider than the main filament, or

it may not be wider at all, as in Q. velutinum (Pl. 2, Figs.

 

1-5).

The ascospores are either simple or septate (two-
celled) and are always observed to be hyaline and thin-
walled. They vary in shape from ovate to fusiform and
such intermediate forms as oblong, ob—ovate, elliptic
and fusiform-elliptic (Figs. l-5). A distinct character—
istic of g. curvulum which easily separates it from the

other Species of Coenogonium is the arcuate Shape of its

 

88

spores (Pl. 2, Fig. 6).

The terms used to describe the shape of the spores
here are taken from the Systematics Association: Descriptive
Terminology of Taxon, 1962. Although this publication is
meant for plane surfaces and not for tri—dimensional forms
like the spores, for instance, it has been found useful and
proper, however, when used to describe optical sections.

The descriptive term, fusiform, has been added to the
terminology to correspond to the forms which are linear in
shape but are broader in the middle and have rounded ends.

The spores may be arranged in various ways in the
ascus. In many species, they are linear and uniseriate;
in others, they are obliquely uniseriate. Biseriate spores
are found in Q. Linkii, Q. curvulum, Q. moniliforme and g,
acrgpephalum. Occasionally the spores are irregularly biseriate,
as has been observed in Q. interpositum and in Q. disjunctum.

.g. velutinum has been described as having its spores arranged

 

in a sub-uniseriate manner in the ascus.

The size of the spores vary from 5-15 u in length
and from 2-4.5 u in width. .9- disjunctum has been reported
and observed by the author to have the largest measurement in
length; 11-15 H; while 9. implexum and Q. Linkii have been ob-

served to measure up to 4.5 u in width.

 

89
Development of the Apothecium

Karsten (1861) described the development of the
apothecium as consisting of a branch cell of the alga
that becomes a free central cell surrounded by hyphal
elements which develop gradually into a globose ball. He
considered the central cell as a sporangium that starts the
development of the apothecial initial. Karsten compared these
structures to the archegonia of higher cryptogams, adding
that some cortical layers below these round apothecial
initials "lift themselves up" and empty their contents into
So—formed openings or holes on the wall of this globose ball,
a process which he interpreted as fertilization.

Schwendener (1862) disagreed with the views of
'I<EiJrsten and considered the "holes" referred to by Karsten
as the lamina of the fungal threads that surround the young
a1 gal branch. Schwendener further disproved the idea of
fertilization Since he did not observe any formation of
supposedly male organs.

Nylander (1862) criticized the descriptions made
by Karsten of the tissues comprising the hypothecium and

his discussion of the development of the paraphyses and

aSci. Regarding the "cellulose layer" or the "cortical

 

9O

envelope of the apothecia" and the "cuticle" mentioned by

Karsten in discussing the fructification of Coenogonium,

 

Ifixrlander believed that these structures have nothing to

Cic> with fertilization nor with the production of the hymenium.
flDrle "central cells" believed by Karsten to be the primordial
czealls of the apothecium are nothing else but the cavities

()1? the young branches which are in the form of small tubercles,
eacncording to Nylander. The "free endogenous cells" observed
133? Karsten in the cuticle of the young apothecium are not
true spermatia.

Karling (1934) presented an interesting contribution
t:c> the studies on the structure and development of Q. Linkii.
116: described the earliest stages in the development of the
aE3<Dthecium as consisting of a whorl of short mycelial branches
Vvlljlch, with subsequent growth, become more and more numerous
Eilici entangled. This apothecial initials grow to considerable

E3i=ze and become definitely spherical. Karling did not describe
t:11€3 origin and development of the ascogenous layer beyond
kless observations on the formation of a compact pseudo-
parenchymatous mass which later becomes the apothecium.

The development of the apothecium in Coenogonium

 

‘3<3es not differ markedly from that of the other related

Sienera of lichens. In most lichens, an apothecium is

 

 

1‘3“.,P»L

91

initiated in the medulla or algal layer of the thallus
and first appears as a closely woven hyphal ball, the
lower side of which may reach the substrate but does not
merge with it. The upper portion grows out of the thallus
and at maturity the apothecium appears to rest on the sur—
face of the thallus.

In the genus Coenogonium, on the other hand, the

 

apothecial initials are distinguished easily at the early
stages of their development as small nodules on the young
branches of the algal filaments. They do not possess any
interior or central cavity at the beginning. These young
branches are globular outgrowths, white and heteromorphous,
that arise from the side of the algal filaments. They are
entirely composed of a fairly obscure tissue of hyphal cells
which are distinctly formed, and which never contain any
angal elements (Pl. 4, Fig. 6). When the young globose
apothecium attains a diameter of about 0.2 mm., there is
Observed in its center the first signs of a hymenial layer
of about 35-40 LL thick. Parallel, short-celled hyphae
perpendicular to the surface project from the middle
portion of this round structure to its edge and form a
layer of about 12—20 11 thick. These are the rudiments of

Paraphyses which later become frequently branched and resemble

 

92

the irregular sterigmata of a fungus fructification. The

internal structure of this young apothecium consists of a

dense, interwoven, entangled fibrous mass oftissue which

later differentiates into the hypothecium.

The surrounding hyphae which appear to support the

apothecia become embedded deeply in the basal part of the

developing hypothecium which, at this stage of development,

is appressed against the loose reticulum of algal cells

beneath it. Commonly the adjacent hypahe around the apothecial

initials become entangled and as the apothecium develops,

become included in the tissues of the fructification.

AS development of the apothecium proceeds, the

differentiation of the tissues takes place in the excipulum.
Certain filaments: near the surface consisting of large

Cells grow at right-angles to the surface and remain distinct

from the deeply-situated hyphae. These hyphae continue to

grow in this course until a thicker growth occurs in the

o“Vt-er part of the excipulum. The peripheral ends of the

threads thus form a fine paraplectenchymatous covering,

which in effect, always differentiates sharply, while the
other cells beneath these expand themselves considerably to

form the paraplectenchymatous tissue of the hypothecium (Pl. 1,

Fig. ,1)... As the apothecium reaches a diameter of 0.4 to

93

(3.5 mm., the hymenium forms a clearly defined layer of about
‘4() to 50 u thick and paraphyses, asci and spores are developed.

This angiocarpic pattern of development of the
aag>othecium is characteristic for most lichens which have a
{Dzroper exciple.

Whether some cells in the apothecial initials function
ans: spermatia, or whether the process of fertilization takes
place in the young apothecium of Coenogonium remains to be
Iparoven. Meanwhile, the assumption that fertilization of a
3((3ung ascogonium does take place is highly probable and
i.t: is hoped that future investigators of this lichen genus
vvjmll turn their interest towards this phase of the fungus

morphology .

Pycnidia
The asexual fruiting body of the Coenogonium is the
E>§?<:nidium. Two forms were observed among the collections
Eitllldied in the present work. One is spherical and the
c>ilfler is pear-shaped. These structures occur attached
‘:(3 the filaments of the thallus on either the upper or lower
a‘~-‘11:~faces of the thallus. The white color of the pycnidium
13EErsists as such even after it has reached maturity or even

141 the dried condition of the plants.

TH!

 

94

The wall of the pycnidium is made up of an inter-

ivc>ven mat of hyphal threads which are thin-walled and septate

'tllroughout. It does not appear to have any markedly different-

j_atted layers, since the outer portions of the wall gradually
nnearge into the fertile layer beneath.

Conidia are formed within the pycnidia in an exo-
koaisidial manner. These conidia are fusiform in Shape, straight,

Eirud are two—celled. They vary in size from 9-11 x 2-3 u.

'Ifiie conidiophores which bear the conidia are very slender

1:11reads, septate, long and flexuous and have been observed to

Ialranch very rarely except near the base (p1, 2, Fig. 7),

Sterile filaments, called "anaphyses" and "sterigmata"

13)? various authors, occur between the conidiophores. These
Slender filaments are elongated and generally simple, but
rnéijg oftentimes be seen to branch.

The pycnidia were found in most forms of the lichens
tzklialli, from the shelf—like forms to the expanded or
l"1<>J:-izontal types of growths. It is obvious that asexual
regEbroduction of this type in Coenogonium is not restricted
‘:<3 the type of thallus nor probably to the species of algal

stbiont present.

In the course of the study of herbarium material,

5‘ number of collections were found with well—developed pycnidia

95

kNJt no apothecia. These collections, therefore, could not be

ixientified but are recorded here.

Material seen: MEXICO: Cosolapa. 1200 m., Veracruz,
G?L1rpus, 3, 1921-22 (us).
On bark of trees, 1100 m., La Palma

COSTA RICA:

Province of Alajuela, Brenes 53, 1925? (NY):

de San Ramon,
Brenes 14484, February 1931

San Ramon, Province of Alajuela,

(IWY); Colinas de San Ramon de San Pedro, Province of Alajuela,

Brenes 18998, 18 January 1934.

CUBA: Woods, 900 m. alt., Cobre Range of Sierra

Leon, Fre. Clement

Maestra, Loma del Gato & vicinity, Fre.

é; IFre. Roca 10117, 11 July—l4 August 1921 (my).

DOMINICAN REPUBLIC: In primitive cloud and.rain

fC>.J:'est on top of Loma Campana, Province of La Vega; alt. about
J-(3136 m., Allard 18709, 1 January 1948 (US); cloud forest,

rchSge above Los Amaceyes, 300—3200 ft., Cordillera

SeaIptentrional, Imshaug 23305, 2 August 1958 (MSC).

PUERTO RICO: Luquillo Mountain, Wilson 125, July

J-SBCDZ (NY); on Acrista monticola, Monte Torrecilla, 900-100 m.

all‘t., Britton, Cowell & Brown 5666, 20 March 1915 (NY).

BOLIVIA: Yungas, 6000 ft., Rusby 280, 1885 (my).

BRAZIL: Bahia, Wilson, (No date) (US).

AUSTRALIA: Tarwin, Leighton, (No date) (NY).

 

96

gm

General

For the taxonomy of the genus Coenogonium, the

writer is inclined to Share the views of Santesson (1952)

that the spores and apothecial characters, rather than the

d ifferent morphology of the symbiotic algae, Should be used

as bases for the distinction of the different families and

for the establishment of the relationships of Coenogonium

 

with the other genera. It is my opinion, however, that this

genus Should remain in the Family Coenogoniaceae and that

Racodium be excluded from this family until the apothecial

 

characteristics of the latter are definitely established.
It is believed that the effect of the fungus on thealgal
symbionts to produce the different types of thalli merits
the separation of this family from the Gyalectaceae.

The following characteristics are being considered

in the present study as of importance in the taxonomy of the

genuS Coenogonium: the type of Spores, external features

of the apothecia (whether sessile, stalked, flat, convex,
lmarginate, etc.), and the special types of thalli formed
with the fungus. As to the Spores, it is especially

the ir septation, Shape and size as well as their arrangement

97

in the ascus that are of taxonomic importance. The septation

of the paraphyses and the Shape of their apices can constitute

some of the diagnostically important species characters.

Other morphological characteristics, such as, the color of

the apothecial disk, the size of the margin, as well as

the size of the apothecia are usually not constant to the

same degree and are, therefore, being considered here as

"erpporting" characters in the delimitation of the different
S[Decies of Coenogonium.

Before identifications of the material were made,
all. the original descriptions of the Species were collected
arms a literature key of the whole genus was made. It was
hfinped that the descriptions might provide an over—all under—
stlandingof the nature and range of variations of the species
so that they could be separated on a comparative basis: and
a1 so that such a method might serve as a guide in determining
t}1€: delimitation of the different species. For this purpose,
:11: XNas necessary that large numbers of specimens had to be

egamined and studied.
The apothecia of the dry Specimens were examined

by soaking them in distilled water and teasing the material

to separate the asci and the paraphyses from the rest of the

fungal tissues. Sections of the apothecia were also prepared

98

for the study of the different anatomical structures.
Below is a list of the herbaria from where the
dried Specimens were obtained. (Abbreviations according
to Lanjouw and Stafleu, Index Herbariorum, Part II, 1954):
M0 - Missouri Botanical Garden, St. Louis, Missouri
NY - New York Botanical Garden, New York, N. Y.
US - National Museum, Smithsonian Institution,
Washington, D. C.
MSC - Michigan State University, East Lansing, Michigan
MICH - University Herbarium, University of Michigan,
Ann Arbor, Michigan
FLAS - Herbarium, Agricultural Experiment Station,
Gainesville, Florida
TENN - Herbarium of the University of Tennessee,
Knoxville, Tennessee
Earlier authors in searching for knowledge and
experience concerning the disposition of the lichenized
Trentepohlia (-Chroolepus) or the sterile forms of Coenogonium,
established a large number of species of this genus. Of the
original forty—eight species described as and referred to
Coenogonium, about, half of this number were found to have
been based on Specimens with a sterile lichen—fungus or a

lichen-fungus with immature apothecia. Four species were

TH!

 

99

determined as belonging to the Fungi Imperfecti (Santesson,
1952) and Six Species have been transferred to the genus

Trentepohlia, three as valid algal species and three as

 

synonyms of Trentepohlia (Hariot, 1890, 1891). Fifteen

 

species are accepted as valid in the present study and a key
to these Species has been made. Two species are treated

as synonyms of g. Linkii and eight species are considered as
doubtful or insufficiently known. Twelve Species are being

rejected as nomina dubia and one Spec1es is considered as a

nomen confusum.

Lichenized Hyphomycetes

The species belonging to this group have been
observed to be lichenized Hyphomycetes with dark brown hyphae
and the fungi probably belong to Racodium or Cystocoleus

in the Fungi Imperfecti.

1. Coenogonium ebeneum (Dillw.) Smith, Handbook of British

 

Lichens. 75. 1921. Conferva ebenea Dillw. British
Confervae, t. 101; Introd. no. 79. 60. 1809.

2. Coenogonium germanicum Gluck, Flora 82:268—285.
1896.

3. Coenogonium nigrum (Huds.) A. Zahlbr. Ann. Naturhist.
.Mns. Wien. 25:241. 1911. Byssus nigra Huds. Flora Anglica.

487. 1762.

100

4. Coenogonium Schmidlei Simmer Allgem. Botan.

 

Zeitschrift. 5:190. 1899.

Non-Lichenized Trentepohlia

 

The following Species attributed in the past to the

lichen—genus Coenogonium have already been transferred to

 

Trentepohlia as valid algal species or synonyms, presumably
on the fact that no trace of fungal hyphae were present on
the thallus:

1. Coenogonium deplanatum Krempelh. Flora 59:250.
1876. Treated by Hariot in 1891 as a synonym of Trente-
gpohlia setifera Farlow.

2. Coenogonium depressum Mfill. Agr. Flora 64:525. 1881.

Transferred to Trentopohlia by Hariot (1890) as a valid

 

algal species.

3. Coenogonium dialeptum Nyl. Ann. Sci. Nat. Bot.

 

IV. 16:90. 1862. Transferred to Trentepohlia by Hariot
(1889) as a valid algal Species.

4. Coenogonium diffractum Krempelh. Flora 59:250.

 

1876. Transferred to Trentepohlia by Hariot (1891) as

 

a valid algal species.
5. Coenogonium effusum Krempelh. Flora 59:250.
1876. Treated by Hariot (1891) as a synonym of Trente-

pohlia setifera Farlow.

101

6. Coenogonium simplex Mfill. Arg. Flora 73:16.
1890. Treated by Hariot (1891) as a synonym of Trente-

pohlia dialepta (Nyl.) Hariot.

Partially-Lichenized Trentepohlia

The following species (or varieties) were described
and established by various authors from thalli without
apothecia or with only immature apothecia. In the absence
of spores, it is impossible to identify these in Coenogonium
and although fungal hyphae were present, it is best to treat
these species in the algal genus Trentepohlia.

1. Coenogonium afrum Mass. Memor. I. R. Instit.
Veneto. 10:43. 1861. Chroolepus afrum Mass. Memor.
I. R. Instit. Veneto. 10:43. 1861. - Orig. coll.:
Africa australis, E§E£Q.

2. Coenogonium cancellatum Leight. Trans. Linn.
Soc. London 27:172. 1869. - Orig. coll.: New
Eliya, Central Province, Ceylon, Leighton.

3. Coenogonium confervoides var. arborum Nyl. Ann.
Sci. Nat. IV. 16:92. 1862. Conferva arborum Ag. Syst.
Algar. 88. 1824. - Orig. coll.: Marianas Is. and Brazil,

Gaudichaud.

102

4. Coenogonium dialeptizum Stirt. Proc. Phil. Soc.
Glasgow 11:103. (1878) 1879. - Orig. coll.: Upper
Amazon, Brazil, Trail.

5. Coenogonium Echinus K. Mfill. Bot. Zeit. 15:387.
1857. Orig. coll.: Brazil, §§r1.Mfiller.

6. Coenogonium heterotrichum Mfill. Agr. Bull. Soc.
Bot. Belg. 32:162. 1893. - Orig. coll.: San Marcos
de Dota, Costa Rica, Tonduz 6115.

7. Coenogonium patagonicum Mfill. Arg. Flora 71:47.

1888. - Orig. coll.: Patagonia and Australia, Caracciolo

8. Coenogonium pulvinatum Krempelh. Verh. Zool. —
Bot. Gesellsch. 26:445. 1876. - Orig. coll.: Lima,
Peru, Barranca,

9. Coenogonium rigidulum Mfill. Arg. Flora 65:490.

 

1882. 4 Orig. coll.: Queensland, Australia, Wilcox.
10. Coenogonium subtorulosum Mfill. Arg. Jour. Linn.
Soc. Lond. Bot. 32:207. 1896. - Orig. coll.: New

Zealand, Colenso 1656.

ll. Coenogonium subvirescens (Nyl.) Nyl. Flora 57:72.
1874. Coenogonium Leprieurii var.subvirescens Nyl.
Ann. Sci. Nat. Iv. 16:89. 1862. — Orig. coll.:

Rio Negro, Amazon, Brazil, Spruce 28.

-103

This species was distinguished from g. Leprieurii
on the basis of its thinner algal threads. As Nylander's
specimens were sterile, it cannot be proved that this species
is not identical to g. Leprieurii.

12. Coenogonium tenuissimum Krempelh. Nuov. Giorn.

Bot. Ital. 7:26. 1875. - Orig. coll.: Borneo, (collector
unknown), 1886.

This species was described by Krempelhuber as a
young form of Coenogonium. According to Mfiller (1890),
the species resembles g. depressum which was, however,
transferred in 1891 to Trentepohlia as an algal species

by Hariot.

Lichenized Coenogonium

The following eight taxa were described from specimens
with ascocarps. No material of these taxa, however, were
available for study and they are not included in the key.

1. Coenogonium boninense Sato, Jour. Jap. Bot.
8:390. 1933. - Orig. coll.: On bark of tree, Bonin:

Mt. Sekimon, Hahazima Island, Japan, Sato 106.

 

Thallus lamellate, spongiose—byssoid, spread in
a flabellate form, 10-25 mm. wide and 5-15 mm. long,

Sometimes imbricate and overlapping, one margin attached

104

to the bark of tree, ashy-green; alga Trentepohlia, cells
24-30 x 7-9 M; hypha covering algal filaments about 15 0
wide. Apothecia about 10-20 on the thallus. Apothecia
round, about 0.5 mm. wide, Situated on both the lower and
upper surfaces of thallus: disk flat to somewhat convex,
carneous or pale yellow, margin thin, pale; hymenium wine
red in iodine; paraphyses wide continuous, 60—65 x 1.5 u,
septate, not branching, apex capitate: asci cylindrical,
45-50 x 4—6 0; 8-spored; Spored colorless, ellipsoid,
simple, 5-8 x 3.5 u.

2. Coenogonium congensis Dodge, Ann. Miss. Bot. Gard.

 

40(4):350. 1953. - Orig. coll.: Belgian Congo, Yangola,
20 km. west of Yangambi, J. Louis 12070.

Thallus dimidiate; apothecia convex, sessile, orange—
yellow, margin concolorous, slightly dentate: paraphyses
unbranched, tips not thickened; spores uniseriate, fusiform,
bilocular, 6 x 2.5 u.

3. Coenogonium Deightoni Dodge, Ann. Miss. Bot. Gard.
40(4):349. 1953. - Orig. coll.: Sierra Leone, Njala
(Kori), F. c. Deighton M4396 a.

Thallus very thin: apothecia sessile, flat, disk
cinnamon-brown, margin paler: paraphyses unbranched, congluti-

nate, tips moniliform but not thickened; spores fusiform,

 

105

simple, 6-8 x 2.5 - 3 u.

,4“ Coenogonium epiphyllum Wain. Ann. Acad. Sci.
Fenrl. 15(6):156. 1921. — Orig. coll.: Luzon,
Phillippines, Robinson.

Thallus a thin crust, adnate, ashy-yellow; apothecia
ndxnate, concave, pale white, margin paler, thin, prominent:
Paraphyses wide, septate, apex capitate-clavate: spores uni-
tO loiseriate; fusiform to fusiform-elliptic, septate, 6-8 x
2.55 - 3 u.

Santesson believes that this represents a good
autonomous species. Wainio reported the type as "parce
lexztum, duobus apotheciis," and growing on leaves of
trees.

5. Coenogonium interplexum f. contextum Stirt.

Prtmn Phil. Soc. Glasgow 10:297. 1877. - Orig. coll.:
WEllington, New Zealand, John Buchan§3,.§fig.

The description given for this species indicated
t1lat it is similar to Q. interplexum except for the reaction
‘3f the apothecium to iodine solution (violet-red color).

DMD further information was given on the characteristics
0f’the lichen-fungus. Only a study of the Spore characters

can the exact relationship of this form of g. interplexum

 

be determined.

106

ES- Coenogonium interponendum Nyl. Jour. Bot. 15:225.
1877 - - Orig. coll.: On trunks and leaves of trees in old

forests, Costa Rica, H. Polakowsky 496.

 

Thallus adnate to effused; apothecia yellowish—
carneous; Spores 8 x 2.5 H, septate, fusiform; algal
filaments 17-20 u wide.

Nylander reported that this species is similar to

lo

interpositum except in the size of the algal filaments
0f the latter which he described as from 14-28 LL wide.
If the separation from g. interpositum is actually based
only on the algal characters, this Species should be a

Synonym of g. interpositum.

 

7. Coenogonium retistriatum Leight. Trans. Linn. Soc.

LOnd. 27:172. 1869. — Orig. coll.: Ceylon, Leighton, s.n.

 

No spore measurements were given in the description
Of this species and it is not possible to identify the
fuhgus on the basis of its straited fungal elements alone.
8. Coenogonium Tuckermani Mont. Ann. Sci. Nat. IV.

7:143. 1857. - Orig. coll.: Venezuela, Fendler, s.n.

 

Based on an imperfect specimen, the description of

this species given by Montagne is very inadequate.

107
Key to the Species of Coenogonium

Sporessimple.....................2
Sporesone-septate...........’.......6
2 . Thallus thin, circular or spongy—byssoid,

imbricate, Shelf-like: apothecia sessile,

flat, or convex, margins thin, pale, uneven:

spores uniseriate, oblique, ovoid-elliptic to

Slightly fusiform-elliptic, 6-10 x 2—4 0;

paraphyses apex clavate or capitate

. . .lg. Leprieurii

2. Thallus amorphous or adnate to effused, expanded or

dense, caepitose—tomentose . . . . . . . . . . . . 3
.Apothecia flat or convex, subsessile, margins thin:
paraphyses not septate, apex not wide, subtruncate . . 4
Apothecia flat to concave, sessile, margins thick,
entire: spore fusiform: paraphyses septate:
thallus dimidiate or caespitose—tomentose . . . . . . . 5
4. Margin entire: spores irregularly biseriate,

elliptic and acute on both ends, 6-10 x 2-3 u:

thallus adnate, expanded . . . . . .‘g. interpositum

 

4. Margin wearing away, fringed: Spores uniseriate,
narrowly fusiform, 8—10 x 2-4 u; thallus

amorphous . . . . . . . . . . . . .g. ornatum

108

Spuores biseriate, fusiform, 7.5-9 x 2.5-3 0;
pairaphyses apices abrupt and widely capitulate:

triallus dimidiate . . . . . . . . . . . .‘C, acrocephalum

 

SENores uniseriate, 8—11 x 2-2.5 u; paraphyses
agpices ob-ovoid; thallus filaments short,
(Zaespitose . g. tomentosum
ES. Thallus prostrate and thin, or adnate, expanded

or spongy—byssoid . . . . . . . . . . . . . . . . . 7
ES. Thallus shelf—like or crust—like . . . . . . . . 13
Spores uniseriate, thallus adnate to effused, pannose,
Gaxpanded or loosely tomentose . . . . . . . . . . . . . 8
:Spores sub—uniseriate to biseriate; apothecia sessile:
jparaphyses not septate . . . . . . . . . . . . - . . 12
£3. Apothecia sessile, flat, margins thick, smooth,

white: Spores ovate or Short—elliptic, 7-12 x

2—4.5 u: paraphyses septate . . . . . . ..g. implexum
£3. Apothecia subsessile or stalked: paraphyses

septate . . . . . . .

.Apothecia flat to concave, margins uneven, pale;

Spores fusiform-elliptic, 6-9.5 x 2-3.5 u . ‘9. complexum

 

Apothecia flat to convex, margins smooth, thin or

Slightly thickened, white or pale . . . . . . . . . . 10

109

 

 

 

 

1C). Spores elliptic or fusiform-elliptic . . . . . 11
1C)- Spores fusiform, 6—7 x 1.5-2 0 . . . . . g, pannosum
11. Spores 6.5-10 x 2—3.5 u . . . . . . . . .< C. interplexum
ll. Sporelex3u. . . . . . . . . . . . . . _C_.botryosum
1J2. Spores sub-uniseriate, fusiform—
elliptic, 9-11 x 3—3.5 u; apothecia
concave to barely convex, margins thin,
smooth, pale; paraphyses apex not at all
wider: thallus spongy-byssoid consisting of
short, aggregated filaments . . . . . . _g. velutinum
212. Spores biseriate, curved, 8-9 x 2-3 0;
apothecia flat to concave, margins thick,
smooth, white: paraphyses apices sub—globose
to clavate: thallus thin consisting of long,
prostrate filaments . . . . . . . . . . ..g. curvulum
13- Apothecia flat, sessile or rarely sub-sessile, margins
thin, smooth, white: spores uniseriate to biseriate,
short-fusiform, 9-13 x 3-5 0; paraphyses non—septate,
apices, globose—clavate: thallus crust-like, filaments
moniliform; phycobiont a Physolingm‘. . . '9. moniliforme
13- .Apothecia flat to convex, subsessile: paraphyses
septate, apices sub-globose or globose to clavate:
filaments not moniliform: phycobiont a Trentepohlia 14

 

110

1L4“ Nbrgin thin, smooth or sometimes rugged, pale:
spores biseriate, ovate or Short-fusiform,
6.8-12.5 x 2-4 0: thallus shelf—like,
imbricate . . . . . . . . . . . . . . . . g, Linkii
214. Margin toothed or slightly toothed,
white: spores irregularly biseriate,
fusiform, 11-13 x 2-4 u: thallus adnate to

effused, pannose, expanded . . . . . g, disjunctum

 

Coenogonium acrocephalum Mfill. Arg.

Fflxxra 63:525. 1881. - Orig. coll.: Apiahy, Brazil,
P_uigqari 156.

Thallus dimidiate, yellow, becoming whitish beneath:
algal symbiont Trentepohlia. Apothecia 7.5 mm. in diameter,
rc>und, sessile: disk flat to concave, orange-carneous to
Palen margin of young apothecium thick, becoming thinner and
tjuen immarginate at maturity, entire: hymenium 55 0 high,
E3E>Iithecium distinct: paraphyses loose, 3-4 u wide, septate,
aFNices abrupt and widely capitulate: asci cylindrical, 8-
SENDred; spores simple, biseriate, fusiform, 7.5-9 x 2-3 0.

This species is related to Q. Leprieurii but differs
from it by its larger spores and wider apothecia with entire

margin. The paraphyses of g. acrocephalum are wider, very

111

loo3e and the apices abrupt. Mfiller described the spores of
this species as 7-8.5 LL long. In my studies of the specimens
from the West Indies, the Puerto Rico material measured
7.4 X 3.3 LL in size, while the collection from Cuba had much
longer Spores, up to 9.3 0..

Material seen. CUBA: El Guama, Palmer & §i_l_e_y
33$, 24 March 1900 (US). PUERTO RICO: Luquillo Mts.,

Film 100, July 1902 (mg).

Coenogonium botryosum Knight

SYnOps. Queensland Flora. I. Suppl. 74. 1886. -
Orig. coll.: On bark of trees, Mt. Perry, Knight.

Thallus dense, velvety, consisting of closely
aggregated short threads together with a few very slender
filaments, both cohering by very short funiculi: yellow-
grESn; algal symbiont Trentepohlia, cells of filaments 1—2
times longer than wide. Apothecia sub-sessile, round:
disk flat, yellow-orange; margin paler, uneven: hypothecium
C01<>r1ess; paraphyses septate, 1.4—1.7 u wide, apices
clavate to globose, capitate, 3.3 - 4 11. wide; asci clavate,
8‘Spored: spores one-septate, elliptic, uniseriate, ca.

10 x 3 0..
Some algal cells in the thallus of this species appear

1ike the elliptic cells in g. moniliforme. I found the algal

112

filamuants in the material slightly narrower than the Size
repor1:ed for the species by Knight but this character is
notza reliable taxonomic difference.

Material seen: MEXICO: Veracruz, Turpus 223 (US).

 

Coenogonium complexum Nyl.

Ann. Sci. Nat. IV. 11:222. 1859. — Orig. coll.: On
trees in Bolivia, Weddell.

Thallus adnate, pannose, expanded, golden—yellow;
alQEfiL symbiont Trentepohlia, filaments fasciculate-conglutinate,
SCaInzely branching, cells 2.5 - 5 times longer than wide.
APothecia round, subsessile; disk flat to concave, carneous-
Yellxmu to yellow-orange: margin paler, thin, uneven: paraphyses
sePtate, 1.5 — 2 11 wide, apices sub—globose, capitate, 3.3 -
41L wide: asci cylindrical, 8-Spored: Spores one-septate,
oblong to fusiform-elliptic, 6—10 x 2.3-3.2 u.

Nylander described the spores as 6-10 x 3-4 u. I
found the Spores of the material from Central America and
thfii West Indies to have the same range in Size (6e8.5 x 3 u)
Enua those from South America to be larger (10 x 3.2 u).

Material seen: COSTA RICA: South bank of Rio

Pejivalle, 650-800 m., Cartago Province, Dodge & Thomas 4408,

 

26 September 1929 (MO 70722); forest near farmhouse in Finca

113

CastiIlla, 30 m., Limon Province, Dodge & Groeger 9207,

 

23 Jilly 1936 (M0 154712). MEXICO: On bark, Cuernavaca,
6500 ft., Pringle, 31 October 1908 (US, MICH). PANAMA:
On thig, drowned forest in valley of R. Puente between the
Tunnral and Natural Bridge, 70 m., Canal Zone, Egggg &
Alleg 8863, 6 January 1935 (M0 154705).
COLOMBIA: Rio Magdalena, ngr, (no date) (NY).
HAITI: On twigs, elfin woodland forest, north
0f I?oret des Pins (Shada station), near Dominican border,
580C) ft. (Dept. de l'Ouest), Wetmore 2931, 14 July 1957
(MSC). TRINIDAD: On Theobroma cacao, St. Pat's Ravine,

AriJna Valley, Fleming (MSC)-

Coenogonium curvulum A. Zahlbr.

Ann. Crypt. Exot. 1:164. 1928. — Orig. coll.: On
leaves, Java, Schiffner 3056.

Thallus prostrate, adhering to substrate, green:
algal symbiont Trentepohlia, filaments more or less dichoto—
mmlsly branched, cells Short-cylindric, 2.5 — 3 times longer
tfiuin wide. Apothecia 0.1 - 0.3 mm. in diameter, biatorine,
Irc>und, sessile, slightly constricted at the small base:
disk slightly concave, pale yellow-carneous: margin white,

Prominent, entire; excipulum paraplectenchymatous: hypothecium

114

colorless: hymenium 40 - 55 0 high; paraphyses filiform,
Simple, 1.5 u wide, apices distinctly sub—globose to clavate,
3.8 u wide, conglutinate; asci sub-elongate to cylindrical,
8-Spored; spores one-septate, biseriate, oblong to elliptic,
round on both Sides, curved or arcuate, 8—9 x 2-3 u.

Zahlbruckner described the size of the cells in the
algal filaments as 14—16 x 17-19 u. In the single Specimen
examined, I found the algal cells to measure 6-8 0 wide and
about 2.5 to 3 times as long. This difference in Size is
not, however, important in the taxonomy of the Species. The
characteristic curved spores are considered the significant
diagnostic feature.

Santesson (1952) reports this species as probably
more or less obligately foliicolous. The material examined
in the present work, however, was found on a bark substrate.

Material seen: PUERTO RICO: Vicinity of Mayaguez,

 

Britton & Marble, 4—10 March 1906 (MICH 13423).

 

Coenogonium disjunctum Nyl.

Bot. Zeit. 20:178. 1862. - Orig. coll.: On trunks and
dead wood, Martinique and Cuba (collecter not known).
Thallus adnate, pannose, expanded, yellow-green:

algal symbiont Trentepohlia, filaments cylindrical, 3—5

 

115

times longer than wide. Apothecia 0.6—0.7 mm. in diameter,
round, stalked; disk flat, carneous-yellow, margin pale,
thin, smooth sometimes rugged: paraphyses septate, 1.5 0
wide, apices sub—globose to clavate, 3.5 - 4.5 0 wide;

asci cylindrical, 8—spored; spores one—septate, irregularly
biseriate, fusiform, 7.5 — 13 x 2-4 u.

This species is easily distinguished from g, Ligkii
by the effused and adnate thallus and the larger size of the
spores.

Specimens from the United States and Central America
had the same spore sizes (ll-l3 x 2-4 0), while those from
the West Indies were smaller, only 7.5 u long. The material
examined from the Philippines had relatively the largest
size of spores, l2—13.2 x 2.8 - 3 0.

Material seen: ALABAMA: (locality and collector

 

not known) (Willey Lichen Herbarium) (US).
COSTA RICA: Morpho Valley, alt. 4400 ft., Danilson
51(Date not known) (MICH). HONDURAS: On twigs, Landetilla

Valley, Chickering 148 d, June-July 1929 (MICH). MEXICO: On

 

tree bark, Jolapa, Pringle, 25 June 1908 (MICH 15407).
PANAMA: Summit forest, Cerro Campana, Bartlett & Lasser,
1 September 1940 (MICH 16936).

PUERTO RICO: On twigs, Rio Icaco and adjacent hills

116

altitude 465 to 720 m., Sierra de Naguabo, Shafer 3636,
30 July - 5 August 1914 (NY).

PHILIPPINES: On trees,and rocks, 2300 ft. elev.,
Lamao River, Mt. Mariveles, Bataan, Central Luzon, Williams

05, 2 January 1904 (NY).

Coenogonium implexum Nyl.

Ann. Sci. Nat. IV. 16:92. 1862. - Orig. coll.:
On barks among mosses, Tarwin, Victoria, Australia, Egrg.
Mfiller.

Synonym: IQ. inflexum Hook. Handbook New Zealand
Flora 581. 1867.

Thallus adnate, pannose, expanded, bright yellow;

algal symbiont Trentepohlia, filament cells 2-4 times longer

 

than wide. Apothecia 1 mm. in diameter, round, sessile:

disk flat, carneous-yellow: margin white, thin, entire:
paraphyses septate, 1.5 - 2 u wide, apices clavate, 2 - 3.5 0
wide: asci cylindrical, 8-spored: spores one-septate, uni-
seriate, oblique, short-fusiform, 7—12 x 2.4 — 4 0.

This Species resembles g, interplexum except for the

 

larger and wider paraphyses of the former. Nylander gives
the measurements of the spores as 8-11 x 3.5 — 4.5 u and

the algal filaments as 10-14 u wide. The spores in the

117

material examined had a wider range and I found the algal
filaments very variable in size. The size of the algal cells
as a diagnostic character cannot be generalized in this case.

This species has been reported as foliicolous by
Nylander and Santesson but I have not encountered any
epiphyllous material.

The collections from South America and Australia
were identical in their size of Spores, and those from
the United States and the West Indies had the same length
of spores: the width of the spores in the West Indies
collection was smaller (2.4 u).

Exsicatti: Malme, Austr. Amer. n. 113.

Material seen: FLORIDA: On twig, southeast Florida,

 

J. D. Smith 5;, March 1877 (US); on bark, (no locality),

 

Curtiss, 1879 (US). TEXAS: On bark, Hardin Co., 7 miles

 

south of Silsbee near Bruce Reid's place: alluvial area,

longleaf pine belt, Whitehouse 25968, 19 November 1951

 

(MO 184372). LOUISIANA: On bark of living tree, woods near
Varnada, Washington Parish, McFarland 187, 24 January 1953
(M0 184371).

BRAZIL: Rio Grande do Sul, Porto Alegre (a), 9
September 1892 (Exp. Regn. I. Lich. n. 478), and Cruz

Alta, 17 April 1893 (Exp. Regn. I. Lich. n. 1255), Malme 113

 

118

(MSC). COLOMBIA: (no locality), Mutis, 1760—1808 (US).
JAMAICA: On bark, Kinloss to Barbecue Bottom,

1300 ft., Trelawny Parish, Imshaug 16060 c, 10 May 1953 (MSC).

 

AUSTRALIA: Tarwin, Tasmania, (Collector not known),
.gg.._2, 1855 (US): on tree, Mt. Arthur, Tasmania, Wilson

23 February 1891 (MSC 3237); Warburton, Victoria, Bastow,

March 1902 (MSC).

Coenogonium interplexum Nyl.

 

Ann. Sci. Nat. IV. 16:92. 1862. - Orig. coll.: On

trunks of trees, Colombia, Lindig 2561.

 

Thallus adnate, pannose, expanded, loosely tomentose,
light yellow: algal symbionts Trentepohlia abietina, T. arborum,
.1. elgngata, or T. ggrga, cell shape and filament size
variable. Apothecia 0.5 - 1 mm. in diameter, round, Sub-
sessile: disk flat to Slightly concave, carneous-yellow or
orange, margin whitish-carneous or white, thin, entire:
hymenium 70-75 0 high; paraphyses slender, about 1 u wide,
septate, apices clavate to obtuse, 3.3 0 wide; asci cylindrical,
8-spored: spores one-septate, uniseriate, linear, short-
fusiform or fusiform—elliptic, 6.5 - 10 x 2.5 - 3 u.

This Species is Similar to g. implexum except for

its consistent smaller spores, thinner paraphyses and somewhat

119

longer algal cells. Nylander (1862) described the spores
as 8—10 0 long but I found spores that were shorter and
slightly wider. Malme (1937) has described the range in
length to be 8—11 u. I found the species not to be variable
as to spore sizes. Specimens from the West Indies, South
America and the United States had the same range, while those
from Central America were slightly longer.

Mfiller (1893) and Santesson (1952) reported folii—
colous specimens from Boruca, Costa Rica, (Pittier no. 6114),
but are sterile and unidentifiable. A collection from Ann
Arbor, Washtenaw Co., Michigan, reported by Fink (1935) as

‘Q. interpositum, has characters that closely agree with this

 

species.
Exsicatti: Malme, Austr. Amer. n. 329.

Material seen: FLORIDA: On bark, among mosses at

 

bases of trees, (no locality), Calkins 170, 1889 (MSC 120023);

 

(no locality), Calkins, 1890 (US); bases of trees on moss,

Jacksonville, Duval Co., Calkins 963, (no date) (MSC); at base

 

of oak trees, Jacksonville, Duval Co., (collector not known)
(Willey Lichen Herbarium) (US); on pine tree near base of
stem, Solution Sink, Millhopper's Hammock, 8 miles east of
Gainesville, Alachua Co., Uyenco_l§, 6 February 1963 (MSC):

on bark of citrus, Low Hammock near Sanford, Seminole Co.,

120

Uyenco 65, 9 February 1963 (MSC); on bark of oak tree

(Quercus virginiana), Lieder Trail, Highlands Hammock State

 

Park, Sebring, Highlands Co., Uyenco lg, 11 February 1963 (MSC):

on bark of Lyssoloma, Rattlesnake Hammock, Everglades

 

National Park, Dade Co., Uyenco 109, 13 February 1963 (MSC);

 

on bark of Lyssoloma, Redd Hammock, Everglades National Park,

 

Dade Co., uyenco 114, 14 February 1963 (MSC). LOUISIANA:

 

Foot of trees in swampy woods around Covington, St. Tammany

Parish, Langlois 799, 1894 (US). MICHIGAN: On old trees,

 

Ann Arbor, Washtenaw Co., Smith, July 1932 (MICH).
‘COSTA RICA: On tree, Cerro de la Carpintera,

Province of Cartago; altitude 1500-1850 meters, Standley

 

35565, February 1924 (US). GUATEMALA: On twig, Dept. Alta
Verapaz: wet forest near Tactic, above the bridge across

Rio Frio, altitude about 1400—1500 m., Standley 90425,

 

30 March 1941 (M0 166103); on twig, Dept. Alta Verapaz:
dense wet limestone forest near Chirriacte, on the Peton

highway, altitude about 900 m., Standley 19142, 9 April 1941

 

(MO 166101). MEXICO: Veracruz, Turpus 125, February 1921-22

 

(US).
PARAGUAY: On trees, Grand Chaco: ad Rio Negro.

15 September 1893. (Exp. Regnell. I.), Malme 330 (MSC).

 

BAHAMAS: On bark, Coppice, Stoppes Hill, Crooked

121

Island, Brace 4823, 9—23 January 1960 (NY). JAMAICA: On

 

bark, Dolphin Head, 1750 ft., Hanover Parish, Imshaug 15627 a,
23 April 1953 (MSC). PUERTO RICO: Ravine on estate of
H. Guggenbfihl, vicinity of Mayaguez, Britton & Marble 651,

4-10 March 1906 (NY): Quebradillas, Bro. Hioram, January

 

1913 (NY).

Coenogonium interpositum Nyl.

 

Ann. Sci. Nat. IV. 16:91. 1862. — Orig° col.:

Reunion (Bourbon) Island, Lepervanche-Mezieres & Louisiana

 

(ex. herb. Tuckerm.).

Thallus adnate, pannose, expanded, yellow-green:
algal symbiont Trentepohlia arborum, filaments sparingly
branched, cells cylindrical, about 3 times as long as
wide. Apothecia 0.7 - 0.8 mm. in diameter, round, sub—
sessile: disk at first concave, becoming convex, carneous-
yellow to pale orange: margin pale, thin, entire, rarely
sub-denticulate; excipulum paraplectenchymatous, cells
angular; hypothecium yellowish: hymenium 70-75 0 high,
colorless; paraphyses simple, conglutinate, 1—1.5 u wide,
apices clavate, 2-4 u wide; asci cylindric-clavate,
8-Spored; spores simple, irregularly biseriate, elliptic

to fusiform-elliptic, 6—10 x 2-3 u.

122

Zahlbruckner (1909) and Malme (1937) used this name

3f3<3r a species with one—septate spores and it was also cited

]:>jy Fink (1935) in the same manner. Nylander distinguished

1:;his Species from Q. disjunctum by its smaller spores and

1t1hinner algal filaments. The latter species has distinctly

<:>ne-septate Spores. The Size of the algal cells, however,

;i_s not believed to be a useful taxonomic character, especially

j_n the case of dried specimens where the filaments do not

iambibe water easily, thus rendering the measurements of

aalgal cells extremely difficult and inaccurate for compara-

‘tive purposes. A comparison of spore sizes among the geo-

graphical areas present interesting information. Specimens

from the United States and the West Indies were exactly the

same in spore measurements; those from the Philippines were

6 x 2.5 u: the collections from Central America were 6.3—8

x 2.8-3 u in size. The Spores in the single fertile

collection from Colombia measured 7.5 - 8.5 x 2.2 u.
Exsicatti: Malme, Amer. n. 330.
Material seen: FLORIDA: On trees (no locality),

Sprague, (no date) (MSC 10877); on twig, in hammocks near

the Homestead road, between Cutler and Longview Camps,

Dade Co., Small & Carter 1347 & 1348, 9-12 November 1903 (NY);

on twig, Royal Palm Hammocks, Dade Co., Lowe, 29 January

123

:1-5316 (Tenn.); on bark of trees, Sanford, Seminole Co., Rapp,

May 1928 (FLAS 29040); on trunk of live oak, in low
hammock, Lake Marion, Osceola Co., McFarlin 8, 21 January

$1.931 (MSC); on sweet gum tree, Low Hammock near Sanford,

Seminole Co., Uyenco 61, 9 February 1963 (MSC); on bark of

ssweet gum tree, Lieder Trail, Highlands Hammock State Park,

£3ebring, Highlands Co., Uyenco 18, 11 February 1963 (MSC).

MEXICO: On living tree, Chichen Itza, Yucatan

IPeninzula, Steere, 13 June 1932 (MICH 1253); on tree in

iforest, San Miguel, Cozumel Island, Quintana Rio, Steere,

6-8 August 1932 (MICH 2726).

COLOMBIA: On tree trunk, Dept. E1 Valle: Cordoba:

altitude 50-100 meters; dense forest, Killip & Garcia 33447,
17 February 1939 (US).

BAHAMAS: (no locality), Eggers 4463, (no date) (NY).

DOMINICAN REPUBLIC: On bark, Santo Domingo, Rose,

March 1913 (MICH 8238); in wooded ravines along trail to

Goodrich rubber grove, vicinity of Piedra Blanca, Prov. of

La Vega: altitude 200—500 meters, Allard 17142, 21

November 1947 (US). HAITI: Mossy tree in deep shade,

ravine on east Slope above road NW of Foret des Pins,

Dept. l'Ouest, Imshagg_22909, 16 July 1958 (MSC). JAMAICA:

Fern gully, near Ocho Rios, 700 ft., St. Ann Parish,

124

M 15797 a, 6 May 1953 (MSC). PUERTO RICO: On twigs,
IL'Ei. Romana, Rio Dulce, Johnston, 7 April 1913 (MICH 8269);

:EBZI. YUnque, Chamberlain, February 1941 (M0 158071).

 

PHILIPPINES: On bark, Lake Manguao, Palawan,

INIKEerrill 9014, April 1913 (US).

Coenogonium Leprieurii Nyl.

Jerrin. Sci. Nat. IV. 16:89. 1862. Holocoenis Leprieurii

 

Clements, Gen. Fung. 174. 1909. g. Linkii var. Leprieurii

 

hdx>nt. Ann. Sci. Nat. III. 16:47. 1851. Orig. coll.:
F rench Guiana, Leprieur .

Synonyms: Dictyonema sericeum var. camerunense Henn.,

 

llitteil. von Forschungsreisend. ind Gelehrten aus dem
IDeutsch. Schutzgebeit. 11:144. 1889.

.9. Leprieurii var. lamellifera Wain. Ann. Acad. Sci.

 

‘Fenn. 15(Ser. A). 6:157. 1921.

.Q- Leprieurii var. panniforme Wain. Jour. Bot.

 

34:204. 1896.

Thallus spongy-byssoid, thin, circular, extending
out horizontally in a prostrate growth, becoming imbricate
With one margin affixed to substrate, shelf—like, yellow—

green: algal symbionts Trentepohlia umbrina or T. odorata.

 

Apothecia situated on both upper and lower surfaces of thallus,

125

1—‘—-ES mm. in diameter, round, sessile or rarely sub-sessile:

d isk flat or somewhat convex, carneous or pale yellow; margin

‘fi7lfitite, thin, uneven: excipulum euparaplectenchymatous:

lfiljgrpothecium colorless, partly paraplectenchymatous: hymenium

about 60 LJ. high; paraphyses Slender, 0.7 — 2.5 LL wide,
£3;j_mp1e, rarely septate, apices capitate, clavate to sub-
§321.0bose, 1.7 - 5 u wide; asci cylindrical or cylindric-clavate,
41-..7 - 6.5 u wide, 8—spored; Spores simple, uniseriate, linear,

c>1310ng—elliptic or fusiform—elliptic, 5.5 — 10 x (1) 2—4 u.

Montagne established 9, Linkii var. Leprieurii on

 

‘tdne basis of the apothecium being plane from the beginning,
{Provided with a rudiment of a stalk, remaining as such or

(Seviating to convex shape, and maintaining hardly a trace of

a.nargin. He also distinguished this variety from g. Linkii

<Dn the color of the apothecium. Montagne, however, made no

Inention about the septation of the spores. Santesson (1952)

Considers the color characteristics as not reliable since

the apothecia of probably all species of Coenogonium get

 

the same color and appearance if they are repeatedly moistened

and dried. Tuckerman (1882) also believed that this

variety cannot be kept apart by the given color differences.
Nylander distinguished g. Leprieurii from Q. Linkii

on the simple spores, the more slender filaments of the

126

tZ-‘l'lallus and of the paraphyses.

Q. Leprieurii var. lamellifera was established by
‘WTainio on the basis of the wide, lamellate thallus. This
<:=haracter is not considered taxonomically Significant.

Wainio established 9. Leprieurii var. panniforme on

 

 

1:he size of the spores, being 1.2—1.4 u wide. This may be a
gyrowth form of Q. Leprieurii and probably belongs to this
species.

I found this species quite constant in the size of
its spores since specimens from Asia, South America, Central
.America, West Indies and the single fertile collection from
West Africa all had the same spore sizes.

Exsicatti: .Mglmg, Austr° Amer. n. 114.

Material seen: COSTA RICA: On tree, Cerro de la
Carpintera, Prov. of Cartago: altitude 1500-1850 meters,
Standley 35596, February 1924 (US); on tree, El Silencio,
Prov. de Guanacaste, near Tilaran, altitude about 750
meters, Standley & Valerio 44619, 13 January 1926 (US);
west bad< of Rio Siquirres, 70-200 meters, Limon Prov.,
Dodge, Catt & Thomas 5574, 7 December 1929 (MO 70721);
flood-plain of Rio Sandalo, peninsula of Osa, 1-10 m.,
Puntarenas Prov., Dgggg & Groeger 9925, 21 August 1936

(MO 154706); flood-plain of Rio Sandalo, peninsula of Osa,

127

3—“-'10 meters, Puntarenas Prov., Dodge & Groeger 10051, 23

 

August 1936 (M0 154707); on twig, flood-plain of Rio Sandalo,
I;><eninsula of Osa, 1-10 meters, Puntarenas Prov., Dodge &
.jgfigroeger 10163, 25 August 1936 (M0 154708); on bark of trees,

<:=,anas GordaS, altitude 1100 m., Quiros 1338, 17 November 1945

 

(HMO 160735); on bark of trees, Penshurst (Limon), M. Q. g.

:1.490, 13 October 1948 (M0 166111). GUATEMALA: Near the

IFFinca Sepacuite, Alta Vera Paz,.gggk & Griggs, 8 April 1902
( US); on twigs, common on jungle, Vaxactum, Bartlett, 20
Idarch 1931 (MICH 15153); on twigs, Monte Santa Teresa,
iLundell, 12 April 1933 (MICH 2721); on twigs, wet forest,

Escoba, across the bay (west) from Puerto Barrios, Dept.

 

Izabal, near sea level, Standley 72905, 3 May 1939 (M0 166110).
HONDURAS: On trunk of palm, Valentin, E1 Cayo

District, Mgigg, 23 June 1936 (MICH 3556). MEXICO: Belleville,

Oaxaca, Orcutt 2934, 23 February 1910 (US); on evergreen

trees in deep jungle, virgin upland evergreen jungle, elev.

920 m., El Suspiro, 10 km. north of Berriozabal, State of

Chiapas, Hale & Sodertrom 20208, 22 March 1960 (US).

 

PANAMA: On tree, common, Hills north of Frijoles, Canal Zone,
Standley 27531, 19 December 1923 (US); on twig Barro Colorado
Island, Canal Zone, Shattuck, 1931 (MO 68651); abandoned

cocoa plantation, Las Cascadas near summit, Canal Zone,

128

.IEESQQgg & Hunter 8841, 23 December 1934 (US); forest along
*Zmalephone trail between the Rio Indio Hydrographic Station

,Eand the Natural Bridge of R. Puente, Canal Zone, Dodge &
gallon 8910, 13 January 1935 (M0 154704); Prov. Coclé:

131 Valle de Anton, altitude 600 m., Martin 2937, 24 July

 

21935 (MO 73207); Las Cruces, Canal Zone, Martin 3073,

 

231 July 1935 (MO 73209): Barro Colorado Island, Canal Zone,
Iflartin 3115, 31 July 1935 (MO 73210); Barro Colorado
iIsland, altitude 20-75 m., Martin 4135, 11 August 1937 (MO
73969); on bark, Barro Colorado Island, Weston, (no date),
(MICH); on tree bark, Barro Colorado Island, Prescott (g),
25 December 1962 (MSC).

BRAZIL: On bark of tree, Rio de Janeiro: Corcovado,
Malme 331 a, 15 August 1892 (MSC 152231); on bark of trees,
Rio Grande do Sul, pr. Sao Leopoldo, Malme 114, 18-26
October 1892 (MSC 151832); on tree trunk, Amazon Basin,
altitude 2510 ft., Rusby 752, 28 September 1921 (NY);
on twig, Rio Grande do Sul, Pareci Novo, giggilg (155),
3 December 1948 (M0 166104); 8. Gabriel, Amazon & Andini,
Spruce 228, (no date) (NY). COLOMBIA: On dry twigs, La
Sierra, Dept. Antioquia: Medellin; altitude about 200 m.,
Archer 1335, 3 January 1931 (US); on tree, E1 Choco: La

Concepcion, Archer 2009 a, 30 April 1931 (US). PERU:

129
(:>II tree trunk, East of Quimiri Bridge, dense forest, Dept.

CIWJnin: La Merced; altitude about 700 meters, Killip &

 

Smith 24039,. May 29-June 4, 1929 (US); on bark, in jungle
(seast of Tingo Maria, Dept. San Martin: altitude 625-1100
130., Allard 20797 b, 30 October 1949-19 February 1950 (US).
ESURINAM: (no locality), Weigelt, 1827 (MO 25591); Tafelberg
(Table Mountain), Maguire_24598 a (B), 1 September 1944 (NY).
“JENEZUELA: (No locality), Fendler, May 1856 (NY); on
‘tree trunks in humid forest, Colonia Tovar & vicinity,

:in meadows & forest: altitude 1700—2300 meters, State

(bf Aragua, Pittier 10033,.28 December 1921 (US); Colonia
Tovar, altitude 1800—2000 m., Allart 349, December 1924

(NY).

BAHAMAS: On alga, Rifle Range Coppice, New
Providence, Britton 564, 6 September 1904 (NY); on wood,
Cherokee, Abaco, Brace 1936 & iggz, 30 December 1904 (NY):
CUBA: On rocks, Valley of the Rio Bayamita, south slope of

the Sierra Maestra; altitude 900-1050 meters, Maxon 3975,

 

5-7 April 1907 (US): on tree trunk, forest slope,

Santa Ana, about 6 miles north of Jaguey, Yateras, Oriente:
altitude 600-625 m., Maxon 4194, 25 April 1907 (US); on

rock, Loma del Gato, around summit of San Juan in mossy cloud

forest with huge boulders and cliffs, Sierra Maestra,

\

130

<2’11‘iente, Imshaug 24695, 12 August 1959 (MSC). DOMINICAN

IE§JEPUBLIC: On bark, in ravine near Goodrich rubber groves,

"I’icinity of Piedra Blanca, Province of La Vega: altitude

32:00—500 meters, Allard 16915, 16922 & 16962 a, 12 November

£1.947 (US). GRENADA: Montane thicket, Summit of Mt. Sinai,

2100 ft., St. David Parish, Imshaug 16180, 13 July 1953 (MSC);
<:>n bark, secondary forest, Mt. Sinai Waterworks, 1300 ft.,

SSt. David Parish, Imshaug 16261, 21 July 1953 (MSC).

IiAITI: On base of shrub in thicket, vicinity of Dondon, Dept.

(in Nord, altitude about 400 meters, Leonard 8736 a, 8
~January 1926 (US); base of tree in dense thicket, Pailborean
Pass, vicinity of Ennery, Dept. de 1'Artibonite, altitude

325-900 meters, Leonard 9552, 5 February 1926 (NY); on twig,

elfin woodland forest, ridge north of Foret des Pins

(Shada Station), 5800 ft., near Dominican border, Dept. de

l'Ouest, Imshaug 22752 & 22720, 14 July 1953 (MSC); on

twig, elfin woodland, ridge about Foret des Pins, 5800-

5900 ft., Dept. de l'Ouest, Wetmore 3178, 18 July 1958 (MSC).

JAMAICA: On slopes above Tweedside, altitude 2000—3000 ft.,

Maxon 969 a, 11 April 1903 (US): on tree, along the trail

from Morce's Gap, 5000 ft., to Vinegar Hill (3950 ft.)

 

Maxon 1517, 23 April 1903 (US); on log, vicinity of Hollymount,

Mount Diablo, Maxon 1903, 8-9 May 1903 (US); exposed roots of

131

forest tree, deep ravine in mountain forest above House

Hill, St. Thomas: altitude 500-700 meters,

Maxon 9117,
€5-—12 June 1926 (NY & US); Lumsden, St. Ann's, Orcutt 6088,

 

1928 (US); on twig, Hollymount, Mt. Diablo, 2754 ft.,

S 1: . Catherine Parish,

Imshaug 14227, 28 January 1953 (MSC).
P UERTO RICO :

Quebrado de Limones, Mesa Mt., vicinity of
Mayaguez,. Britton & Marble, 4—10 March 1906 (NY); vicinity
of Utuado, Britton & Cowell 53, 15-20 March 1906 (NY); ravine
near Utuado,‘ Wheeler 991, 15—20 March 1906' (NY & MICH 13295);

Finca Alvarez, Quebradillas, Bro. Hioram 5;, January 1913
(NY); on tree, mountain forest, 905 m. altitude, Barrio

des Maizales, Sierra de Naguabo, Britton & Hess 2299, 9
March 1914 (NY):

 

Rio Icaco & adjacent hills, altitude 465-

720 m., Sierra de Naguabo, Shafer 3759, 30 July - 5 August

1914 (NY); on twig, Rio de Maricao, 600—720 m. altitude,

Britton & Cowell 4234, 14 February 1915 (NY & MICH 14433):

on wood & trees, Yunque, Fink 781, 11 December 1915 (MICH 8558);

on trees in forest, Chamberlain, April 1941 (MICH).

BORNEO: On twig, Gat,

Upper Rej ang River,

Sarawak,
3L

& M. g. Clemens 5901, 15 July 1929 (NY). CEYLON:

(no locality), Thwaites g_34, (no date) (NY).

CHINA: On
nmssy slope, 480 m. altitude, Ta Chai Shan (Yung Hsien),

Kwangshi, Cheo 2622, 26 August 1933 (NY). PHILIPPINES: On

132
thvigs of living Shrubs, Lamao, Province of Bataan, Merrill

35568, October 1903 (US); Apo, 6500 ft., (collector not

 

'}<nown), 2676, 1 April 1905 (NY). WEST AFRICA: Prince's

IIsland, Marm, 1861 (NY).

Coenogonium Linkii Ehrenb.

in Nees ab Esenb. Horae Phys. Berol. 120, P1. 27. 8120. —

(Orig. coll.: Santa Catharina, Brazil, Chamisso.

Synonyms: ‘9. andinum Karst. in Nyl. Bot. Zeit.
20:178. 1862.

.Q- confervoides Nyl. Flora 41:380. 1858. —
IOrig. coll.: Nbxico, Er. Mfiller (Schimper Herb.).

Rejected name: ‘g. controversum Pers. in Gaudich.
‘Voyage Uranie, Bot. 7:214. 1826. Nomen novum for Q. Linkii
and Conferva arborum Ag. — Orig. coll. of g. arborum: Maria-
nas Island, Gaudichaud.

Thallus shelf—like, pendant, lamellate, spongio-
byssoid, orbicular, occasionally dimidiate, attached at
one side to the substratum in the manner of some polypores:
reniform or round, Sometimes zonate, yellow—green; algal
symbionts Trentepohlia arborum, T. ggrgg or T. elongata,
covered by hyphae 2—3 u wide. Apothecia on different levels

on the thallus, either on the upper or lower surface, 0.6 -

133

1.2 mm. in diameter, round: young apothecia white, nearly
,pedicellate, top or cup-shaped, gradually widening and be—
c:oming subsessile: disk flat or convex, carneous or pale—
yrellow becoming red; margin pale, thin, entire; excipulum
eeuparaplectenchymatous; hypothecium white or colorless,
thin; hymenium gelatinous, about 40-60 N high; paraphyses
septate, 1-3 u wide, apices capitate, sub-globose to
clavate, 2.3 - 5 u wide: asci cylindrical to fusiform-cylindric,
about 4-7 u wide, 8—spored; spores one-septate, biseriate,
straight to oblique or slightly curved, ovate to short-
fusiform, 6.8 - 12.5 x 2—4 u.

Ehrenberg illustrated the Spores of this species
as Simple; Santesson believes that this might be incorrect
on account of the poor optical equipment used by Ehrenberg.
Nylander (1862) described this species as similar to g.
Leprieurii except for the septate Spores and the wider
filaments and paraphyses of the former. He also mentioned
simple spores with only poorly developed septa apparent.
I found this character occurring in some of the materials
examined and unless the very thin septa were carefully stained,

the two—celled nature of the spores was likely to be over-

looked.

134

Coenogonium andinum was separated from Q. Linkii

 

on the basis of its wider algal filaments and paraphyses.
Pas this character does not seem sufficient to establish
aanother species, the writer believes that g, Engiggm should
Tbe a Synonym of g, Linkii.

Nylander distinguished Q. confervoides from g, Linkii
solely on the wider algal filaments and more distinct confer-
void cells of the former. Since algal characters cannot be
used in maintaining a separate species, g. confervoides is
treated here as a synonym of Q. Linkii.

The spore sizes of this Species were observed to be
quite variable in the collections studied and a comparison
can be made of these variations among the geographical areas
represented. Below is a summary of the range in length and
width of the spores:

United States - 7—9.3 x 2-3 0

South America - 6.8—10.4 x 2—3.5 u

Central America - 7.5—12 x 2—3.5 0

West Indies — 12.5 x 2.5-4 0

Philippines — 8.7-9.3 x 2.3-3.4 u

Australia - 11.9 x 4 u

Exsicatti: Malme, Austr. Amer. no. 113.

135

Material seen: FLORIDA: On bark of Carpinus and
.Magnolia, on moss, 6 mile Cr., Calkins 8;, 29 &.ll§a (no
aates) (MO 33775 & MICH 962 a, US); on bark of C_a_r_\La
Sgomentosg, Millhopper Hammock, in limestone sink, 8 miles
tnorthwest of Gainesville, Alachua Co., Uyenco 11, 6 February
1963 (MSC); on bark of sweet gum tree, Black Bottom
Hammock, 8 mi. west of Sanford, Seminole Co., gyggggpgg,

9 February 1963 (MSC); on bark of oak among liverworts, Low
Hammock near Sanford, Seminole Co., Uyenco gg, 9 February
1963 (MSC); on twigs of sweet gum tree, Lieder Trail,

Highlands Hammock State Park, Highlands Co., Uyenco_ZZ,

11 Feb. 1963 (MSC): on twigs and bark of Lyssoloma,

 

Rattlesnake Hammock, Everglades National Park, Dade Co.,

Uyenco 107, 13 February 1963 (MSC); on bark of Ficus tree,

 

Redd Hammock, Everglades National Park, Dade Co., Uyenco 116,

 

14 February 1963; on bark of Dipholis salicifolia, Fire Road

 

near Gate 3, Redd HammoCk, Long Pine Key, Everglades National

Park, Dade Co., Uyenco 117, 14 February 1963 (MSC).

 

LOUISIANA: Caris Creek, near St. Francesville, West Feliciana
Parish, Flint, 15 July 1938 (Prescott).

COSTA RICA: On twigs and tree, E1 Muneco, on the
Rio Navarro, Prov. Cartago, altitude 1400—1500 meters,

Standley & Valerio 51055, 6—7 March 1926 (US). MEXICO:

136

Veracruz, Turpus 222, 1921-22 (US). PANAMA: Marraganti,
fiilliams'logz, 8 April 1908 (1w); on trees, Achajuela,
Egornman 3QQ_, 3 November 1917 (MD 158769).

BRAZIL: On barks of trees, Rio Grande do Sul:

Cascada de Herminegilda pr. Pelotas, Malme 329, 11 December

 

1892 (MSC); on Piptadenia durissima, Porto Alegre, Rio
Grande do Sul,.§gmbgugg, 1931 (MO 71064). COLOMBIA:
Vicinity of Medellin, Rev. Charetier 208, 1911 (US): on
twig, Bosques de Yanaconas, altitude 1760, Agggl, 5
February 1938 (US): Cordillera Oriental, Dept. Norte de
Santander: region del Sarare: Alto de Sta. Ines, bosques

2150—2250 meters, Cuatrecasas, Schultes & Smith 12440,

 

 

19 October 1941. ECUADOR: Camino de los Colorados,
PrescottT-48, 10 February 1958. PARAGUAY: (no locality),
Morong 1441, 1888-1890 (NY). PERU: On Pajnail trail 3500-
5000 ft., Bassler, October 1922 (NY & MO 166292); on tree
trunk, Dept.Ayacucho: Aina, between Huanta and Rio Apurimac,
altitude 750-1000 meters, open woods, Killip & Smith 23146,
7-17 May 1929 (MO 70247 & US); on twigs, Dept. Junin, Pichis
Trail, Enenas, altitude 1600—1900 meters, dense forest,
Killip & Smith 25658, 30 June—2 July 1929 (US). VENEZUELA:
On tree trunks in humid forest, Colonia Tovar & vicinity,

in meadows & forests: altitude 1700—2300 meters, Aragua

137

State, Pittier 10034, 28 December 1921 (US).
CUBA: (No locality), Wright, (no date) (US); on
‘twig, south of lumber camp, crest of Sierra Nipe, altitude

600-700 meters, Oriente, Morton & Acuna 3240, 16—17 October

 

1941(US): on bark, near Topes de Collantes, Trinidad Mts.,
Las Villas, Imshaug 24628, 2 August l959 (MSC).

DOMINICAN REPUBLIC: On twig, cloud forest, ridge above
Los Amaceyes, 300—3200 ft., Cordillera Septentrional,
Imshaug 23276, 2 August 1958 (MSC): on twig, wet montane
mist forest, Casabito, 4000 ft., between El Rio and
Jayaco, La VEga, Wetmore 3433, 4 August 1958 (MSC).

HAITI: On bark, elfin woodland forest, ridge north of
Foret des Pins (Shada Station), 5800 ft., near Dominican
border, Dept. de l'Ouest, Imshaug 22753, 14 July 1953
(MSC): on tree branch, shrub thicket at summit of Montagne

Noire, 5500 ft., Kenscoff, Dept. de L'Ouest, Wetmore 2774,

 

7 July 1958 (MSC). PUERTO RICO: Mt. Morales, near Utuado,
upper slopes of tree, Howe 1153, 19 March 1906 (NY & MICH
13296): (no locality), g. g. g. _2_8_§_fl_, 15 July 1913 (MICH
7042); on twigs, Rio Icaco & adjacent hills, 465-720 meters,
Shafer 3736, 30 July - 5 August 1914 (NY & MICH 13422):

near La Juanita, Britton 905, 7 February 1915 (NY): on tree

in mountain forest, Monte Cerrote, near Adjuntas, 900-1050

138

meters altitude, Britton & Brown 5466, 15 March 1915 (NY &
MICH 14434); on Acrista, Monte Torrecilla, 900-1100 meters
altitude, Britton, Cowell & Brown 5666, 19e20 March 1915
(MICH 14435). TRINIDAD: On a small tree, forest, Broadway,
February 1928 (M0 943076).

PHILIPPINES: Bontoc Subprovince, Luzon, §£-.Y§22‘
verbergh 874, October 1910 (US): on trees, Baguio, (no
collector's name),-1§1§, 20 September 1904 (US).

AUSTRALIA: Tasmania, Lindley, (no date) (NY).

Coenogonium moniliforme Tuck.

Proc. Am. Acad. Sci. 5:416. 1862. — Orig. coll.:
Cuba, (unknown locality), corticolous, Wright n. ;_g,

Synonym: Biatorinopsis torulosa Mfill. Arg. Rev.
Myc. 10:114. 1888. - Orig. coll.: Guarapi, Paraguay,
corticolous, Balansa 4165.

Thallus effused, crust-like, margin widely pallid,

glaucous-green to deep yellow; algal symbiont Physolinum

 

monilfi;(De Wildem.) Printz, filaments branched, cells sub-
globose to elliptic, about 18 u in diameter, elliptic cells
10-15 x 22-31.5 u. Apothecia appressed on the thallus,
round, 0.2 - 0.4 mm. in diameter: disk flat, carneous—red,

turning paler when wet with water: margin white, thin,

139

eentire: excipulum and hypothecium colorless; hymenium 50-60
0 high; paraphyses simple, 1.2—2 u wide, apices slightly
capitate, sub—globose to clavate, 2-4.5 u wide; asci sub-
cylindric to clavate, 8-spored; spores one-septate, uni-

to biseriate, oblong— elliptic to fusiform-elliptic, 9-13 x
3.2 - 5 u.

Santesson (1952) reports this species as facul-
tatively foliicolous. Most of the materials I examined had
larger spore sizes than those mentioned by Nylander in his
re-description of this species (1862). No variation in the
range of spore sizes was observed in collections from
Central America, United States and the West Indies. The
original descriptions made by Mfiller and Tuckerman did not
mention the size of the spores except to state that they
were 3-4 times longer than wide. Malme (1937) gave the
measurements of the spores in the specimens he examined from
Paraguay as 9—11 x 3—3.5 u. Knight (1886) and Shirley (1889)

mentioned this species as closely allied to g, botryosum

 

on account of the elliptic cells found in their short

filaments. The algal filaments are Trentepohlia unlike

 

those of g, moniliforme which are Physolinum monile.

 

 

The spores of g, botryosum are uniseriate and 10.4 x 3 u

 

in size, while those of g. moniliforme are irregularly

 

140

biseriate and relatively larger, 9—13 x 3.2 - 5 u.
Exsicatti: Malme, Austr. Amer. 111 & 112.

Material seen: FLORIDA: On trees, (no locality),

 

Austin, (no date) (MSC 13140): Pine Island, Charlotte
Harbor, Charlotte Co., g. Q. S. & C. 5., March 1878 (NY);

on bark, Caloosa River, Polk Co., Austin, 1878 (US):

 

southwest Florida, Austin 42, March 1878 (US): on bark of

Ficus, Fuch's Hammock, Homestead, Dade Co., Uyenco 100,

 

12 February 1963 (MSC).
GUATEMALA: On dead branch, Dept. Chimaltenango,
along Rio Guacalate, southeast of Chimaltenango, altitude

about 1700 m., Standley 80054, 14—23 December 1940 (M0 166105).

 

PARAGUAY: On bark, Aregua pr. Asuncion, (Exp.
Regn. I. Lich. n. 1626),21 August 1893, Mglmg, (no date)
(MSU 151829). SURINAM: On bark, (no locality, collector
not known, no date), (Willey Lichen Herbarium) (US).

CUBA: (no locality), Wright, (no date) (US).
HAITI: elfin woodland forest, north of Foret des Pins
(Shada Station), near Dominican border, 5800 ft. (Dept. de

l'Ouest), Wetmore 2878, 14 July 1958 (MSC). JAMAICA: On

 

trees, (no locality), Harris, (no date) (MICH 12378); on
stems of Pisonea aculeata, Hope Gardens, 680 ft. alt.,

Harris, 14 January 1915 (NY).

141

Coenogonium ornatum Mfill. Arg.

 

Bull. Herb. Boissier. 4:96. 1896. - Orig. coll.:
On bark, Queensland, Australia, Shirley n,;1§1.

Thallus amorphous, filaments closely aggregated,
not at all fasciculate, bright-green: algal symbiont

Trentepohlia, cells of filaments 2—3 times longer then

 

the width. Apothecia 0.4—0.6 mm. in diameter, round, sessile;
disk flat to convex, carneous-purple; margin concolorous with
disk, wearing away at maturity: paraphyses simple, 1.5 u
wide, apices clavate, sub-truncate, 1.5 - 2 u wide; asci
clavate, 8-spored; spores simple, uniseriate, fusiform,
8—10 x 2.3 — 4 u.

Nylander (1862) stated that this species was closely

related to C, interpositum. It is believed that the

 

distinct immarginate structure of the apothecium, the
arrangement of the spores in the asci and the texture of
the thallus merit the separation of this species from the
other members of the group.

I found the spores in the materials from the
West Indies of the same length but slightly narrower than
those from South America. Specimens examined from Central

America had longer Spores than those from either of the two

142

but the widths were identical with the specimens from the
West Indies, 2.5 u.
Exsicatti seen: Malme, Austr. Amer. no. 113.

Material seen: MEXICO: Veracruz: Mirador, Baldwin

 

Jr. 14406, 20 May 1950 (vs).

 

PARAGUAY: On bark of Acacia, 9 August 1893,
(Exp. Regn. I. Lich. n. 1552), Malme 112 (MSC 151830).
VENEZUELA: On tree trunks, Colonia Tovar, 2000—2100

m., Pittier 10033, 28 December 1921 (NY).

 

JAMAICA: (no locality), Cummings 170, February

 

and March, 1905 (NY). PUERTO RICO: (no locality), Britton,

(no date) (MICH 11758 b).

Coenogonium pannosum Mfill. Arg.

Flora 64:234. 1881. - Orig. coll.: On bark, near

Apiahy, Central Brazil, Puiggari 1026.

 

Thallus adnate, slightly tomentose, upper surface

appearing wrinkled, yellow-green: algal symbiont Trentepohila,

 

cells barrel—shaped. Apothecia 0.3 - 0.5 mm. in diameter,
round, sub-sessile: disk flat, rarely convex, pale to whitish-
orange: margin paler, thin, entire: paraphyses slender, 1.5 u
wide, apices obovoid, 2.7 - 3.5 u wide: asci cylindrical,
8—spored: spores one-septate, uniseriate, fusiform, 6-7 x

1.5 - 2 u.

143

The immarginate structure of the apothecium and
the very narrow spores distinguish this species easily from
the others which have one-septate spores.

Material seen: GUADELOUPE: Souface, Ep. 400 m.,
‘Questel 3616, 20 January 1939 (MD 161488). JAMAICA: East
slope of John Crow Mts. above Ecclesdown, 2300 ft., Portland
Parish, Imshaug 14544 a, 11 February 1953 (MSC). PUERTO
RICO: On tree, Sierra de Naguabo 600 m. altitude, Britton

& Cowell 2194, 8 March 1914 (MICH 13294 & NY).

Coenogonium tomentosum Mfill. Arg.

Jour. Linn. Soc. Lond. 32:206. 1896. - Orig. coll.:

On bark, New Zealand, Colenso 1615.

 

Thallus tufted, tomentose, bright-yellow: algal
symbiont Trentepohlia, filaments short, closely aggregated,
not interwoven cells twice longer than wide. Apothecia
0.3 - 1 mm. in diameter, round, sub-sessile: disk flat
to slightly concave, white when young, becoming carneous-
orange as it matures: margin white, prominent, entire:
parphyses slender, septate, capitate, globose: asci clavate,
8-spored: spores simple, uniseriate, fusiform, 8 - 10.5 x

2.5 u.

144

The original description for this species described
the size of the spores as 10-11 x 2.5 u. The specimen
from the Dominican Republic was observed to have shorter
spores (7.6 u) but the characteristic uniseriate arrangement
of the spores in the ascus and the globose-capitate apex of
the paraphyses place it in this species.

Material seen: BAHAMAS: Soldiers Road, New

 

Providence, Britton 3180, 30 January 1905 (NY): Mt. Pleasant,

 

New Providence, Britton 760, 13 September 1904 (NY).

 

DOMINICAN REPUBLIC: In heavy woods near Goodrich
rubber grove, vicinity of Piedra Blanca, Province of La
Vega, altitude 200-500 meters, Allard 18987, 19 January 1948

(US).

Coenogonium velutinum A. Zahlbr.

 

Nat. Hist. Juan Fernandez 2:332. 1924. — Orig. coll.:
Masatierra, Juan Fernandez, Skottsberg, s.n.
Thallus adnate, cushion—shaped, round, bright-

yellow: algal symbiont Trentepohlia, filaments short,

 

closely aggregated, cells cylindrical. Apothecia 1-2
mm. in diameter, round, biatorine, sessile: disk concave
to slightly convex, orange-yellow, pruinose: margin pale,

whitish, thin, entire: excipulum and hypothecium

145

paraplectenchymatous, the latter pale yellow: hymenium
colorless to slightly yellow near the surface, about 100 u
high: paraphyses slender, 1.3 - 2 u wide, simple, apices not
at all wider: asci cylindrical to clavate, 8—spored: spores
one-septate, irregularly uniseriate, elliptic to fusiform-
elliptic, 9—11 x 2.6 - 3.5 u.

This species is easily distinguished from the others
by the pruinose apotheciaand the almost truncate shape of
the apices of the paraphyses. It is similar to g, disiunctum
except for the uniseriate spores, the slightly smaller size
of the spores and the aseptate paraphyses.

Material seen: JUAN FERNANDEZ: Yunque Mts., 500

 

m., Masatierra, Skottsberg, 24 April 1917 (NY).

 

JAMAICA: South slope of Mossmans Pk., 500 ft.,

Blue Mts., St. Thomas Parish, Imshaug 14616, 19 February

 

1953 (MSC).

DISCUSSION

It is often difficult to establish whether an organism
is a lichen or not. Easily misinterpreted situations occur

frequently in the genus Coenogonium where the algal symbiont

 

is more conspicuous than the mycobiont. In the past, the
algal characters, e.g., shape of cells, cell sizes of the
filaments and the shape of the thallus, have been used to

identify the species of Coenogonium. The problem of taxonomy

 

has been further aggravated wherever slightly or partially

lichenized algae (Trentepohlia or Physolinum) were encountered

 

 

and names were given to the consortium based on the algal
component alone.

Some authors have assigned equal importance to
the alga and fungus in lichen taxonomy: others have given
scientific names to the consortia as a whole. Thus, two
consortia with the same fungus but with different algal
symbionts may have different scientific names. The
International Code of Botanical Nomenclature, Art. 76 (1950)
states,in part: "For nomenclatural purposes names of lichens

shall be taken as applying to the fungal component." Such

146

147

provision should clarify the misinterpretations raised

by taxonomists who maintain that lichens form a taxonomic
unit. As is well known, a consortium is not a genetical

unit and cannot therefore be considered taxomonically as

such.

It has been shown that neither the lichen—fungi
nor the lichen-algae form distinct taxonomic groups but
that they show relationships with many groups of non—
lichenized fungi and algae. The names of the lichen-fungi
have usually been thought of as the names of the lichens.
Lichens, however, are biological entities, not taxonomic
units. The fundamental fact which must be observed in the
taxonomy, therefore, is that the lichen-fungi, not the
lichens, are being classified.

The name Coenogonium has been erroneously used to

 

refer to the members of the Trentepohliaceae which are more
or less surrounded by fungal hyphae. It is evident that
this designation was given for the consortium in which the
taxonomical position and the morphology of the algae have
been the bases for the classification of the genus. Since,
as has been discussed above, the lichen is considered as a
biological group and not a taxonomic unit, the generic name

Coenogonium must, from a strictly taxonomic point of view,

148

belong to the fungal component.

The present investigation is the natural result of
combined herbarium, laboratory and field studies. The
importance of culture studies in this group of lichens
cannot be overemphasized. Here is a situation where the
experimental method has proven useful as a means of solving
a taxonomical problem. It not only supplements the classi—
cal morphological methods in taxonomic studies (herbarium

and field work) but also shows that the genus Coenogonium

 

can be treated in the same manner as the other members of
lichen groups where the fungal characters are the primary
bases for their classification. The results obtained in
the laboratory reveal the role of the fungus in the
symbiotic relationship, i.e., in determining the shape of
the thallus, and therefore, permits its use as a taxonomic
character. It was demonstrated that the alga does not
control the shape and development of the lichen. It has

been shown how a shelf-like Coenogonium, for instance,

 

could be formed by a lichen-fungus producing apothecia that

bear either simple spores (g, Leprieurii) or septate spores

 

(g. Linkii). The expanded, loose growth on a piece of

bark may be a Coenogonium with simple spores (g, interpositum),

 

or one-septate spores (g. interplexum). A cushion—shaped

 

149

thallus with wrinkled or tomentose surface could be that

with simple spores (Q. tomentosum) or one-septate spores

 

(Q. botryosum). Thus, it is unwise to identify the species

 

of Coenogonium in nature or to rely on measurements of algal

 

cells. The size of the algal filaments do not have any
taxonomic importance in so far as shape of the thallus is
concerned. It was demonstrated that the lichenized

Trentepohlia does not branch freely and its cells are generally

 

smaller in size than those of the free—living algae. If the
extent of the lichenization of the filaments has an evident
influence on the size of the algal cells and the branching
of the filaments, it is apparent, therefore, that the fungus
has some inhibitory effect on its symbiont. This could.
account for the different types of thalli in the genus.

The results of the culture studies further show that the
reproductive characteristics of the lichen-fungus are of
real importance for the species taxonomy of Coenogonium,

the algal characters not being reliable as such. The
biological characteristics of the fungus, the special sub-
strata and geographical distribution do not seem to be
equally important in establishing the specificity require—
ments of the lichen—fungus as to its algal symbiont. The

morphological characteristics are believed to be more

150

important in taxonomy.

All the specific epithets given to sterile or

 

undeveloped Coenogonium species have been rejected in the
present study. It is believed that a partially lichenized

Trentepohlia with only a sterile lichen-fungus, should not

 

be named as a Coenogonium but rather, the Species of
Trentepohlia should be identified. It is suggested that
material with sterile lichen-fungi can be conveniently dis—
posed of by simply indicating them as "slightly or heavily
lichenized Trentepohlia."

Zahlbruckner (1926) based his classification system
of orders and suborders on the fungal characters, but in
the case of families, the algal characters have been
the primary basis. The genus Coenogonium provides a
case where the lichenized fungi do not show distinct
specialization for individual species of Trentepohlia.

The thalli of Coenogonium vary considerably in
size according to their age and location. Specimens
with mature apothecia have been found to vary from 10—25
mm. and this range was observed in several species. At
what age of the thallus the apothecia first appear is un-
certain. The thalli grow and persist for several years,

and whether or not the fungus fruiting bodies appear annually

151

or at certain seasons of the year, may provide an interesting
study for future investigators. The collection data reveal
that fertile specimens were encountered at all months of the
year for Coenogonium Leprieurii and at scattered periods in
other species. This information, of course, depends on

the intensity of the collection and, therefore, cannot be
generalized.

The algae have been demonstrated to be capable of
growing apart from the fungus. Whether they can produce a
concentric or any specific shape of thallus if maintained
at the suitable conditions of growth required by them over
a considerable length of time remains to be studied.

The nature of the relationship between the lichen—
algae and the fungal hyphae and the penetration of the
hyphae among the algal cells have yet to be established.
Haustoria have not been observed in the material examined
and apparently the fungus has no effect other than the partial
inhibition of branching of the alga. Haustoria in lichens
are usually not discernible except by electron microscopy.

It is hoped that this comparative study of the
morphology, including the reporductive structures, of
both the algal and fungal components of the lichen-genus

loosely referred to as Coenogonium, may provide a basis for

 

152

future work towards the completion of a monograph of this

genus.

SUMMARY

One species of Trentepohlia can form lichen thalli with

 

more than one fungus species.

One fungus (Coenogonium) can form lichen thalli with

 

more than one species of Trentepohlia.

 

The shelf-like form of lichen thallus is a character—

istic of the fungal component since both the shelf—form

and the non-shelf form of thalli have identical algal
species.

Characters of algal cells, such as, size measurements

and shape, cannot be accepted as valid in the determination

of the species of Coenogonium.

 

Six species of Trentepohlia are recognized for the first

 

time as phycobionts in the lichen-genus Coenogonium.

 

Four species of Trentepohlia: namely, T. abietina, I.

 

 

arborum, T. aurea, and T. elongata are being reported

 

for the first time from Florida.
Physolinum monilggoccurs as the phycobiont of g,

moniliforme.

 

153

154
Of the original forty-eight species referred to as

Coenogonium by earlier authors, eighteen species have

 

 

been recommended for transfer to the genus Trentepohlia

as valid algal species or as synonyms. Four species are
considered as belonging to the Fungi Imperfecti and two
species are treated as synonyms of Coenogonium Linkii.

Eight species are considered doubtful and one species

is rejected as having been based on two discordant elements.
Fifteen species are recongized and accepted as valid

and a taxonomic key has been made for these.

Acharius,

Agardh, C. A.

Ahmadjian, V.

E.

Bold, H. C.

E.

Bornet,
Brand, F.

 

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LIST OF DISTRIBUTION MAPS

Distribution of known Coenogonium species
Black circles represent material seen and examined:

light circles,

 

type localities, no material Seen

acrocephalum Mfill. Arg. (dark circles)

 

botryosum Knight (light circles)

 

(dark squares)

 

(light triangle)

 

pannosum Mfill. Arg. (light squares)

 

tomentosum Mfill. Arg. (dark triangles)

 

(dark squares)

 

(dark triangles)

 

Coenogonium

Coenogonium

Coenogonium complexum Nyl.
Coenogonium curvulum A. Zahlbr.
Coenogonium

Coenogonium

Coenogonium disjunctum Nyl.
Coenogonium implexum Nyl.
Coenogonium moniliforme Tuck.

(dark circles)

 

Coenggonium
Coenogonium

interplexum Nyl. (dark circles)
ornatum Mfill. Arg. (dark triangles)

 

Coenogonium

interpositum Nyl. (dark triangles)

 

Coenogonium

 

Leprieurii Nyl. (dark circles)

 

Coenogonium

velutinum A. Zahlbr. (dark circles)

 

Coenogonium

Linkii Ehrenb. (black squares)

 

170

171

Distribution map of known Coenogonium species. Black
circles represent material seen and examined: light
circles, type localities, no material seen.

 

Dd y all. llll III.
111.611.1111.: gilt

151“] . g UM‘KOmu—lum figs

 

 

 

 

 

Distribution map of:

Coenogonium
Coenoggnium
Coenogonium
Coenogonium
Coenogonium
Coenogonium

172

acrocephalum Mfill. Arg. (dark circles):
botryosum Knight (light circles):
complexum Nyl. (dark squares):
curvulum A. Zahlbr. (light triangle):
pannosum Mfill. Arg. (light squares):
tomentosum Mfill. Arg. (dark triangles).

"l‘lts
{I’ll-3‘11)!

 

 

 

 

ill-
ell-.111} ggaag
Ill-5:.- 3\..I. 2 E o. i a. 111 on a. on. no. I.

. ~ I . i
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173

Distribution map of:

Coenogonium disjunctum Nyl. (dark squares):
Coenogonium implexum Nyl. (dark triangles): and
Coenogonium moniliforme Tuck.(dark circles).

 

 

 

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174

Distribution map of:

Coenogonium interplexum Nyl. (dark circles): and
Coenogonium ornatum Mfill. Arg. (dark triangles).

 

 

 

 

 

 

 

GOODE'S SERIES OF BASE MAPS

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175

Distribution maps of:

Coenogonium interpositum Nyl. (dark triangles) and
Coenogonium Leprieurii Nyl. (dark circles.

 

 

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176

Distribution maps of:

Coenogonium velutinum A. Zahlbr. (dark circles): and
Coenogonium Linkii Ehrenb. (black squares).

 

 

 

 

 

 

 

 

 

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177

Plate 1
Fig. 1. Convex, stalked apothecium with thin margin.
(Coenogonium Leprieurii Nyl.)

Fig. 2. Convex, sub-sessile apothecium (g. interplexum
Nyl. or Q. interpositum Nyl.)

 

 

Fig. 3. Flat, stalked apothecium with thick margin.
(9, disjunctum Nyl.)

Fig. 4. Convex apothecium with thin margin.
(9, Linkii Ehrenb. or Q. pannosum Mfill. Arg.)

 

Fig. 5. Flat, sessile apothecium with thick margin.
(9. implexum Nyl. or g, acrocephalum Mfill. Arg.)

 

 

Fig. 6. Concave, subsessile apothecium. (Q. complexum Nyl.)

 

Fig. 7. Longitudinal section of a mature apothecium
showing the hymenium, excipulum and the
hypothecium (g, Linkii). x 100.

  
 
      

     

   

2
4
5 <5
h m
”mm ‘ Y
MM “HQ/M‘IIIIJ/ /fl/,.
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Fig. 1.
Fig 2.

Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.

178

Plate 2

Ascus and paraphysis of Q. Leprieurii Nyl. with
simple, ovate and linear spores: paraphysis septate
with clavate apex. x 1000

 

Ascus and paraphysis of g. velutinum. A. Zahlbr.

 

with fusiform-elliptic and irregularly biseriate
spores: paraphysis simple with apex not wider than
main filament. x 1200

Ascus and paraphysis of g. moniliforme Tuck. With
1-septate, fusiform and biseriate spores: paraphysis
septate, with globose-clavate and capitate apex.

x 875

 

Ascus and paraphysis of g. implexum Nyl. with 1-
septate, elliptic, oblique and uniseriate spores:
paraphysis septate, apex globose and capitate.

x 1125

 

Ascus and paraphysis of g, Linkii Ehrenberg with
l-septate, short-fusiform and biseriate spores:
paraphysis septate, apex sub-globose. x 889

Ascus and paraphysis of Q. curvulum A. Zahlbr.
showing arcuate spores. x 1500

 

Conidia and conidiophores. x 1000

 

_ J F J

 

 

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QRDW 4

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Fig.

Fig.

Fig.

Fig.

Fig.

179

Plate 3

Portion of a filament of T, arborum (Ag.) Hariot
with three sporangia attached to stalk cells. x 600

Trentepohlia abietina (Flot.) Hansgirg phycobiont
of g. interplexum, showing lateral and terminal
sporangia. x 300

 

 

Trentepohlia aurea (Linn.) Mart., phycobiont of
.g. Linkii, showing young, globose gametangia. x 706

 

Portion of a filament of Trentepohlia umbrina
(Kfitz.) Born. showing lateral, globose sporangia.
x 900

.2. aurea, phycobiont of g, interplexum, with terminal

 

stalk cell bearing sporangium. x 590

“T. aurea, phycobiont of Q. Linkii, showing right-
angled branching habit of the filament and the
pectose cap on the tip cell. x 756

180

Plate 4

Fig. l. Portion of a filament of T. elongata, symbiont of
g, interplexum, showing a young branch and lateral,
globose sporangia at different age stages. x 1064

 

 

Fig. 2 .T. abietina (Flot.) Hansgirg, with terminal, globose
sporangia. x 300

 

Fig. 3. _T. elongata, phycobiont of g, Linkii. x 750

 

Fig. 4. .2. elongata, phycobiont of g. interplexum. x 526

 

 

Fig. 5. Portion of a filament of Physolinum monilg;(De
Wildem.) Printz, showing sporangia containing
aplanospores. x 1000

Fig. 6. Lichenized algal filaments with two apothecial
initials. x 500

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

O +
326

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

10.

181

Plate 5

A portion of a filament of T, arborum showing an
empty gametangium. x 600

Young isogametes. x 1000

A single zoospore. x 778

Isogametes gametes in contact prior to fusion. x 750

Zygote. x 750

Young isogametes of T. aurea. x 1500

Isogametes in contact prior to fusion. x 1125

Zoospores of T. elongata, phycobiont of g. Linkii.
x 1000

 

Portion of a filament of T. aurea showing sessile,
globular gametangium. Note germinating spores in
vicinity of the filament. x 750

Trentepohlia odorata (Wiggers) Wittrock with rough

 

wall and terminal and lateral sporangia. x 900

182

Plate 6

Coenogonium Linkii Ehrenb. showing shelf-like habit of

 

thallus and apothecia on the upper surface. x 2.5
Azael, 5 February 1938 (US).

 

183

Plate 7

Coenogonium Leprieurii Nyl. showing shelf-like habit of
thallus and apothecia on the lower surface. x 2. 2676
1 April 1905 (NY).

 

 

184

Plate 8

Coenogonium Leprieurii showing zonateappearance of thallus.
x 2. Spruce 228 (NY).

 

185

Plate 9

Fig. 1. Trentepohlia aurea (phycobiont of Coenoggnium
interplexum) with young apothecial initial and a
young branch.

 

Fig. 2. .T. aurea with older apothecial initial.

 

Fig. 1.

Fig. 2.

186

Plate 10

Hymenium of Coenogonium Linkii showing asci and
paraphyses with globose apices.

Trentepohlia arborum (phycobiont of Coenogonium

Linkii) with young sporangium.

U:

)4

. 1 ,

 

187

Plate 11

Fig. l. Trentepohlia elongata filaments grown in culture.

Note extended tip cells of filaments (Low power
objective).

Fig. 2. Lichenized filaments of T. elongata showing hyphae
surrounding vegetative cells. (High power objective).