Occasional papers of the Farlow Herbarium of Cryptogamic Botany.

occasional papers of the

Farlow.
HerbariumM of cryptogamic botany

No. 15 April, 1980 Harvard University, Cambridge, Massachusetts

Taxonomic Studies in the Phacidiales: The Genus
Martha A. Sherwood ,
Coccomyces (Rhytismataceae)

Edited by: Donald H. Pfister
Kathryn Rollins

ISSN: 0090-8754

occasional papers of the

Farlow.
HerDarlumM of cryptogamic botany

No. 1. Sylvia A. Earle: Hummbrella, a New Red Alga of Uncertain Taxo-
nomic Position from the Juan Fernandez Islands (June 1969).

No. 2. |. Mackenzie Lamb: Stereocaulon arenarium (Sav.) M. Lamb, a
Hitherto Overlooked Boreal-Arctic Lichen (June 1972).

No. 3. Sylvia A. Earle and Joyce Redemsky Young: Siphonoclathrus, a
New Genus of Chlorophyta (Siphonales: Codiaceae) from Panama
(July 1972).

No. 4. |. Mackenzie Lamb, William A. Weber, H. Martin Jahns, Siegfried
Huneck: Calathaspis, a New Genus of the Lichen Family Cladonia-
ceae (July 1972).

No. 5. I. Mackenzie Lamb: Stereocaulon sterile (Sav.) M. Lamb and Stereo-
caulon groenlandicum (Dahl) M. Lamb, Two More Hiterhto Over-
looked Lichen Species (March 1973).

No. 6. I. Mackenzie Lamb: Further Observations on Verrucaria serpuloides
M. Lamb, the Only Known Permanently Submerged Marine Lichen
(April 1973).

No. 7. Bruce H. Tiffney and Elso S. Barghoorn: The Fossil Record of the
Fungi (June 1974).

No. 8. Donald H. Pfister: The Genus Acervus (Ascomycetes, Pezizales).
|. An Emendation. Il. The Apothecial Ontogeny of Acervus flavidus
with Comments on A. epispartius (May 1975).

No. 9. Donald H. Pfister: A Synopsis of the Genus Pulvinula. A New
Combination in the Genus Gymnomyces. Norton G. Miller: Studies
on North American Quaternary Bryophyte Subfossils. |. A New Moss
Assemblage from the Two Creeks Forest Bed of Wisconsin (July
1976).

No. 10. Emmanuel Sérusiaux: Some Foliicolous Lichens from the Farlow
Herbarium (August 1976).

Continued on back cover

LE ll OS

SE oe | ee ee aries Leo RA ie

TAXONOMIC STUDIES IN THE PHACIDIALES:
THE GENUS COCCOMYCES (RHYTISMATACEAE)

MartHaA A. SHERWOOD

TABLE OF CONTENTS

SE eee ee eds a eg 4K Ok i ye SR ee wm Be ee eS 1
erOCOC Oty Rea &. & 6.0% Bok AX Soe Bees 8 ee ]
PRIStORY eet & coe seca: 8. SS eS ACR Re DO ee OS 2
Relationship of Coccomyces to other genera. ........... 4
Key to the genera of Rhytismataceae with orbicular ascocarps
Whiehropetvoy feet 6 acne ood Aone Ose Bh ee Qe we BOR 6
Materials an MnetNGUS. 4.4%. eae wd De Ode ow SRE SESS 9
MGIDNOlOC§ is + oie a4 ow Ee ERAS Bw A oe Peeters 10
Pista Pano ow 8 ee 3k Be eS OK GO ee 13
COGCOINUCCS Ce ANNs: bos he ee OAS 2S & GOES 1S
Key to-serepted Species: 2. 486 2 kee aw OO Se 16
Included and excluded species .............0 0008. 18
Allied genera and excluded species ................. 105
MEV CO DiIOMCuS SDECIGS. 45-4 9% bow Ve 2 Se ee Re Ses 105
Key to Myriophacidium species .............000 00% LZ
POSE Le OSs ok Shy ok a He BA eS Ne aOR GE 116

Ptteratiiee Cuetec e se 4 6 Kw kk eK a 4 Re Ac ee oe es 119

TAXONOMIC STUDIES IN THE PHACIDIALES: THE GENUS
COCCOMYCES (RHYTISMATACEAE)

by

1
MartHa A. SHERWOOD

SUMMARY

This paper reviews the history, nomenclature, morphology, and relationships of
Coccomyces (Phacidiales: Rhytismataceae) and provides keys to and detailed descrip-
tions of the fifty species accepted in the genus. Also included are descriptions of
Biostictis tjibodensis, four species of Karstenia, Cerion leucophaeum, Myriophacidium
corticola and M. tridentatum. Of the fifty accepted species of Coccomyces, 29 were
previously accepted in the genus, five (C. bipartitus, C. castanopsidis, C. foliicola,
C. petersii, and C. philippinus) are new combinations, and 16 (C. annulatus, C.
antillarum, C. arbutifolius, C. concolor, C. crystalligerus, C. duplicarioides, C. farlowii,
C. irretitus, C. monticola, C. palmicola, C. parvulus, C. radiatus, C. tesselatus, C.
tympanidiosporus, C. urceolus, and C. venezuelae are described as new.

INTRODUCTION

The Rhytismataceae (Ascomycetes: Phacidiales) (= Hypodermata-
ceae of most authors) constitute a well-defined family of immersed
discomycetes characterized by reduced apothecia imbedded in an
intramatrical stroma which opens at maturity to expose the hymenium;
thin-walled, inoperculate, J— asci without an obvious apical pore;
abundant paraphyses; and colorless, sheathed ascospores.” The history
and nomenclature of the group have been adequately reviewed by
Darker (1932, 1967), Terrier (1942), Nannfeldt (1932), and Tehon
(1935); only those aspects of the history relevant to Coccomyces will
be reviewed here.

Most of the twenty-two genera accepted by Darker (1967) in the
Rhytismataceae occur either primarily or exclusively on conifer nee-
dles. These conifericolous species, many of which are parasitic, were
ably revised by Darker (1932, 1967); recent additions to the group
are summarized by Hunt and Ziller (1978). In contrast, the non-conif-
ericolous species are poorly known, and generic concepts are based
on studies of a few, not necessarily representative, taxa. Tehon’s (1935)
monograph of Lophodermium, for example, emphasizes conifericolous
taxa, completely ignoring several common and distinctive tropical
species on broad-leaved angiosperms. Hypoderma was revised by
Powell (1974) in an unfortunately as yet unpublished thesis. There
is no modern treatment of Colpoma, Rhytisma or their segregates.

Of the remaining genera of the Rhytismataceae, the most serious
taxonomic problems involve Coccomyces, a large assemblage of species

"Present address: Department of Botany, University of Liverpool, P.O. Box 147, Liverpool L69
3BX, England.

? The status of the alleged brown-spored species of the group, (Criella (Sacc.) P. Henn., Phaeorhytisma
P. Henn., and Nymanomyces P. Henn.) deserves reinvestigation).

2 MARTHA A. SHERWOOD

which occur on leaves, herbaceous stems, bark, and wood, and which
have circular ascocarps opening by teeth, cylindrical or clavate asci,
and cylindric-clavate to filiform ascospores. This assemblage is the
subject of the present revision.

HISTORY

De Notaris (1847) erected Coccomyces as a monotypic genus of
the Hysteriaceae, typified by Hysterium tumidum Fr. (= Coccomyces
tumidus (Fr.) de Not.). At the same time he explicitly excluded three
species now considered to belong in Coccomyces, C. coronatus (as
Lophodermium phacidium), C. dentatus, and C. delta, placing them
in Lophodermium p Phacidioides. Darker (1967), who considered C.
coronatus and C, tumidus to be synonyms, regarded this arrangement
as “curious.” In fact, C. tumidus and C. coronatus are at least as
different from each other as Hypoderma rubi (Pers. ex Chev.) de
Not., type of Hypoderma, and Lophodermium arundinaceum (Schrad.
ex St. Amans: Fr.) Chev., type of Lophodermium, two genera which
have been considered distinct since 1826. In addition to the differences
in asci and ascospores (Hypoderma and C. tumidus have long-stalked,
clavate asci and short-cylindrical ascospores; C. coronatus and Lopho-
dermium arundinaceum have nearly sessile, cylindrical asci and
filiform ascospores), C. tumidus lacks the preformed, regular ascocarp
dehiscence mechanism found in C. dentatus and C. delta. Hence,
under a modern arrangement of these fungi, paralleling Darker’s (1967)
treatment of the hysterothecial Rhytismataceae, de Notaris’s arrange-
ment (with Lophodermium p Phacidioides raised to generic rank) might
well be preferable to the arrangements of Nannfeldt (1932) and Terrier
(1942).

Such an arrangement, however, results in a number of unfortunate
nomenclatural difficulties. The most familiar species of Coccomyces
are relegated to Coccomycella Hohnel, a name not accepted by any
author other than its originator. Coccomyces tumidus, on the other
hand, appears to have no close relatives among the leaf and bark-
inhabiting Coccomyces species.

Coccomyces, in the sense of modern authors, developed by gradual
accretion of taxa, many of them well-known but assigned to other
genera. Until the general acceptance of ascospore characteristics as
a means of classifying discomycetes, together with the use of micro-
scopes capable of resolving the small, colorless ascospores of the
Rhytismataceae, the orbicular, discoid species continued to be separat-
ed from the hysterothecial type of the genus. Coccomyces coronatus
and its allies were considered (e.g., Fries, 1849; Fuckel, 1870) to
be typical members of the genus Phacidium, now generally considered

THE GENUS COCCOMYCES 3

to have been typified by von Hohnel (1917) with P. lacerum. However,
when Fries (1822-23) validated Phacidium, he explicitly excluded
one of the two species on which the genus was founded, leaving
only P. coronatum in Phacidium. If this constitutes valid typification
of Phacidium, the generic name should be proposed for conservation
in von Hohnel’s sense.

De Notaris (Ardissone et al., 1859) subsequently recognized the
similarities between Coccomyces tumidus and Phacidium coronatum,
and transferred the latter species to Coccomyces. Karsten (1871) made
the first systematic attempt to bring together under Coccomyces all
of the filiform- and fusiform-spored Phacidium species, including
C. pint (now Therrya), C. juniperi, C. quadratus, C. macer (a Karstenia),
C. coronatus, with Phacidium trigonum (= C. tumidus) reduced to
infraspecific rank, and C. rubi (= Coleroa). He also transferred the
genus from the Hysteriaceae to the Phacidiaceae.

Rehm (1887-96) added a number of additional species, again consid-
ering Phacidium trigonum as a subspecies of Coccomyces coronatus,
while recognizing C. tumidus as a species of Lophodermium. At the
same time he errected the Hypodermataceae (a later, more familiar
name for the Rhytismataceae), but included Coccomyces in the Pha-
cidiaceae. Saccardo (1889) recognized C. coronatus, C. trigonus, and
C. tumidus as distinct species, accepted other species included by
Karsten and Rehm, and added a number of extralimital species of
Phacidium not included by those authors in their regional floras.
Minor contributions by other authors, consisting of descriptions of
additional species, are discussed under the species concerned.

After Duby (1861), whose treatment of Coccomyces was identical
to that of De Notaris, the first major attempt to reorganize the fungi
now included in the Rhytismataceae was that of von Hohnel (1917),
who divided the Phacidiales into six families on the basis of orientation
with respect to the host. The Schizothyrieen consisted entirely of
loculoascomycetous leaf parasites. The Leptopelteen were of a mixture
of subcuticular fungi including Coccomyces, with its correct type
species C. tumidus, Rhytisma, Bifusella, Duplicaria, Lophodermina,
and a variety of Loculoascomycetes. The intraepidermal Dermatopel-
teen included Coccomycella (= Coccomyces s.1. pro parte) and the
majority of Rhytismataceae. The subepidermal Phacidiaceae were a
mixture including Phacidium, Lophodermium, and Coccomycetella,
a genus not now considered (Nannfeldt, 1932) to be related to
Coccomyces. The Phacidiostromaceen were characterized by deeply
sunken leaf-inhabiting stromata. The Cryptomyceteen, with deeply
sunken bark-inhabiting stromata, included Sporomega, Colpoma, and
Therrya. This arrangement is untenable by modern systems of classi-
fication, since it juxtaposes within the same family genera now

4 MARTHA A. SHERWOOD

considered to belong to separate subclasses.

Nannfeldt (1932) rejected this classification of the Phacidiales,
removed the most discordant elements included by von Hohnel in
the group, and recognized a single family, Phacidiaceae, including
hysterothecial forms as well as Phacidium and Coccomyces. Darker’s
(1932) treatment of the Hypodermataceae of conifers included only
hysterothecial forms traditionally included in that family.

Terrier (1942) examined the genera of Phacidiaceae sensu Nannfeldt
and separated them into three families. The Hypodermataceae were
characterized by a complex covering layer of radiating, often carbonized
hyphae, solitary ascocarps, and J— asci. The Rhytismataceae possessed
ascocarps like those of the Hypodermataceae immersed in a complex,
compound stroma. The Phacidiaceae were characterized by a covering
layer of vertically-oriented, prismatic cells, and J+ asci. Although
the orientation of cells in the covering layer of Coccomyces is often
vertical, and Terrier’s suggestion that the Phacidiaceae are ascolocular
is evidently incorrect, the distinction between the Hypodermataceae
and the Phacidiaceae on the basis of ascus characters appears to be
a valid one. More recent authors (Darker, 1967; Dennis, 1977; Korf,
1973) have united the Rhytismataceae and Hypodermataceae. Terrier
briefly redescribed several species of Coccomyces, including C.
coronatus and C. dentatus but not C. tumidus.

Since 1942 no further revisions of Coccomyces have appeared.
Darker (1967) discussed the genus briefly, with Coccomyces coronatus
incorrectly considered as its type and C. tumidus as a synonym. Dennis
(1977) used the name C. coronatus for C. tumidus. B. Eriksson (1970)
provided a useful account of the species on Ericaceae in Fennoscandia.

RELATIONSHIP OF COCCOMYCES TO OTHER GENERA

According to the most current system for delimiting the genera
of the Rhytismataceae (Darker, 1967), genera having ascocarps which
open by teeth are set aside definitively from those genera with
hysterothecial ascocarps. Hence, it should be possible to consider
the generic limits in one group without reference to the other. It
is questionable, however, whether this is an appropriate model for
evolutionary relationships in the group. The majority of wood-inhabit-
ing species of Coccomyces, as well as Sporomega, Duplicaria, some
taxa assigned to Colpoma, and a few species of Lophodermium, lack
any preformed dehiscence apparatus and split open irregularly, usually
by a longitudinal slit if the substrate has a strong longitudinal grain,
otherwise by teeth. According to D. C. Minter (personal communica-
tion) two species of Lophodermium which occur on both needles
and cones of Pinus may exhibit stellate dehiscence when on cones.

THE GENUS COCCOMYCES 5

In contrast, the majority of the hysterothecial Rhytismataceae and
a large proportion of the foliicolous Coccomyces species have distinct,
differentiated, often rather elaborate dehiscence mechanisms. In the
former undifferentiated group the mode (linear or stellate) of ascocarp
dehiscence is probably of less significance taxonomically than in the
latter group.

The species of Rhytismataceae which have a well-defined preformed
dehiscence mechanism have a number of other common characters,
including a foliicolous or acicolous, intraepidermal habit, production
of pycnidia, bleaching of the substrate to form discrete lesions bounded
by a black line, and a tendency toward parasitism. If these characters
are accorded equal weight with the ascocarp shape in determining
the relationships of the genera of Rhytismataceae the effect is to group
the foliicolous species of Coccomyces with the foliicolous species
of Lophodermium and not with the wood-inhabiting Rhytismataceae
which open by teeth. More information is needed on taxa which occur
on Angiosperms, particularly in the tropics, before this question can
satisfactorily resolved.

If the rhytismataceous fungi which open by teeth are considered
in isolation, there remain a number of problems in the delimitation
of genera and their relationship to one another.

The characters used by Darker (1967) for defining genera of hys-
terothecial Rhytismataceae included ascus shape (whether cylindrical,
cylindric-clavate, or clavate), ascospore morphology, the shape of the
apothecia (whether elliptic of nervisequious), the number of apothecia
in a stroma and the orientation of the ascocarps with respect to the
surface of the substrate. Ascospore characters considered of generic
importance included shape (rod-shaped, clavate, filiform, bifusiform)
and septation. Other authors have regarded paraphyses characters to
be of generic importance. For example, Reid and Cain (1961) consid-
ered the knobbed paraphyses immersed in a brown gel to be diagnostic
of Therrya, and Dennis (1958) separated Cerion from Colpoma on
the grounds that the former lacked the circinate paraphyses of the
latter.

With the exception of ascus and ascospore shape, these characters
appear to be independent of each other. Therefore, if each is considered
by itself to be diagnostic at the generic level, as shown in Table
1, the potential number of genera in the group is high, being the
product of septation level (2 states), spore shape (4 states), paraphyses
type (4 states), ascocarp shape (4 states), and orientation with respect
to the surface of the substrate (4 states), or 512 genera if all the
slots are filled, a figure exceeding the number of species now known
in the Rhytismataceae.

Rather than carry such a scheme to its absurd logical conclusion

6 MARTHA A. SHERWOOD

TABLE 1. CHARACTERS OF GENERIC IMPORTANCE IN THE RHYTISMATACEAE
(AFTER DARKER, 1967).

Spores Spore Shape Paraphyses Ascocarp Shape Orientation
septate rod-shaped simple nervisequious subcuticular
nonseptate clavate branched linear-elliptic intraepidermal
filiform circinate compound subepidermal
bifusiform saa in circular intracortical
a ge

and distribute the species of Coccomyces among a myriad of segregate
genera, a system which shows little potential for being natural or
elucidating evolutionary relationships, I have adopted a conservative
approach in which all of the species of Rhytismataceae having
ascocarps which open by teeth have been retained in Coccomyces
except those which belong in genera already accepted in the literature.
This approach is admittedly conservative. In order to avoid creating
large numbers of new combinations in a group where future research
may well again suggest altering generic limits, I have sacrificed a
degree of uniformity in generic definition. The distinctions between
the accepted segregate genera (Cerion, Duplicaria, Duplicariella,
Lasiostictis, Moutoniella, Myriophacidium, Naemacyclus, Sporomega,
Therrya, and Tryblidiopsis) and Coccomyces are discussed briefly
below, as are the reasons for transferring those genera in the above
list, which are traditionally considered to belong to other groups,
to the Rhytismataceae.

KEY TO THE GENERA OF RHYTISMATACEAE
WITH ORBICULAR ASCOCARPS WHICH OPEN BY TEETH
The key also includes selected non-rhytismataceous genera of similar growth habit;

their names appear in parentheses. For a more complete key to immersed, stromatic
discomycetes, see also Sherwood (1977a).

1. On wood, bark, or coniferous cone scales ...... 2.00. 2.
1’. On leaves, rarely on herbaceous stems .. 2... 00000000000 ee ee 9.
2(1). Stroma fleshy, thick, well-developed, noncarbonized or with a carbonized
crust and gelatinous interior, without a preformed dehiscence mechanism . . 3.
2’(1). Stroma heavily carbonized or much reduced ................ 5.

3(2). Covering stroma colorless or nearly so, of small, vertically oriented cubical cells;
asci saccate to clavate; spores closely septate; hymenial gel sometimes blueing in iodine

Glaic Secane Ao thant Aye, @ tele e's Ge WY © cern ws es es ae dr ed ee ee (Karstenia).
3’(2). Stroma pigmented. Spores usually nonseptate and sheathed; asci clavate or
cylindrical, often pointed at the apex... 2.0. 2. 4.
4(3’). Ascospores long-clavate to filiform. Ascocarps immersed, not erumpent,
tending to be elongate. On Ericaceous twigs ............. Sporomega.

4'(3’). Ascospores oblong to clavate. Ascocarps erumpent, pseudostipitate. On
twigs of various hosts, typically conifers ............... Tryblidiopsis.

THE GENUS COCCOMYCES 7

5(2’). Covering stroma heavily carbonized, fringed internally with a conspicuous layer
of hyphal hairs visible to the naked eye. On cone scales of Pinus ..... Lasiostictis.
5’(2’). Covering stroma without a prominent internal fringe of hairs, sometimes with
gelatinoussperipNySOldSs.,, Sagat ts, she doh usec viv aula A i ee eee are es 6.
6(5’). Covering layer essentially absent, reduced to a layer of crystals .... .
Se errr gto cake rte ihc Meaty, 2) Shs a iekl ce, Ge te oh, Ss can coy Spel cd ve ee ee Naemacyclus.
6'(5’). Covering layer well-developed ........0...0........004.% 7.
7(6’). Apothecia erumpent, eventually nearly superficial, large, with a bright red hyme-
nium and black rugose border. Ascospores long-filiform. Australia, tropical America

Food Covet yg SeRE Na SC alos eerey Ace Uy Ley PANS 86 Wi) Pe Ale) te A He ot sa oy RR GUE ro ede Cerion.
7'(6’). Apothecia remaining immersed, or, if erumpent, covered by stroma when dry;
hymenium variously pigmented but not bright red... 2... 0 8.
8(7’). Epithecium of inflated paraphyses cemented in a brown gel; ascospores
Septate <OMCONMErS Wee cP cane o nS: G's) <a ck Vay Te, gio ee oe Therrya.
8’(7’). Epithecium colorless or yellowish. Ascospores generally nonseptate. On
VariousiMOsts, me, ae ss 6 le Sh oe ob Ob ieee ee Coccomyces.
9(1’). Covering stroma reduced or absent, often crystalline... ........2... 10.
9’'(1’). Covering stroma well-developed ......................04. 12.
10(9). Paraphyses cemented apically in a brown gel; ascospores filiform. Java
Soup IRMECEE Sco Dee te% a: ic ve. Se a, ee) 4, Sy A oe Moutoniella.
10’(9). Paraphyses not cemented in a brown epithecial gel .......... ie
11(10’). Asci cylindrical, with a prominent apical cap. Ascospores closely septate.
Parasitic om living leaves = 2:%. 4 06 5 heck hres ede eo a eee we eo es ees (Biostictis).
11’(10’). Asci without a prominent apical cap; spores nonseptate or distantly septate
ree anes eevee gee asarMree a eaMis sic! fe we ai Ga Gi Fs, eto et SE oie eee Naemacyclus.
12(9’). Ascospores bifusiform .................0000- Duplicaria.
12’(9’). Ascospores some other shape .............0 0000000 13.
13(12’). Ascospores ovoid .. 0... 0 0 0 14.
13’(12’). Ascospores long-cylindrical, clavate, or filiform. ........ Coccomyces.
14(13). Ascocarps subcuticular. Stroma fleshy, without a preformed dehiscence
MECHANISM Wa. -vole. 8) fo ca e e fhe & Ae en Duplicariella.
14’(13). Ascocarps intraepidermal. Stroma carbonized, with a preformed dehis-
cence mechanism................0.. 0000004 ee Myriophacidium.

Of the above genera, Biostictis Petr., discussed because it includes
a species described as a Coccomyces, is evidently Ostropalean. The
affinities of Karstenia are obscure, but do not appear to lie with the
Rhytismataceae.

Sporomega has, in the past, been synonymized with Colpoma Wallr.
and included among the hysterothecial Rhytismataceae. Eriksson
(1970) considered it distinct from Colpoma on the grounds that it
lacked a distinct dehiscence apparatus and lip cells. The ascocarps
are elongate but occasionally open by teeth. The rather fleshy stroma
separates Sporomega from the majority of the wood-inhabiting species
of Coccomyces. Microanatomically it is rather similar to Coccomyces
tumidus, which is, however, totally dissimilar ecologically.

Tryblidiopsis Karst., and the closely related, possibly synonymous
genus Discocainia J. Reid and Funk, have usually been assigned
to the Helotiales by modern authors (Korf, 1973; Reid & Funk, 1966).
Both genera, however, share several characters which suggest that
they are related to the corticolous species of Coccomyces. Stalked
clavate asci with no demonstrable apical pore apparatus, sheathed,

8 MARTHA A. SHERWOOD

colorless spores, and fruit bodies which open by splitting a partially
carbonized covering layer radially, all suggest rhytismataceous affini-
ties. The taxonomy of this group of fungi will be discussed in greater
detail in a separate paper.

Lasiostictis, formerly included in the Stictidaceae, was considered
by Sherwood (1974) and Dennis (1977) to belong to the Phacidiaceae,
a position based on the vertical orientation of the covering layer and
an alleged J+ reaction of the ascus tip. Since many indubitable species
of Coccomyces have vertically oriented cells in the covering layer,
this character cannot be used by itself to determine to which family
an immersed phacidealean fungus belongs. As for the asci, their
structure more closely resembles the asci of the Rhytismataceae and
the observations on the ascus pore are open to question since the
alleged pore region is minute and a diffraction pattern might well
be mistaken for a blueing reaction. Di Cosmo (1979) considers the
genus to be rhytismataceous.

Naemacyclus species whose ascocarps open by teeth are closely
related to Coccomyces, differing in having a much reduced covering
layer and an invariably subepidermal habit. This group was partially
revised by Sherwood (1977b) under the name Propolis. Cerion is
discussed under a separate heading, below.

Both Nannfeldt (1932) and Reid and Cain (1962) doubted that
Therrya belonged in the Rhytismataceae. Neither, however, was able
to place it satisfactorily in another family of discomycetes. The two
known species are quite isolated and share a combination of many
distinctive characters, but most of these characters (somewhat truncate
asci, septate, sheathed spores, inflated paraphyses imbedded ina brown
epithecial gel, and a massive corticolous stroma) can be found else-
where in the family. The appendages on the spores of T. fuckelii
are the only character unique to Therrya. Surely the genus is better
accomodated near the wood-inhabiting species of Coccomyces than
with the foliicolous needle pathogens with amyloid asci of the
Phacidiaceae s. str.

Moutoniella is essentially a Propolis with a brown agglutinated
epithecium, therefore differing from that genus no more than Cocco-
myces clusiae differs from the other foliicolous species of Coccomyces.
Duplicaria and Duplicariella differ from most species of Coccomyces
(but not from its type, C. tumidus) in having fleshy, subcuticular
stromata lacking preformed lines of dehiscence; additionally, Dupli-
caria has bifusiform ascospores and Duplicariella ovoid ascospores.
Myriophacidium is discussed under a separate heading, below.

Mention should be made here of Godronia (Leotiaceae: Scleroderri-
doideae). The species accepted by Groves (1965) in the genus are
all easily distinguished from Coccomyces by their substipitate asco-

THE GENUS COCCOMYCES 9

carps with a well-developed exipulum (which may be toothed) and
typical helotialean short-stalked asci with an obvious J+ pore, but
there remains a residue of species formerly assigned to the genus
for which Groves could locate no satisfactory alternative placement.
Of these, only G. castanopsidis, G. treleasei, G. splendida, G. jamai-
censis and G. juniperi appear from their descriptions and/or examina-
tion of specimens to be rhytismataceous. Groves’s list of excluded
species, however, should be consulted for erumpent filiform-spored
discomycetes which cannot be identified here.

MATERIALS AND METHODS

The observations in this paper are based primarily on dried her-
barium specimens. Coccomyces ascocarps, adapted to alternate wetting
and drying in nature, preserve well and retain most of their diagnostic
characters.

Ascocarps were rehydrated in water, sectioned at 15 or 20 jm on
a freezing microtome, and mounted in Melzer’s reagent or lactophenol-
cotton blue for microscopic observation. Additional crush mounts
were used for observation of asci and spores. For more routine
identification gross morphology, ascospore dimensions, paraphyses
characters, and host are usually adequate for determining a Coccomyces
to species; sections provide additional characters of stroma and
exciple.

The only species cultured during the study was Coccomyces strobi.
Mass ascospore isolates of this species produced compact, slow-grow-
ing mycelial colonies, initially colorless but darkening with age, on
potato dextrose agar. No conidia were produced.

Because they are inconspicuous and occupy a specialized habitat,
Coccomyces and its allies often escape the attention of the general
mycological collector. They are, however, worthy of attention, since
they are diverse, preserve well by drying, and can be found fruiting
in areas and at times of the year when few other fungi are available.

Leaf inhabiting species are easily recognized in the field because
of the distinctive bleached lesions they cause. Fruitbodies are generally
found only on the upper surface of leaves in the topmost layer of
litter. Host species should be recorded if possible, and collections
from different hosts kept separate. To avoid collecting effete material
the collector should select leaves which are not in an advanced state
of decay. If the hymenium is exposed it will normally be a color
other than black; specimens which appear entirely black in wet weather
are probably not sporulating.

Twig-inhabiting species are less conspicuous. Small twigs of trees
whose lower branches are dying are a favored habitat. The majority

10 MARTHA A. SHERWOOD

of species occur on conifers, a few on Fagaceae. The species parasitic
on ericaceous twigs should be taken from living stems; ascocarps
on dead twigs are usually overmature. Again, the host should be
recorded. Even if the same fungus appears to be attacking several
different hosts, collections should be made from all host species.

If present, rhytismataceous fungi usually occur in quantity. Since
it is normally impossible to determine collections to species on field
characters alone, it is always advisable to make large collections, both
to ensure that fertile material is obtained and to provide adequate
herbarium specimens for future workers in the event that the species
is uncommon.

MORPHOLOGY

Vegetative morphology. The vegetative thallus of a Coccomyces
consists of intramatrical, colorless or pigmented hyphae of narrow
diameter, often accompanied by extruded crystals, presumably of
calcium oxalate. Most leaf-inhabiting species form discrete lesions,
often bounded by a thin, black line of carbonized fungal cells and
disintegrating host tissue. The entire structure is roughly analogous
to the ‘substratal stroma’ (Whetzel, 1945) of the Sclerotiniaceae.
Microsclerotia of unknown function may accompany the line.

Imperfect stage. A few species of Coccomyces produce small,
flattened, lenticular brown subcuticular or intraepidermal pycnidia
with simple, colorless, subulate phialides in a single basal layer
producing minute, unicellular, bacilliform colorless conidia. These
pycnidial stages are usually assigned to the form genus Leptothyrium,
although Darker (1967) expresses doubt that they really belong there.
Their morphology is so uniform as to be scarcely diagnostic in
differentiating between species of Coccomyces, but their presence
or absence is an important factor in the classification of species. Since
conidia are usually produced before the apothecia are developed
(perhaps they function as spermatia) the presence or absence of a
pycnidial stage could not be verified in some infrequently collected
species.

In addition to the Leptothyrium stages, also ascribed to Coccomyces
by Tulasne and Tulasne (1865), Nannfeldt (1932) and others, Lind
(1907), von Hohnel (1925) and Nannfeldt (1932) suggested that Ceuth-
ospora imperfect stages were associated with Coccomyces coronatus
aggr. The material I examined gave no evidence of an association
between Coccomyces and Ceuthospora. Since Coccomyces species
often grow in association with obviously unconnected fungi (e.g.,
Mycosphaerella) great care is needed in interpreting perfect-imperfect
stage connections. Ceuthospora is generally considered to be the

THE GENUS COCCOMYCES ll

imperfect stage of Phacidium. The fungus which Grove (1928)
interpreted as Schizothyriella quercina Thum. and claimed to be the
imperfect stage of Henriquesia quercina (= Coccomyces tumidus)
is simply immature Coccomyces tumidus. Schizothyriella quercina
(FH-Thum. Mycotheca Universalis #1864, isotype) is an entirely
unrelated coelomycete.

Perfect stage. Following the interpretation of most recent authors
(Nannfeldt, 1932; Terrier, 1942; Powell, 1974) of a rhytismataceous
ascocarp, the fruitbody of Coccomyces is viewed as an apothecium
immersed in a stroma. The entire structure is shown diagrammatically
in Figure 1-I. A young apothecial stroma is initially intramatrical,
either subcuticular, intraepidermal, or subepidermal (on leaves), intra-
cortical (on bark) or immersed in decorticated decaying wood. The
position of the developing apothecium in relation to the substrate
has been used as a criterion for delimiting genera in the group (von
Hohnel, 1917; Tehon, 1935), but is used here only at the specific
level, where it appears to be a constant character.

The stroma consists of a covering layer (b) composed of one or
more layers of thick-walled, dark, globose, radiating, or epidermoid
cells, a basal layer (h) constructed like the covering layer, and (at
least in most species) an internal differentiated layer of tissue (here

Fic. 1-I. Cross section of a rhytismataceous ascocarp (diagrammatic):—a. lip cells.—b. covering
stroma.—c. periphysoids.—d. proper exipulum.—e. hymenium.—f. internal matrix of stroma.—g.
subhymenium.—h. basal stroma.—i. host tissue. Fic. 1-II. Asci of immersed discocarpous Ascomy-
cetes:—a. Rhytismataceae.—b. Phacidiaceae or Dermateaceae.—c. bitunicate ascus.—d. Stictidaceae.

12 MARTHA A. SHERWOOD

termed the intra-stromal matrix) which breaks down as the developing
apothecium enlarges and ruptures the overlying covering layer. This
layer may be absent in the mature apothecium. Its structure may
be hyphal or pseudoparenchymatous and appears to be a useful feature
in defining species.

In a mature apothecium the covering layer may consist only of
stromatic, heavily carbonized cells, or may be differentiated, with
a layer of colorless, fleshy pseudoparenchymatous tissue or periphy-
soidal hairs (c) on its inner face. These structures, often gelatinous,
swell when rehydrated and appear to aid in opening and closing
the stroma.

In hysterothecial Rhytismataceae (e.g., Lophodermium spp.) the
apothecia open by splitting the overlying covering layer longitudinally
along a preformed line of thin-walled, noncarbonized cells which
may later give rise to a compact layer of differentiated hyphae termed
periphyses (Tehon, 1935) or lip cells (a). The opening mechanism
of Coccomyces is rarely so elaborate. Most wood and bark inhabiting
species appear to open irregularly, without any distinct dehiscence
apparatus. Coccomyces delta and allied species have an opening
mechanism consisting of a stellate figure of thin-walled, noncarbonized
cells flanked by thick carbonized ribs, as in Lophodermium; however,
there is no subsequent elaboration of the marginal cells. Some species
also appear to have a flexible “hinge” of colorless cells near the junction
of the covering and basal layers which probably aids opening and
closing of the stroma.

A final form of dehiscence mechanism involves either a central
thickening in the covering layer which appears as a papilla in the
immature apothecium, or a system of thickened stellate ribs. The
covering layer tends to tear along the junction between thicker and
thinner portions of the stroma, but more irregularly than in the
preceding group.

Coccomyces apothecia may be orbicular, polygonal, or irregular
and elongate in face view. Shape was much emphasized as a taxonomic
criterion in the early literature, but appears to be variable in most
species. The color of the fresh hymenium ranges from pale grey to
yellow, orange, olive green, dark grey, or brown, and appears to be
sufficiently useful as a taxonomic character to warrant taking accurate
field notes.

The apothecium consists of a usually colorless, small-celled and
nondescript subhymenium (g), asci, paraphyses, and, in some cases,
a rudimentary proper exciple (d) consisting of a few layers of aggluti-
nated septate hyphae. The paraphyses, which are usually nonseptate,
provide some of the most valuable diagnostic characters for differen-
tiating species. The apices may be simple, inflated and knoblike,

THE GENUS COCCOMYCES 13

cemented in a colorless or pigmented gel, branched, or circinate.
The asci (Figure 1-Ila) range from cylindrical and nearly sessile to
long-stalked and clavate, are uniformly thin-walled, J—, somewhat
pointed at the apex, and lack an obvious apical pore. This characteristic
rhytismataceous ascus is quite different from the asci of Phacidium
and immersed Dermateaceae (b), which have a distinct apical appara-
tus and usually J+ pores, microthyriaceous leaf parasites (c), which
have thick-walled, bitunicate asci, and the Stictidaceae (d) which have
long-cylindrical asci with a distinct apical cap. Members of these
unrelated groups are occasionally confused with Coccomyces.

Ascospores of Coccomyces range from short-fusiform to long-fili-
form, are colorless, usually nonseptate, and frequently conspicuously
sheathed. As Darker (1932) and Tehon (1935) have pointed out, both
asci and ascospores of the Rhytismataceae are variable in their dimen-
sions even in apparently mature apothecia. Evidently the asci continue
to elongate after spores have been delimited. All measurements of
asci and spores in this study were made from rehydrated herbarium
specimens mounted in Melzer’s reagent. Darker (1932) mentions that
ascospores are narrower in diameter after drying. Differences in
methods of treating material probably account for differences in
measurements in recently published accounts and those reported here.
Small differences in measurements of asci and spores have not, alone,
been used to delimit species in the present treatment. A second source
of variability in recorded ascospore dimensions can be accounted
for by the presence of many species which normally produce both
4- and 8-spored asci. This feature is often overlooked in species with
filiform spores. Ascospores in 4-spored asci are invariably larger than
those in 8-spored asci of the same species.

HABITAT AND RANGE

Knowledge of the ecology of Coccomyces is limited, since no detailed
studies exist of the life cycles of any of the species now considered
to belong in the genus. A few inferences can be drawn from the
morphology and collection data, however, and by analogy with other
members of the Rhytismataceae.

Coccomyces species are often found in rather dry climates or in
microhabitats such as standing dead twigs which are too dry to support
many groups of fungi. Situations exposed to alternate wetting and
drying pose problems which fungi have overcome by a number of
adaptive strategies. It is reasonable to suppose that the covering stroma
of Coccomyces, which opens when wet and closes upon drying in
response to changes in rehydration of the periphysoidal layer or
pseudoparenchymatous matrix on its inner face, is an adaptation

14 MARTHA A. SHERWOOD

protecting the persistent hymenium during periods of drought.

The majority of species of Coccomyces fruit on recently dead plant
material. It would be worth investigating whether the mycelium could
be recovered from living plant parts. Naemacyclus niveus (Pers. ex
Fr.) Sacc. and Lophodermium pinastri (Schrad. ex Fr.) Chev., two
other members of the Rhytismataceae, occur as asymptomatic infections
on living conifer needles, fruiting only on dead material (F. Carroll
et al., 1977; G. Carroll & F. Carroll, 1978).

Data from frequently collected species of Coccomyces suggest that
most species in the genus exhibit some degree of host specificity,
but that host preference is determined more by the texture of the
invaded substrate and the ecology of the host than by taxonomic
affinity. Many authors have interpreted this apparent inability to
discriminate between unrelated hosts as evidence that the species
are plurivorous, and have consequently reduced distinct species to
synonymy. For example, Nannfeldt (1932), observing that Coccomyces
could be found growing on leaves of various trees in the same natural
collection, concluded that there were neither morphological nor bio-
logical factors that would separate C. coronatus, C. dentatus, C.
quercinus (= C. delta) and C. coronatus var. trigonus (= C. tumidus).
In addition to being distinct morphologically the four species exhibit
subtle differences in host and geographical range. Coccomyces corona-
tus, which fruits in August and September, occurs on thin-textured,
rather well-rotted deciduous leaves, chiefly in northern parts of Europe
and eastern North America, on Fagus, Betula, Castanea, Quercus
(chiefly red oaks), Populus, and Acer. Coccomyces dentatus occurs
on sound, often evergreen leaves of Quercus (red, white, and live
oaks), Castanea, Rhododendron, Lithocarpus, Berberis, Arbutus, Fagus,
Gaultheria, Myrica, and Lauraceae. It is a warm-temperate species,
common in southern Europe and the western U.S., fruiting in late
summer in northern areas but nearly year-round in California. Cocco-
myces delta is apparently confined to evergreen hosts (Quercus and
Lauraceae) in southern Europe and Macaronesia, and fruits throughout
the winter rainy season. Coccomyces tumidus occurs in late fall on
thin deciduous leaves of Fagus, Quercus (red and white), Betula,
and Populus (rarely on canes of Rubus) in Europe and northeastern
North America, and is often collected with C. coronatus.

Common species with restricted host ranges include: Coccomyces
strobi, which apparently occurs only on bark of Pinus strobus and
introduced white pines and never on other conifers; C. triangularis,
on twigs of Quercus alba; C. arctostaphyli, on leaves of Arctostaphylos
uva-ursi; C. quadratus, on twigs of Vaccinium spp.; and C. ledi on
twigs of Ledum spp. Although the hosts of most tropical collections
are unfortunately unidentified, some of the leaf fragments are suffi-

THE GENUS COCCOMYCES 15

ciently distinctive to suggest that the tropical species also exhibit
host specificity.

None of the species investigated appeared to be capable of colonizing
both bark and leaves. Rather, bark-inhabiting and _ leaf-inhabiting
species tend to fall into two distinct taxonomic groups. Those on
bark are characterized by small spores, irregular dehiscence of the
ascocarp, lack of imperfect stages, and a strictly saprophytic habit.
Those on leaves are characterized by long-filiform spores, regular
dehiscence along preformed lines of specialized cells, pycnidial stages,
and parasitism in a few species. Emphasis on host species, but not
substrate, has led in the past to lumping of unrelated species, such
as Coccomyces quadratus on stems and C. quadratus var. arctostaphyli
(= C. arctostaphyli) on leaves of Ericaceae, and C. boydii on bark
and C. boydii var. foliicola (= C. foliicola) on leaves of Myrica.

Coccomyces is unusual among rhytismataceous fungi in being rare
on monocotyledonous hosts and unknown on conifer needles. This
may reflect the strong linear grain of these substrates, which encourages
a hysterothecial growth habit.

Coccomyces is worldwide in its distribution, and probably abundant
in most areas with woody, broad-leaved vegetation, although specimens
are unavailable from some poorly collected areas. Several species occur
on Ericaceae in arctic and alpine environments, others in temperate
deciduous forest and others on the sclerophyllous leaves of the
Mediterranean chaparral. It is one of the few genera of saprophytic
discomycetes commonly encountered in tropical lowland rain forests.

Individual species tend to have restricted distributions related to
their preference for a single host or group of hosts (not necessarily
taxonomically related) which tend to co-occur. The effect of climate
on distributions is difficult to assess, since climate affects the phenolo-
gy and morphological adaptations of host as well as fungus. Foliicolous
species of Coccomyces in the eastern deciduous forest fruit on year-old
fallen leaves at the onset of leaf senescence in the autumn. This
ensures that ascospores are discharged into the air and may enable
the fungus to colonize leaves before they reach the litter layer. This
phenological adaptation would be less advantageous in an evergreen
forest, which may account for the absence of C. coronatus from warm
climates.

Coccomyces de Notaris

Coccomyces de Notaris, Giorn. Bot. Ital. 2(7-8): 14 & 38(1847).
= Lophodermium Chev. p Phacidioides de Not., 1.c. p. 42.
= Coccomycella Hohnel, Ann. Mycol. 15: 323 (1917).
Holotype species: Coccomyces tumidus (Fr.) de Not.
Apothecia imersed in an intramatrical stroma. Stromata orbicular

16 MARTHA A. SHERWOOD

to angular or somewhat elongate, never linear, at least partially
carbonized, usually consisting of a layer of carbonized hyphae running
parallel to the surface of the substrate, a basal carbonized layer, and
an internal matrix of fleshy pseudoparenchymatous cells, opening
either irregularly or along preformed lines of dehiscence composed
of thin-walled cells, the covering layer usually split into three or
more teeth, rarely opening by a longitudinal slit, the torn edges of
the teeth typically not lined with a well-defined layer of lip cells.
Apothecia ascohymenial, consisting of a subhymenium, asci, and true
paraphyses, sometimes with a rudimentary proper excipulum in addi-
tion to the stromatic margin. Asci uniformly thin-walled, J—, usually
pointed, without an obvious apical dehiscence apparatus, cylindrical
and nearly sessile to clavate and long-stalked, 4- or 8-spored. Ascospores
colorless, cylindrical, fusiform, or filiform, usually sheathed, simple
or septate. Paraphyses simple, inflated, branched, or circinate, free
or cemented in a gelatinous epithecium.

On leaves, twigs, bark, and old wood, rarely on Rubus canes,
saprophytic or less commonly parasitic, throughout the vegetated parts
of the world.

KEY TO ACCEPTED SPECIES

1. Ascocarps on leaves, fern rachides, or Rubus canes ...............0.. 21.
1’. Ascocarps not on leaves; on wood or bark... 2... ..0.000000 0000020 2.
2(1’). Parasitic, fruiting on living twigs of Ericaceae .............., 3;
2’(1’). Saprobic, or in any case not fruiting on living twigs of Ericaceae .. . 4.
3(2). On Ledum. Spores less than 50 wm long, broadly sheathed ........ C. ledi.
3’(2). On Vaccinium, Gaultheria, and Rhododendron. Spores more than 50 pm long
pret Gi SB, gots Nahata Gen sib Abo og “g™ ds ol kay @ E.G abe e Gice wT alse ee, @ eee ae C. leptideus.
4(2’). Ascospores all less than 50 pm long. .................... 5,
4'(2’). Ascospores all or almost all over 50 pm long .............. 13.
5(4). Ascocarps developing beneath a common black stromatic crust ......... 6.
5’(4). Ascocarps not developing beneath a common black stromatic crust ..... . 7.
6(5). Ascospores fusiform, 3-4 wm broad... .............. C. atactus.
6’(5). Ascospores cylindric-fusiform, 1-1.5 wm broad ........ C. parvulus.
7(5’). Ascocarp with lip cells. On cactus pads, sometimes opening by a longitudinal
SMG. de Fe He Ge Gee ow A ee ale Oe Be bo ee es C. wagnerianus.
7'(5’). Ascocarps without lip cells 2.2... 00000000000 000000000004 8 8.
8(7’). On recently dead, corticate twigs of white (5-needle) pines ....... 9.
8'(7’). On other hosts 2... 0 10.
9(8). Occurring in North America, chiefly on Pinus strobus. Basal stroma and ac-
companying matrix not well-developed... ..............0 2.0004 C. strobi.
9’(8). Occurring in Pakistan on P. wallichiana. Basal stroma massive, subtending a
matrix of thick-walled hyphae ... 2... ..0..00020.0.... 0.00040. C. papillatus.
10(8’). Ascospores more than 2 wm broad ................00.4. 11.
10’(8’). Ascospores 2 wm broad or less... 2... 12.

11(10). Paraphyses nodose. On Pseudotsuga and Abies, western North America... .

i Be eek Ae oR Ae, 9s em Ge TR, Ho ey en hs Ae wT Gt ore A ow Se C. pseudotsugae.

11’(10). Paraphyses circinate. On Tsuga spp., western North America .........

thesis eee a Re ee ew ae ne ee ee we ae ee oe cfr. C. heterophyllae.
12(10’). On Juniperus, Europe (if from North America see C. petersii) .... .
ey a anaeur de 8 aie He Ge Sh eee GRAN G WER Ga eee aw ae oe C. juniperi.
12’(10’). On Tsuga heterophylla, western north America . . . C. heterophyllae.

THE GENUS COCCOMYCES 17

1a?" |. “Paraphysescireindte: 0 Sc 6 a ewace gk ede wea ewe BE a ba 14.
13’(4’), Paraphyses noncircinate ........00 000.0000 cee ee 18.
14(13);. Periphysoids present «5-66 4c os so a oo RO De 15.
12 (13); PenripnysOils Qbsent yo 5 5% 4d & 4-0 ance bX NN ee 17.
15(14). Periphysoids netlike. On conifers, eastern North America .... . C. irretitus.
15’(14’). On Fagaceae. Periphysoids not netlike .................... 16.
16(15’). Outer surface of covering layer pruinose. On Quercus alba, eastern
THRU ICIS, eC. cp -cae hee ep k dy Ga eep a aed We, Xe C. triangularis.
16’(15’). Outer surface of covering layer not pruinose. On Quercus and Castan-
opsis, western North America ..................., C. castanopsidis.
17(14’). Basal stroma prominent, heavily carbonized. On Pinus cembra, Europe .. .
eae ood es eecet ue Cla Seee edo ne es gece ah hy as aah, > bene Sa C. cembrae.
17’(14’). Basal stroma reduced to a subiculum. On Juniperus virginiana, eastern North
PRIME IUC A ates ti emeen ya Ce ELON e oat: ok. % yk ink ooo Noah 6 RR C. petersii.
18(3’). Ascospores septate, eventually budding to form ascoconidia..... . .
Tee MM MED ee re Aref yt og ch! Meteo “ee C. tympanidiosporus.
18’(3’). Ascospores nonseptate, not forming ascoconidia ........... 19.
19(18’). Paraphyses branched apically. Apothecia in swarms beneath a common black
StrOMalIC CrUSt 5 co tae rei aerk nee es Syke a8) eh Sash gs gen C. bipartitus.
19’(18’). Paraphyses unbranched, apothecia solitary. ................. 20.
20(19’). Spores 60-100 x 2.0 um. On Erica, Azores ........... C. ericae.
20'(19’). Spores 50-55 x 1.0 um. On Myrica, British Isles... . . C. myricae.
21(1’). Apothecia subcuticular... 2... 0.0.0.0... ee ee ee 22.
21’(1’). Apothecia intraepidermal or subepidermal .................. 24.
22(21). Ascospores cylindrical, 3-4 4m broad ..........2.2.. C. tumidus.
22° (21). Aseospores filiform «2. .....066 6 64 4% Gwe be es 23.

23(22’). Ascospores under 100 1m long. On Andromeda polifolia, Siberia. ......
eee ee ne arate RG: es ah Gs Eyes ds. wx 4 wl wee ¥ ge C. duplicarioides.

23’(22"). Ascospores over 100 1m long. On Carex, Scandinavia ....... C. insignis.
24(21’). Apothecia intraepidermal ....................0 000. 25:
24'(21’). Apothecia subepidermal ......................... 45,

25(24). Paraphyses inflated at the apex, firmly immersed in a gel, forming an epithecium

ae ePrice ie Noe toe nn Es MeN Se ea Ss. ea an de gg he nat ne Ton ah Ts Et Sy © “ee? 26.

25’(24). Paraphyses apices not immersed in a gel, although sometimes inflated . . 28.
26(25). Epithecial gel brown ........................ C. clusiae.
26'(25). Epithecial gel hyaline or yellow... .................. 27.

27(26"). Apothecia with a distinct dark brown proper exciple, tropical America .. . .
C. leptosporus.

27'(26’). Apothecia without a proper exiple, Azores ............., C. foliicola.
28(25'). Apothecia on bleached spots bounded by a black line or reddish
CISCOOTOG Abed: epee ek 4 eS baa bk we bE RH Ae oh Gp 29.
28’(25’). Apothecia not on bleached spots, or bleached spots not bounded by
a black line or discolored area... 2... 4].

29(28). Ascospores more than 2 wm broad... ......2..2.2..2....- 0-004. 30.

29'(28). Ascospores 2 um broad or less ... 2.00.00... .00 0000 eee Syl
30(29). Apothecia irregular in form, without a preformed dehiscence mechanism:
pycnidia absent, strictly temperate .................., C. coronatus.
30'(29). Apothecia regular in form, with a preformed stellate dehiscence mechan-
ism, accompanied by pycnidia, temperate or tropical... 2... . C. dentatus.

31(29’). Spores 1.0 wm broad or less... 0.0 32.

Od (29 ho Spares 1-2 jm DiOad ok bce aed ew ka ba @ a Bee 38.
32(31). Parasitic. Lesions bounded by reddish discolored leaf tissue, without
Se AMIE A Slama aes ek ne <8. 2 4 de W- 44S Sal 4 el ee ee C. vilis.
32’(31). Not parasitic; spots bounded by a black line ............. 3D:

33(32"). Hymenium bright yellow-orange when dry, diffusing a yellow pigment in

NA AUOT BOOS EA STICTIOIS tel ae Se eg, goog. 48 XR a Row weed Bhs Ch SO C. tesselatus.

33/(32’). Epithecium not diffusing a yellow pigment in water... ........., 34.
34(33’). Paraphyses broader than 3 pm at the apex .............., 3D:

34’(33’). Paraphyses less than 3 1m broad at the apex ............ 36.

18 MARTHA A. SHERWOOD

35(34). Pycnidia produced, Philippines ...............-.. C. philippinus.
35’(34). Pycnidia not produced, tropical America ............05., C. limitatus.
36(34’). Apothecia very regular in outline, with a preformed dehiscence mechan-
PSAs is eves eke Oe se ee eee ea ate Gta Re ees Gee ig, Uae ana ae 37.
36’(34’). Apothecia without a preformed dehiscence mechanism, more irregular
in Outline. “6% Go. a8 we Ses ee a Gd be Pee we Oo ess 8 38.
37(36). On Lauraceae and Quercus, Europe, North Africa, Atlantic Islands . . C. delta.
37’(36). On various hosts, tropical America... 2... 2... eee C. spegazzinii,
38(36’). With a proper exciple, remaining open when dry .......... 40.
38'(36’). Without a proper exciple, closing when dry ............-. 39.
39(38’). On palm fronds, tropical America... 2...) ee ee ee C. palmicola.
39’(38’). On Canarium, Philippines .... 2... 02 ee eee C. canarii.
40(38). On Quercus, eastern North America ..........-54. C. farlowii.
40’(38). On evergreen hosts, South America... .......... C. monticola.
41(28’). Ascospores more than 21m broad .. 1... ee 42.
41’(28’). Ascospores 2 wm broad or less... 1 43.
42(41). Parasitic on Arbutus menziesii, pycnidia present... . C. arbutifolius.

42'(41). Saprobic, on Eryngium and monocots, tropical America. Pycnidia absent
ae wa ad Graeme Oe Gate ob Ohi e Se Ge ten ee C. pampeanus.

43(41’). Paraphyses branched apically . 2... 2... ee ee C. venezuelae.
43'(41’). Paraphyses not branched, more or less inflated... 2... .....0.0. 44.
44(43’). Proper exciple present . 2... 20.0.0... -55 5255005 C. antillarum.
44'(43’). Proper exciple absent,on Musa ............., C. annulatus.
45(24’). Ascospores more than 2m broad .... 1... 2... eee eee 46.
45'(24’). Ascospores 2 wm broad or less... 2 es 48.
46(45). Paraphyses circinate .. 2... 6... ee ee ee ee C. arctostaphyli.
46'(45). Paraphyses not circinate .. 2... 0... ee ee es 47.
47(46’). In bleached spots bounded by a reddish discolored line .... . C. puiggarii.
47'(46’). Not in bleached spots .. 2... eect eee eee C. consociatus.
48(45’). Ascospores 1 ym broad or less . 2... 2 2 ee 49.
48'(45’). Ascospores 1-2 pm broad .... 1... ee ee ee es 51.
49(48). Paraphyses with a solid refractive tip ..............05-. C. urceolus.
49’'(48). Paraphyses apices not solid and refractive .. 2.2... 6. eee ees 50.
50(49’). Asci 85-120 x 5.5 pm, on Palmae, tropical Africa... . . . C. cocoes.

50'(49’). Asci 180 * 7 pm. On dicotyledonous leaves, tropical America .. . .

ean ale oda. dite Sie ate ne Ad wie Foe SS hig ead ak ee ela ee C. concolor.
51(48’). Paraphyses simple, inflated at the apex, on Rhododendron, eastern North
AMe@riCa. «34. 6.5.4 466.6 Oa bea Ad RS BRS OE Oe ak oe we Ee C. radiatus.
51’(48’). Paraphyses branched apically, on ferns, South America . . C. crystalligerus.

INCLUDED AND EXCLUDED SPECIES

All of the published combinations and species included in Cocco-
myces traced, together with new species and new combinations of
species here transferred to Coccomyces, are listed below in alphabetical
order. Descriptions are provided for fifty accepted or provisionally
accepted species as well as for a few excluded taxa. Abbreviated
collection data are provided for all specimens examined during the
study, except that, in order to avoid unnecessary repetition, only
selected specimens of common species are cited. If more than one
specimen from a state or other subdivision is cited, the subdivision
is given only for the first specimen listed. Herbarium abbreviations
are taken from Holmgren and Keuken (1974). Exsiccati specimens
are indicated by title of the set and specimen number.

THE GENUS COCCOMYCES 19

The following two species, described in 1979, appeared in print
too late to be included in this paper: Coccomyces araucariae Butin
& Speer, Sydowia 31: 23 (1978 [1979]), on leaves of Araucaria
angustifolia, Brazil; Coccomyces aurei L. Vasilyeava, Mykol. Fitopatol.
13: 280 (1979) on Rhododendron aureum Siberia.

1. Coccomyces acerinus (Kunze & Schm.) Quélet, Enchirid. Fung.

337 (1886).
= Pilidium acerinum Kunze & Schm., Myc. Hefte 2: 92 (1323),
Leptothyrium acerinum (Kunze & Schm.) Corda, Icon. Fung.
2: 25 (1838).

According to the original description and illustrations, the name
Pilidium acerinum applies to a fungus with the external appearance
of a Phacidium, but with small fusiform spores not contained in
asci. Quelet (l.c.) did not provide a redescription of the species as
an ascigerous fungus or justify transferring it to Coccomyces. This
is the type species of Pilidium Kunze, a genus of Coelomycetes (Sutton,
1977).

2. Coccomyces albidus (Phill. & Harkn.) Sacc., Syll. Fung. 8: 746
(1889).
= Phacidium albidum Phill. & Harkn., Grevillea 12: 83 (1884).
= Naemacyclus phacidioides (Fr.) B. Erikss., Symb. Bot. Upsal.
19(4): 49 (1970).

For a redescription and additional synonyms of this species, see
Eriksson (1970) and Sherwood (1977b). Those Naemacyclus spp.
which open by teeth appear to be rather closely related to Coccomyces,
differing principally in their more deeply immersed, subepidermal
fruitbodies, crystalline margins, and lack of a stromatized covering
layer.

Specimen examined: NORTH AMERICA. USA: On dead leaves of Vaccinium ovatum,
California, Dr. H. W. Harkness, 1884, holotype of Phacidium albidum (k).

3. Coccomyces annulatus Sherwood, spec. nov. Figure 2

Ascocarpi primo immersi, dein aperientes, tetragoni vel hexagoni, 0.8-1.2 mm diam,
per lacinias aperientes, in macula pallida stromate non obvallato insidentes. Margo
superior stromatica 50-60 um crassa, ex hyphis intertextis carbonaceis constata. Periphy-
soidei nulli. Margo inferior stromatica 5 um crassa, ex hyphis carbonaceis constata.
Excipulum nullum. Paraphyses filiformes, apice ad 4.0-4.5 jm incrassatae, achromae.
Asci 140-160 xX 8.5-10 ym, haud pedicellati, cylindrici, 8-spori, in iodo non caerule-
scentes; sporis filiformibus, continuis, in tunica gelatinosa inclusis, 120 x 2.0 jm.
In foliis dejectis Musae paradisicae, Colombia, Amer. Austral.

Holotypus: Ny, Fungi of Colombia 5228, on banana leaves, ca. 20
km from Aguazul on the Aguazul-Sogamoso road, Dpto. Boyaca, elev.
ca. 2500 ft, leg. K. P. Dumont, S. E. Carpenter, M. A. Sherwood

20 MARTHA A. SHERWOOD

Fic. 2. Coccomyces annulatus:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

& L. A. Molina, 14 June 1976. Isotypi: COL, FH.

Etymology: annulatus (L), annulate, referring to the arrangement
of the ascocarps in a ring.

Ascocarps intraepidermal, arranged in a rough circle near the margin
of bleached spots not bounded by a black line, predominantly quadrate,
sometimes pentagonal or hexagonal, 0.8-1.2 mm diam, black, shining,
with a distinct, light-colored preformed dehiscence mechanism, open-
ing regularly by teeth. Covering stroma 50-60 wm thick, of heavily
carbonized cells 5 wm diam, lined on the inner face with colorless
pseudoparenchyma, lacking periphysoids. Excipulum absent. Basal
stroma carbonized, 5 wm thick. Subhymenium 10-15 wm thick, color-
less, without imbedded crystals. Asci 140-160 X 8.5-10 um, cylindrical,
short-stalked, J—, 8-spored. Ascospores 120 X 2.0 wm, distinctly

THE GENUS COCCOMYCES 21

sheathed. Paraphyses filiform, enlarged to 4-4.5 pm at the knoblike
apex, not imbedded in a gel.

On banana leaves, Colombia. The species is similar to Coccomyces
dentatus, but differs in having longer ascospores, no imperfect stage,
and no black line bounding the lesions. Other tropical species of
Coccomyces having inflated paraphyses have narrower ascospores
lacking a prominent gelatinous sheath. As indicated below under
Coccomyces clusiae, the name C. musae applies to a different species.

4. Coccomyces antillarum Sherwood, spec. nov. Figure 3

Ascocarpi primo immersi, dein erumpentes, trigoni vel tetragoni, 0.2-0.5 mm diam,
per lacinias aperientes, in macula pallida stromate non obvallato insidentes. Margo
superior stromatica 20-25 jm crassa, ex hyphis intertextis brunneis constata. Periphysoi-

ahaa 3
OOO CRS

Se

ra
)

Fic. 3. Coccomyces antillarum:—a. cross section of apothecium, X375,—b. covering layer in
face view, X1000. Dotted lines represent hyphae below plane of focus.—c. asci, paraphyses, and
spores, X750.—d. habit sketch, X7.5. Drawn from the holotype.

22 MARTHA A. SHERWOOD

dei hyalini, 7 wm longi. Margo inferior stromatica 5 jm crassa, ex hyphis intertextis
brunneis constata. Excipulum hyalinum. Paraphyses filiformes, apice ad 3-5 pm
incrassatae, in epithecio gelatinoso non inclusae. Asci cylindrici, haud pedicellati,
115-140 x 6-6.5 ym, in iodo non caerulescentes; sporis filiformibus, continuis, in
tunica gelatinosa inclusis, 90-110 x 1.0 um. In foliis dejectis ignotis, Antilles infer.

Holotypus: On undetermined leaves, Grand Etang, Grenada, British
West Indies, 1912-13, leg. R. Thaxter (FH). Isotypi: UC, CUP, NY.

Etymology: antillarum (L), referring to the Antilles, where the
species occurs.

Apothecia intraepidermal, epiphyllous or hypophyllous, in diffuse
bleached spots not bounded by a line, triangular or quadrate, 0.2-0.5
mm diam, opening by teeth along indistinct preformed lines of
dehiscence, the lobes black, shining, the disc yellowish. Covering
layer 20-25 ym thick, of dark brown cells 3-6 um diam, not heavily
carbonized, lined on the inner face with short hyaline periphysoids
7 X 5 pm. Lower stroma | cell layer thick, of dark carbonized cells
5 pm diam. Subhymenium colorless, 15 wm thick. Proper exciple
colorless, ca. 20 pm thick above, of septate hyphae 2.5-3.5 um diam,
with pointed ends like those of the paraphyses. Asci 115-140 x 6.0-6.5
wm, cylindrical, short-stalked, 8-spored. Paraphyses filiform, inflated
to 3-5 ym at the lanceolate apex, not firmly cemented in a gel.
Ascospores nearly as long as the asci, 1.0 1m broad, nonseptate, the
sheath apparent as a distinct refractive cap at the apex of the spore.

On coriaceous leaves, Caribbean islands, frequent. The host seems
to be identical in all of the collections cited, but I am unable to
identify it from leaf fragments. Slightly broader spores with a distinct
sheath, lanceolate paraphyses, and the absence of a line stroma
distinguish Coccomyces antillarum from other members of the C.
leptosporus complex.

Specimens examined (see also holotype, above): WEST INDIES. GUADELOUPE: Grand

Etang, 6.1.1974, Pfister 1006 (FH, Cup); Transverse Road, 4.1.1974, Pfister 844 (FH,
cup). Dominica: St. Paul’s Parish, 29.VI.1970, Korf et al. (cup-po 295).

5. Coccomyces arbutifolius Sherwood, spec. nov. Figure 4

Ascocarpi primo immersi, dein erumpentes, irregulariter orbiculati, 0.2-1.0 mm diam,
per lacinias irregulariter aperientes. Margo superior stromatica 25-30 mM crassa, ex
hyphis brunneis constata. Periphysoidea nulla. Margo inferior stromatica nulla. Excipu-
lum nullum. Paraphyses filiformes, apice ad 2.5 wm incrassatae, achromae. Asci 125-150
x 15-20 pm, clavati, pedicellati, in iodo non caerulescentes; sporis filiformibus,
continuis, in tunica gelatinosa inclusis, 75-100 x 2.5 wm. Pycnidia subepidermalia,
brunnea; conidia hyalina, continua, 3.5-5 x 1.0 pm.

In foliis vivis Arbuti menziesii, Vancouver, British Columbia.

Holotypus: FH, on Arbutus menziesii, 1938, leg. I. Mounce, distrib-
uted by DaoM as Coccomyces quadratus, ex DAOM 5347. Isotypi: BPI,
DAOM.

THE GENUS COCCOMYCES 23

Fic. 4. Coccomyces arbutifolius:—a. cross section of margin, X375.—b. ascus, paraphysis, and
spores, X750.—c. conidiophores and conidia, *750.—d. habit sketch, 7.5. Drawn from FH-ex
DAOM 5347.

Apothecia subepidermal, dark brown, densely gregarious, on discol-
ored spots not bleached or bounded by a line stroma, but marked
by a faint, whitish, superficial bloom, 0.2-1.0 mm diam, irregular
and often somewhat elongate in outline, without a preformed dehis-
cence mechanism, opening irregularly by teeth (sometimes by a
longitudinal slit). Covering layer 25-30 pm thick at the center,
becoming nearly obsolete at the edges, of dark brown, slightly carbon-
ized cells 3-5 wm diam, lined on the inner face with a layer of globose
colorless cells; periphysoids lacking. Subhymenium colorless, 20 wm
thick, the cells 3-5 pm diam, resting directly on disintegrating host
tissue; basal stroma absent. Asci 120-150 x 15-20 wm, clavate,
long-stalked, 8-spored. Ascospores 75-100 xX 2.5 jm, nonseptate,

24 MARTHA A. SHERWOOD

prominently sheathed. Paraphyses filiform, simple or occasionally
once-branched, enlarged to 2.5 pm at the apex.

Pycnidia hypophyllous, subepidermal, scattered among the apothe-
cia, flattened, irregular in outline, dark brown, consisting of a covering
layer of dark brown isodiametric cells 2.5 1m diam and a basal palisade
of colorless, simple conidiophores 15 X 1.5 wm, with a narrow neck
and no collarette; conidia simple, colorless, bacilliform, 3.5-5 x 1.0
Lm.

On leaves of Arbutus menziesii, Vancouver Island, British Columbia.
DAOM 5162, cited below, appears to have been collected on living
leaves.

Specimens examined (see also holotype, above): NORTH AMERICA. CANADA: Vancouver
Island, British Columbia, leg. F. L. Drayton, 7.1V.1938, distributed as C. quadratus,
ex DAOM 5162 (MICH).

6. Coccomyces arctostaphyli (Rehm) B. Erikss., Symb. Bot. Upsal.
19(4): 11 (1970). Figure 5
= Coccomyces quadratus [Schm. & Kunze] Sacc. var. arcto-

staphyli Rehm, Ber. Bayer. Bot. Ges. 13: 130 (1912).

Apothecia epiphyllous or hypophyllous on dead (exceptionally
living) leaves, the spots not bounded by a black line, subepidermal,
0.3-0.7 mm diam, black, shining, triangular to quadrate or circular,
opening by 3-5 teeth, without an obvious preformed dehiscence
mechanism. Covering stroma 30 pm thick, black, carbonized, the
hyphae 3-5 pm diam, fringed below with sparse hyaline periphysoids
5 X 2 um. Lower stroma ca. 5 pm thick, discontinuous in the center.
Subhymenium colorless, ca. 30 wm thick. Asci 90-110 x 10-12 pm,
cylindric-clavate, stalked, slightly thick-walled when young, 4- (rarely
8-) spored. Paraphyses filiform, 1-1.5 wm diam, simple, weakly
circinate. Ascospores 45-55 X 2-2.5 um, unicellular, multiguttulate,
prominently sheathed.

On leaves of Arctostaphylos uva-ursi, northern and central Europe
and western North America, late summer and fall. A relatively in-
conspicuous fungus, it is probably more common and widespread
than the few records indicate. Eriksson (1970) discusses this species
and the question of synonymy with Coccomyces ursinus (q.v.).

Specimens examined: EUROPE. SWEDEN: Harjedalen, Mt. Funasdalsberget, I. Nordin
3399, 10.X.1965 (ups). swiTzERLAND: auf Arctostaphylos uva-ursi, Schynige Platte,
15.7.1905, leg. O. Jaap, holotype of C. quadratus var. arctostaphyli (s); Zermatt, Ct.
Valais, Terrier, 23.X.1938 (zr); Ct. Wallis, Aletschwaldreservat, Muller, 9.9.1962 (z1);
Ct. Graubunden, H. Hess, 10.X.1958 (zr). NORTH AMERICA. CANADA. Alberta: Kananaskis,
E. Muller, 6.9.1977 (zr, FH). USA. Oregon: Sahalie Falls, Sherwood & Pike, 5.VIII.1978

(FH).

THE GENUS COCCOMYCES 25

Fic. 5. Coccomyces arctostaphyli:—a. cross section of margin, X375.—b. asci, paraphysis, and
spores, X750.—c. habit sketch, *1.5.—d. conidia found associated with lesions. Drawn from
FH-Muller 6.9.1977.

7. Coccomyces atactus Rehm, Rabenh. Krypt.-Fl. ed. 2, 1(3): 81
(1888). Figure 6
Apothecia at first immersed in blackened decorticated wood, gre-
garious, becoming confluent, 0.5-1.5 (—2.5 fide Rehm) mm diam,
generally somewhat elongate, black, shining, opening irregularly by
4-6 teeth to expose the greyish-yellow disc. Covering layer and basal
stroma heavily carbonized. Covering layer ca. 50 wm thick, the cells
5 wpm diam, vertically arranged, carbonized above, paler beneath.
Periphysoids absent. Excipulum absent. Subhymenium colorless, 15
um thick, separated from the basal stroma by ca. 50 um of colorless
subiculum. Asci clavate, long-stalked, 90-100 x 10-14 wm, 8-spored.
Ascospores fusiform, unicellular, prominently sheathed, 20-25 x 3-4

26 MARTHA A. SHERWOOD

Fic. 6. Coccomyces atactus:—a. cross section of apothecium, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

um. Paraphyses filiform, simple, enlarged to 3 pm at the apex.

On spruce wood, Tyrol, known only from the type specimen, The
unusual fusiform ascospores will immediately separate Coccomyces
atactus from other species of Rhytismataceae which occur on wood.
Like allied species (C. bipartitus, C. parvulus) it differs considerably
from typical leaf-inhabiting species of Coccomyces, and might with
equal justification be referred to Colpoma, The type of Colpoma,
however, has elongate hysterothecia with a well-defined opening
mechanism consisting of a longitudinal slit flanked with prominent
lip cells.

Specimen examined: EUROPE. AUSTRIA: auf Fichtenstock in Langenthal bei Keihter,

c. 5000 ft. in Tyrol oetz. 8/1872, Dr. Rehm, holotype of C. atactus (s).

THE GENUS COCCOMYCES ot

if]
a

OSS fe
oye, ROSS (is
BS.

Fic. 7. Coccomyces bipartitus:—a. cross section of apothecium, *300.—b. detail of an ascus,
paraphysis, and spores, x 750.—c. habit sketch of rehydrated material, X7.5. Drawn from the holotype.

8. Coccomyces bipartitus (Kauffm.) Sherwood, comb. nov.

Figure 7
= Schizoxylon bipartitum Kauffm., Pap. Mich. Acad. Sci. 9: 184
(1929).

Apothecia gregarious beneath a common black stromatic crust, 1-2
mm diam, often somewhat elongate with the grain of the wood, opening
irregularly by teeth or by a longitudinal slit to expose the grey
hymenium, without a preformed dehiscence mechanism. Covering
stroma ca. 30 wm thick, heavily carbonized, the cellular structure
obscured in mature specimens. Lower stroma black, carbonized, of
loosely consolidated hyphae 3-4 wm diam, with a few crystalline
inclusions. At the margin of the apothecia the space between the
covering layer and basal stroma is filled with colorless, somewhat

28 MARTHA A. SHERWOOD

gelatinous hyphae 2.0 pm diam. Subhymenium colorless, ca. 75 pm
thick. Asci clavate, stalked, uniformly thin-walled, 170-200 x 12-14
wm, 8-spored. Ascospores 60-80 X 1.5-2.0 xm, nonseptate, distinctly
sheathed, in two irregular series. Paraphyses filiform, colorless,
branched and enlarged to 3.0 pm at the apex, forming an epithecium.

On decorticated conifer wood, Michigan and New York, common,
according to Kauffman (l.c.). According to the original description
the ascospores disarticulated in the middle, but this is not evident
in any of the material I examined. The ascus structure alone definitively
excludes this species from Schizoxylon. It is somewhat intermediate
between Coccomyces and Colpoma.

Specimens examined: NORTH AMERICA. USA. New York: Warrensburg, 11.X.1934, A.

H. Smith 888, 898; Warrensburg, J. Lowe (micH). Michigan: on Picea nigra?, Rock
River, C. H. Kauffman, Sept. 6, 1927, holotype of Schizoxylon bipartitum (MIcH).

9. Coccomyces boydii A. L. Smith, Trans. Brit. Mycol. Soc. 3: 39
(1907). Figure 8
Apothecia intracortical, immersed, raising the bark into small pus-
tules and opening by cracking irregularly to expose the pale yellow
hymenium, 0.8-1.5 (—3 fide Smith, l.c.) mm diam, orbicular to irregular
in outline, sometimes becoming confluent, without a preformed dehis-
cence mechanism. Covering stroma up to 40 pm thick, slightly
carbonized, the cells 5-10 um diam. Periphysoids and excipulum
absent. Lower stroma incomplete, replaced by an almost colorless
pseudoparenchyma 50 pm thick, the cells 3-5 1m diam. Subhymenium
ca. 50 pm thick, colorless. Asci 90-115 xX 8 pm, clavate, stalked,
8-spored. Ascospores 50-55 xX 1.0 1m, obscurely sheathed. Paraphyses
filiform, enlarged to 2.0 pm at the apex, bent but not circinate.
On bark of Myrica gale, Great Britain, apparently known only from
the type specimen. The species is similar in apothecial morphology
to C. strobi, but the spores are larger.

Specimens examined: EUROPE. GREAT BRITAIN: ad corticem Myrica gale, Killin, Perth-
shire, D. A. Boyd, July, 1907, Holotype of C. boydii (x).

10. Coccomyces boydii f. foliicola Dennis & Spooner. See C. foliicola.
11. Coccomyces brasiliensis Karst. See Cerion leucophaeum.

12. Coccomyces brasiliensis Speg. See Coccomyces spegazzinit.

13. Coccomyces bromeliacearum Theiss. See C. dentatus.

14. Coccomyces canarii Rehm, Phillip. J. Sci. 8:403 (1913).
Figure 9

THE GENUS COCCOMYCES 29

Fic. 8. Coccomyces boydii:—a. cross section of apothecium, *375.—b. asci, paraphysis, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

Apothecia intraepidermal, scattered in prominent bleached spots
bounded by a delicate black line, on dead coriaceous leaves, triangular
to quadrate, 0.3-0.5 mm diam, black, shining, without an obvious
preformed dehiscence mechanism, opening by 3-4 teeth (rarely by
a longtitudinal slit). Covering layer 25-30 wm thick, consisting of
a heavily carbonized outer layer 15-20 wm thick and an inner layer
of vertically oriented hyphae ca. 3 pm diam. Lower stroma 10 ym
thick, carbonized only on its lower surface. Subhymenium 10 pm
thick. Proper excipulum absent. Asci cylindrical, short-stalked, J—,
8-spored. Ascospores nearly as long as the asci, 0.5-0.8 wm diam,
not obviously sheathed. Paraphyses filiform, 1.5 wm broad, simple,
not inflated.

On dead leaves of Canarium, Philippine Islands, apparently known

30 MARTHA A. SHERWOOD

\N\\
RRS Y:
NS
"%,) MAH

om \

\ \\ |
\

af

Fic. 9. Coecomyces canarii:—a. cross section of apothecium, *375.—b. asci, paraphysis, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

only from the type collection. It is rather similar to some members
of the C. leptosporus complex in South America, but does not agree
in all characters with any one of them. Rehm (l.c.) stated that the
spores were 0.05 um broad, surely an inadvertent error.

Specimen examined: Asia. PHILIPPINES: Luzon, Prov. Laguna, Los Banos, Baker 703
in herb. Rehm, holotype of Coccomyces canarii (s).

15. Coccomyces carbonaceus ([Fr.] ex Fr.) Quélet, Enchirid. Fung.
337 (1886).
= Phacidium carbonaceum [Fr.] ex Fr., Sclerom. Sueciae 210
(1821); Syst. Mycol. 2(2): 574 (1823).
= [Xyloma carbonacea Fr., Obs. Mycol. 2: 359 (1818)].

e

THE GENUS COCCOMYCES 31
= Discella carbonacea ({Fr.] ex Fr.) Berk. & Br., Ann. and Mag.
Nat. Hist. ser. 2, 5: 377 (1850).

The specimen of Scleromycetes Sueciae I examined was sterile.
Barkeley and Broome (l.c.) and Saccardo (1884) used the name for
conidial Cryptodiaporthe salicella (Fr.) Petr., but according to Sutton
(1977), Fries’s type is a Rhabdospora.

Specimen examined: EUROPE. SWEDEN: Fries, Sclerom. Sueciae 210, isotype of Xyloma
carbonacea (FH).

16. Coccomyces castanopsidis (Seaver) Sherwood, comb. nov.
Figure 10

Godronia castanopsidis Seaver, Mycologia 37: 351 (1945).
= Colpoma californicum Cash, Mycologia 50: 645 (1958).

"
ik
é:
ay
9
4
—

AXH
ETE
SSareesaNh

Laks 9%,
ey

Te

ig
Ta ; AN Ty ho5

0
enka

ae Giqierlne SN

\\

\
\
Fic. 10. Coccomyces castanopsidis:—a. cross section of apothecium, x 150.
paraphysis, and spores, X750.—c. habit sketch, X7.5. Drawn from FH-Suttle L. VHI.1978.

b. detail of ascus,

32 MARTHA A. SHERWOOD

Apothecia intracortical, rarely imbedded in stromatized wood, long
enclosed by the orbicular, black, carbonized stroma, finally opening
by 3-6 irregular teeth, sometimes by a longitudinal slit, without
preformed lines of dehiscence, 1.5-4.0 mm diam, the disc bright yellow
when fresh. Covering stroma 250 wm thick, the cells 5-8 pm diam,
the outermost 40 pm heavily carbonized, the inner face fringed with
gelatinous periphysoids 25 X 4.0 pm, agglutinated and appearing
pseudoparenchymatous. Basal stroma reduced toa subiculum of coarse
brown hyphae intermingled with pieces of disintegrating wood.
Subhymenium colorless, 100 xm thick. Asci 200-250 x 12-15 pm
(300 x 16-18 fide Seaver, l.c.), stalked, clavate, uniformly thin-walled.
Paraphyses filiform, circinate, 1.5-2 4m diam at the apex, mostly
unbranched, with yellowish granular contents. Ascospores 8, 100-150
x 2.0 pm, eventually becoming multiseptate, the cells 6-10 4m long,
with a prominent sheath.

On dead bark and wood of Castanopsis and Quercus chrysolepis,
western North America. A conspicuous species when fresh because
of its large size and bright yellow hymenium, it is probably uncommon,
but the limited distribution records suggest a preference for dry areas
which have not been extensively sampled by mycologists. Apparently
saprophytic.

Specimens examined: NORTH AMERICA. USA. California: W. B. Cooke, Mycobiota North
America 157, on Castanopsis chrysophylla, Mt. Shasta, Siskiyou Co., elev. 6000 ft,
WBC 15725, 8-18-41, isotype of G. castanopsidis (ru); California Fungi 1132, on Q.
chrysolepis, Butte Co., May-June 1936, Copeland, isotype of Colpoma californicum

(FH). Oregon: Jefferson Co., Suttle Lake, on Castanopsis, Sherwood & Pike, 15.XIII.1978
(FH); Government Camp, 29. V.1932 (osc 27,497).

17. Coccomyces cembrae Rehm, Hedwigia 24: 232 (1885).
Figure 11

Apothecia densely gregarious, developing beneath a common black
stromatic crust on decorticated wood, irregular in outline, opening
irregularly, without a defined preformed dehiscence mechanism,
0.5-1.0 mm diam, the disc dark yellowish ochraceous when dry.
Covering layer up to 60 pm thick, of uniform, isodiametric, thick-
walled brown cells 4 1m diam, carbonized above, the lowermost cells
faintly pigmented and vertically oriented but not definitely periphy-
soidal; lower stroma 5-10 wm thick, the cellular structure obscure
in mature specimens, heavily carbonized. Subhymenium 50 ym thick,
brown below, hyaline above. Paraphyses filiform, simple, weakly
circinate, enlarged to 2-3 um above. Asci clavate, long-stalked, 75-120
x 9-10 pm, uniformly thin-walled, J—, 8-spored. Ascospores 40-55
x 1.5-2.0 (—2.5 fide Rehm) wm, prominently sheathed, nonseptate
(becoming septate according to Rehm, l.c.).

On pine wood, Tyrol, apparently rare. The name has been misapplied

THE GENUS COCCOMYCES 33

Fic. 11. Coccomyces cembrae:—a. cross section of apothecium, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from s, Rehm Ascomyceten 801.

in herbaria to a number of other species of Coccomyces on decorticated
wood, all of them readily separable on ascospore and paraphysis
characters.

Specimen examined: EUROPE. AUSTRIA: Rehm, Ascomyceten 820, an faulenden ent-

rindeten Aesten von Pinus cembra, oberhalb der Gamperhofe bei Sulden am Ortler
(Tyrol), ca. 1900 m. 7/1884, Dr. Rehm, isotypes of C. cembrae (s, FH).

18. Coccomyces clematidis (Phill.) Sacc. See Karstenia clematidis.

19. Coccomyces clusiae (Lév.) Sacc., Syll. Fung. 8: 747 (1889).
Figure 12
= Phacidium clusiae Lév., Ann. Sci. Nat. Bot. sér. 4, 20: 291
(1863).

34 MARTHA A. SHERWOOD

Fic. 12. Coccomyces clusiae:—a. cross section of apothecium, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, ¥7.5. Drawn from FH-Lindig 2821.

Phacidium musae Lév., Ann. Sci. Nat. Bot. sér. 3, 5: 303 (1846),
nom. rej. prop. (Art. 69).

= Coccomyces musae (Lév.) Sacc., Syll. Fung. 8: 752 (1889).
Phacidium pluridens Berk. & Curt., J. Linn. Soc. (London)
10: 371 (1868).

= Coccomyces pluridens (Berk. & Curt.) Sacc., Syll. Fung.

8: 747 (1889).

Apothecia epiphyllous or hypophyllous, intraepidermal, on dead
leaves, in bleached spots not bounded by a black line, black, shining,
0.7-1.5 mm diam, orbicular or polygonal, the covering layer with
a distinct preformed stellate dehiscence mechanism of light-colored
thin-walled cells, opening by 4-10 (usually more than 5) teeth to
expose the dark brown hymenium. Covering layer ca. 30 ym thick,

THE GENUS COCCOMYCES 3D)

consisting of an upper layer 10 wm thick of carbonized cells 2-3
wm diam, fringed below with a layer of hyaline periphysoids 20 x
3-5 wm, unbranched, widely spaced ina gel. Basal stroma carbonized,
10-20 pm thick. Excipulum absent. Subhymenium colorless, ca. 20
um thick. Paraphyses filiform, enlarged to 5-6 um at the knobbed
apex, imbedded in a dark brown gel, forming an epithecium. Asci
cylindrical, short-stalked, 75-120 x 7-8 um, J—, 8-spored. Ascospores
50-60 X 1.0 um, nonseptate, obscurely sheathed.

On dead, fairly sound leaves of Clusia spp. and unidentified
Ericaceae, in humid forests at middle elevations, throughout tropical
America, common, conspicuous, and unmistakable on account of the
inflated paraphyses imbedded in a brown gel, a character found in
no other leaf-inhabiting species of Coccomyces.

The earliest validly published name for the species is Phacidium
musae, based on a specimen collected by Bonpland in tropical America.
The specimen deposited under this name in pc is annotated in
Leveille’s handwriting and appears to be the holotype. The host,
however, is not Musa, but Clusia. To the best of my knowledge
this species never occurs on Musa. Léveillé described the species
as having “disco nigricante,” a character fitting the species on Clusia
but none of the species known to occur on Musa. The evidence,
then, suggests that this is the correct application of the name Cocco-
myces musae.

According to the revised wording of Article 69 of the International
Code of Botanical Nomenclature, ““A name must be rejected if it has
been widely and persistently used for a taxon not including its type”
(Stafleu et al., 1978). Where the name Coccomyces musae has been
used in the literature, it has been used for species on Musa rather
than species on Clusia (see, for example, Seymour, 1929; Stevenson,
1926; Wardlaw, 1935). I would argue that among parasitic fungi, or
fungi which for other reasons are considered to be host specific and
are routinely identified through the use of a host index, an initial
misdetermination of the host may lead to persistent use of a name
in a sense that excludes its type, and that this is the case here. I
therefore propose that the name C. musae be rejected under the revised
wording of Article 69 of the International Code of Botanical No-
menclature.

Specimens examined: CENTRAL AMERICA. COSTA RICA. Prov, Cartago: Gomez 6897 (cr).
Prov. Heridia: Gomez 6898 (cR), CARIBBEAN ISLANDS. PUERTO RICO: Maricao, 16.VI.1970,
Korf et al. (cup-pR 4151). cuBpa: Fungi Cubensis Wrightiani 581, 873, 713, 724, C.
Wright (FH-cuRTIS); Fungi Cubensis Wrightiani 533, isotype of Phacidium pluridens
(FH-CURTIS). GUADELOUPE: La Soufriere, Pfister et al., Jan, 1974 (FH-GUADELOUPE 764,
774). SANTO DOMINGO: Cifferi, Mycoflora Domingensis 2241, Moca, on Clusia rosea

(BPI); San Jose de las Natas, Skuen, 11.4.1930 (pri); Mao near Santiago, V.1929,
Cifferi (zr); San Jose de las Natas, Ekman 4-1930 (imi 21430); Santiago, Cifferi, 1924 (im1

36 MARTHA A. SHERWOOD

101418). Haiti: Departement du Nort, Leonard 7020a, on C. rosea (ZT). COUNTRY UNKNOWN:
Herbier de la Amerique equatorial donné par M. Bonpland, presumed holotype of
Phacidium musae (pC). SOUTH AMERICA. COLOMBIA: Villeta, Lindig 2280, 1400 m (FH);
Villeta, Lindig 2821, 1861, alt. 1400 m, lectotype, designated here, of Phacidium clusiae
(pc); Antioquia, Dumont et al., 1976 (Nv-co 6240, 6561, 6265, 6245); Cauca, Dumont
et al. (Ny-co 1272). VENEZUELA: Sydow, Fungi Exotici 862, Puerto de la Cruz, on C.
rosea (PC, FH, BPI); Sydow, Fungi Venezuelani 36a, Catuche, on C. rosea (BPI, FH);
Eralia, Dpto. Federal, Dennis 4.7.1958 (zr); Dpto. Federal, Dumont et al. (Ny-ve 562,
832, 90).

20. Coccomyces cocoes Dennis, Kew Bulletin 1953: 50 (1953).
Figure 13

Apothecia immersed in obscurely bleached spots not bounded by
a line (type) or bounded by a delicate black line, on long-dead leaves,
quadrate to hexagonal, subepidermal, dark brown when closed, 0.2-1.0
mm diam, without an obvious preformed dehiscence mechanism,

if )
Lai
NN
;
is

TEE

f{?
LJ

wate

4

Fic. 13. Coccomyces cocoes:—a. cross section of apothecium, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

THE GENUS COCCOMYCES Br

opening irregularly by 4-6 teeth. Covering layer devoid of differentia-
tion, consisting of 10-15 pm of carbonized hyphae 3 jm diam.
Excipulum rudimentary, consisting of a few layers of colorless closely-
septate hyphae. Basal stroma complete, carbonized, 10-15 wm thick,
the cells 5 um diam; subhymenium 30 pm thick, colorless. Asci
cylindrical, short-stalked, 18 * 7 wm, 8-spored. Paraphyses filiform,
enlarged to 3-5 pm at the apex, not cemented in a gel. Ascospores
70-90 X 1.0 wm, nonseptate, obscurely sheathed.

On leaves of Palmae, west Africa. Similar morphologically to C.
spegazzinii and C. limitatus, differing from both principally in its
host and geographical range, and its subepidermal habit.

Specimens examined: aFRICA. GOLD Coast: Asanta, on Cocos nucifera, S. J. Hughes

687, 14.5.1949, holotype of Coccomyces cocoes (kK). GABON: Libreville, on Eremospatha
macrocarpa, G. Gilles 29.4.1979 (FH).

21. Coccomyces comitialis (Batsch) Dearn. & House. See C. tumidus.

22. Coccomyces concolor Sherwood, spec. nov. Figure 14

Ascocarpi primo immersi, orbiculati, 0.5-0.8 mm diam, per lacinias 4-6 irregulariter
aperientes, in macula pallida inconspicua stromate obvallato insidentes. Margo superior
stromatica ex hyphis intertextis brunneis constata, inconspicua. Periphysoidei nulli.
Margo inferior stromatica 5 jm crassa, ex hyphis carbonaceis constata. Excipulum
brunneum. Paraphyses filiformes, apice ad 5-6 wm incrassatae, achromae, in epithecio
gelatinoso non inclusae. Asci 85-120 xX 5.5 ym, cylindrici, haud pedicellati, 8-spori,
in iodo non caerulescentes; sporis filiformibus, continuis, in tunica gelatinosa inclusis,
70-100 X 0.8-1.0 pm. In foliis dejectis ignotis, Venezuela.

Holotypus: cup-Fungi of Venezuela 4266, on a leaf blade, Rancho
Grande, Aragua, K. P. Dumont, June 12, 1968.

Etymology: concolor (L), concolorous, since the apothecia appear
nearly the color of the leaf they are growing on rather than appearing
black against a bleached background as in allied species.

Apothecia immersed, hypophyllous, subepidermal, brown, in in-
conspicuous bleached spots bounded by a delicate black line, orbicular,
0.5-0.8 mm diam, without a definite preformed dehiscence mechanism,
opening by 4-6 irregular teeth to expose the pale yellowish-grey
hymenium. Covering layer nearly obsolete, consisting of 1-2 layers
of loosely interwoven brown noncarbonized hyphae 2-3 pm diam,
not accompanied by crystals. Lower stroma 5 um thick, black, carbon-
ized, separated from the subhymenium by a subiculum of hyaline
hyphae 2 pm diam. Proper excipulum 40-50 ym thick, of septate
hyphae 3-4 pm diam, brown above. Subhymenium 20 pm thick,
colorless. Asci 80-120 X 5.5 um, cylindrical, very short-stalked, J—,
8-spored. Paraphyses filiform, colorless, inflated to 5-6 wm at the
apex, not imbedded ina gel. Ascospores 70-100 X 0.8-1.0 um, obscurely
sheathed, nonseptate.

38 MARTHA A. SHERWOOD

Fic. 14. Coccomyces concolor:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, * 10, Drawn from the holotype.

On undetermined leaves, Venezuela. The almost complete absence
of a covering layer, subepidermal habit, and absence of epithecial
gel distinguish this species from Coccomyces leptosporus.

Specimen examined: see holotype, above.

23. Coccomyces consociatus Petrak, Sydowia 2: 66 (1948).
Figure 15

Apothecia scattered on discolored spots on living leaves (the spots
said to be caused by Ophiodothella sydowii), not accompanied by
stromatic structures or pycnidia, 0.5-0.8 mm diam, without a preformed
dehiscence mechanism, subepidermal, irregularly polygonal, raising
the substrate into small pustules and at length becoming erumpent,
the overlying epidermis colorless, splitting irregularly by 3-5 teeth

THE GENUS COCCOMYCES 39

Fic. 15. Coccomyces consociatus:—a. cross section of apothecium, *375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, 7.5. Drawn from Sydow 815a (z1).

to expose the pale yellow hymenium. Covering layer consisting of
1-2 layers of hyaline pseudoparenchyma, the cells 3-4 1m diam, lined
on the inner face with a compact layer of periphysoids 10 * 2.5-3.5
um. Subhymenium colorless, 15 wm thick, resting on a subiculum
of colorless hyphae 1.5-2 .m diam intermingled with crystals, becom-
ing increasingly crystalline with age. Basal stroma absent. Asci 90-130
x 8-10.5 wm, cylindrical, short-stalked, 8-spored. Paraphyses filiform,
slightly enlarged to 2.5 um at the apex, not cemented ina gel. Ascospores
60-90 x 1.5-2.3 um, nonseptate, obscurely sheathed.

On dead leaves of Cavendishia, Ecuador. The isotype specimen
examined is only in fair condition, and the above is taken in part
from Petrak’s original description. The species is intermediate between
Coccomyces and Naemacyclus; it lacks the prominent pigmented

40 MARTHA A. SHERWOOD

stroma of most Coccomyces species but has a more differentiated
covering layer and lacks the branched paraphyses of a typical Nae-
macyclus.

Specimen examined: SOUTH AMERICA. ECUADOR: in dem durch Ophiodothella sydowti
verursachten Blattflecken auf Cavendishia sp., Ekuador, Prov. Tungurahua, Hacienda

San Antonio bei Banos, leg. H. Sydow nr. 615 a, 28.XII.1937, isotype of Coccomyces
consociatus (FH).

24. Coccomyces coronatus ({Schum.] ex Fr.) de Not., Erbar. Critt.
Ital. ser. 1, 236 (1859).” Figure 16
= [Ascobolus coronatus Schum., F1. Saell. pars post. 437 (1803)].

b

i
|

Fic. 16. Coccomyces coronatus:—a. cross section of apothecium, X375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, X10. Drawn from FH, Thaxter, Intervale VIII.1901.

For other names not listed above, see discussion of dubious synonyms and misapplied names
following the species description.

THE GENUS COCCOMYCES 4]

= [Phacidium coronatum (Schum.) Fr., Obs. Myc. 1: 167
(1817)]; ex Fr., Scleromycetes Sueciae 165 (1821); Syst.
Mycol. 2(2): 577 (1823).

= [Peziza connivens Martius, Fl. Erlang. 463 (1817)].

= Lophodermium phacidium de Not., Giorn. Bot. Ital. 2(7-8):
42 (1847).

= Coccomyces epiphyllus Velenovsky, Monogr. Discom. Bohem.

43 (1934).

Apothecia in bleached spots bounded by a delicate line stroma
and accompanied by microsclerotia, on long-dead, well-rotted leaves,
orbicular or polygonal in outline, black, shining, without an obvious
preformed dehiscence mechanism, 0.6—-1.2 mm diam, splitting open
by 4-6 irregular teeth to expose the pale orange (when dry) hymenium,
intraepidermal. Covering layer ca. 20 wm thick, of brown, carbonized
cells 5-10 wm diam; periphysoids absent. Basal stroma 10 um thick,
carbonized, the cells 5 wm diam. Subhymenium colorless, 25 jm
thick, separated from the basal stroma by a hyaline subiculum with
a few crystalline inclusions. Proper exciple absent. Paraphyses filiform,
with a pronounced apical swelling 4-5 um diam, somewhat agglutinat-
ed, forming an epithecium. Asci cylindric-clavate, short-stalked, 100-
130 xX 11-13 wm, J—, 8-spored. Ascospores 60-80 X 2.0-2.5 im,
nonseptate, narrowly sheathed.

On well-rotted leaves of many species of deciduous trees, particularly
Betula and Quercus, mid-autumn. Very common. Collection records
from Europe and North America suggest that it prefers more northern
and less maritime areas than Coccomyces dentatus, with which it
is often confused.

Considerable confusion exists in the literature regarding the identity
of Coccomyces coronatus, although the above interpretation has been
included by most authors in their concept of the species. The specimen
which Fries distributed as Phacidium coronatum in Scleromycetes
Sueciae 165 is this species. I designate the specimen of Scleromycetes
Sueciae 165 in ups to be the lectotype of Phacidium coronatum. If
one interprets Article 9 of the International Code of Botanical No-
menclature as implying that all of the material included by Fries
in his concept of a species, including descriptions and specimens
of pre-starting point authors, form an integral part of the protologue
and are eligible for selection as lecotypes, then Scleromycetes Sueciae
165 must be considered a lectotype and not (collectively) the holotype
of the species.

Schumacher’s published description of the species is at best ambig-
uous. His unpublished illustration, deposited in the library of the
University of Copenhagen, which shows strongly erumpent apothecia
with a tan disc and irregular dehiscence, may represent Coccomyces

42 MARTHA A. SHERWOOD

tumidus. In view of the International Code of Botanical Nomenclature’s
recommendation not to typify species with pre-starting point descrip-
tions, and the absence of a type specimen, the Friesian type has
been chosen for C. coronatus.

Several pre-starting point names listed by Fries in his synonymy
of Phacidium coronatum appear, as well as can be determined from
illustrations of macroscopic characters, to belong to species other than
Coccomyces coronatus. These are omitted from the synonymy above.
Peziza comitialis Batsch may be C. tumidus; this is the interpretation
of Dearness and House (1925). Xyloma pezizoides Pers. opens by
a longitudinal slit, a character rarely found in C. coronatus. Peziza
laciniata Alb. & Schw., cited as a synonym by Karsten (1871, as
C. coronatus var. laciniatus) and Rehm (1887-96) is a stalked superficial
fungus now referred to Discocainia. Phacidium trigonum Kunze &
Schm., also included by Karsten in his concept of C. coronatus (as
var. trigonus) is C. tumidus. Coccomyces dentatus, C. tumidus, and
C. quercinus (= C. delta), considered to be synonyms by Nannfeldt
(1932), are distinct species readily separable on morphological grounds.

Imperfect stages have been ascribed to Coccomyces coronatus (cfr.
Nannfeldt, 1932). None of the specimens I examined had a pycnidial
stage co-occuring with ascocarps in the same lesion. Reports of
Leptothyrium stages may have resulted from confusion with C.
dentatus.

Specimens examined:* EUROPE. SWEDEN: Fries, Sclerom. Sueciae 165 (FH, ups, lectotype
of Phacidium coronatum Fr.); on Quercus, Uppsala, O. Eriksson (UME). FINLAND:
Messkuby, Sept., 1860, on Betula, Karsten, Fungi Fenniae 340 (FH). DENMARK: On
Quercus, Sord, 19.X1.1978, Pfister (FH). GERMANY: Sydow, Mycotheca Germanica 2713,
on Quercus, Brandenberg (Bp1, FH); Allesch. & Schnabl., Fungi Bavarici 457, on Quercus,
pp (FH). AUSTRIA: on Fagus, Sonntagberg, 1885 (Bp1). POLAND: Bidlowieza, Muller, 5.1X.1966
(zr); Mikldjka, Muller, 2.1X.1966 (zr). czEcHOSLOvAKIA: Bohemia, Jevany, in foliis Fagi
sylvaticae, IX. 1922, leg. Velenovsky, holotype of Coccomyces epiphyllus (prM 148699);
Bohemia, on Quercus, leg. M. Svrcek (PRM 756276, 756274, 620800, 714315, 714316,
625807); Bohemia, on Fagus, leg. Svrcek (prM 612637, 756282); Bohemia, on Betula
(PRM 756284, 149921); Moravia (pRM 150513). traty: Herb. Crittog. Ital. 256, Riva di
Valsesia, on Fagus, Cesati, Carestia (FH). FRANCE: Herb. Barb. Boiss. 1010b, Normandy
(FH). GREAT BRITAIN: Phillips, Elvellacei Britanici 201, on Quercus, Herfordshire (ppt,
FH); Hockfall, Yorkshire, Hughes & Webster, 25.1X.1948 (Bri). NORTH AMERICA, CANADA,
Nova Scotia: on Fagus, Wehmeyer 1467 (mMicu). Ontario: on Quercus, Lake Timagami,
Thompson & Pady, 25. VIII.1930 (micu, Bet). USA. Maine: on Quercus, Shear 23.VIII.1940
(BPI); on Betula, ibid. (pri); Mt. Katahdin, Shear, 1940 (Bp1); Whetzel & Sprosten, 1940
(cup 29190). New Hampshire: White Mts., O. F. Cook, VIII, 1889 (spi); Intervale,
on Fagus, Thaxter, 1901 (FH); Chocorua, on Sarracenia, Farlow (FH); on Betula, Thaxter,
1885 (FH); on Betula, Farlow, 1885 (FH), Massachusetts: Newton, Farlow, VIII.1878,
on Quercus (FH). New York: on Acer, Warrensburg, Rogerson, 3.X.1959 (Ny); Lake
Placid, on Fagus, Kauffman & Mains, 15.1X.1917 (micH); Saranac L., Mains, 25.XII.1914
(micH); Ithaca, Whetzel, 6.X.1903 (cup 1838). Iowa: G. W. Martin, 22. VIII.1936 (spi).
Michigan: Marquette, on Populus, Mains 1935 (micu); Ann Arbor, 15.1X.1938, Smith

4 . 7 . ; F 5 ; ‘ ,
For exsiccati specimens distributed as C. coronatus see also C. tumidus and C. dentatus. ‘pp
indicates a mixed collection of two or more Coccomyces spp.

THE GENUS COCCOMYCES 43

(micH). Wisconsin: Lake Geneva, VII.1903, Harper (F). SOUTH AMERICA. BRASIL: Rio Grande
do Sul, Rick, VII.1906 doubtfully referred here (s, HERB. REHM).

25. Coccomyces coronatus f. rubi Rehm, Ber. Bayer. Bot. Ges. 13:
129 (1912).

Rehm based this form on a specimen collected by Morthier in
Schweizer-Jura in 1864 and distributed in Herbier Barbey-Boissier
as Phacidium coronatum. No specimen answering this description
could be found in s or FH. The name may apply to Coccomyces
tumidus, which occasionally occurs on Rubus canes, has ascospores
of the dimensions given by Rehm, and was included by Rehm in
his concept of C. coronatus.

26. Coccomyces crustaceum Curt. in Reid & Cain. See C. strobi.

27. Coccomyces crystalligerus Sherwood, spec. nov. Figure 17

Ascocarpi primo immersi, irregulariter orbiculati, 0.7-1.2 mm diam, per lacinias
irregulariter aperientes, in macula pallida non insidentes. Margo superior stromatica
20 wm crassa, ex hyphis intertextis carbonaceis constata. Excipulum brunneum.
Periphysoidei nulli. Margo inferior stromatica 20 wm crassa, ex hyphis intertextis
carbonaceis constata. Paraphyses filiformes, ramosae, achromae, in epithecio gelatinoso
non inclusae. Asci 180-225 * 10-11 ym, cylindrici, haud pedicellati, 8-spori, in iodo
non caerulescentes. Sporis filiformibus, continuis, in tunica gelatinosa inclusis. In
laminae et petiolis filicis ignotis, Colombia, Amer. Austral.

Holotypus: ny-co 6111, on indet. fern leaf (Polypodiaceae) ca. 82
km from Medellin-Pto. Valdivia rd., Dpto. Antioquia, elev. ca. 9400
ft, leg. Dumont, Carpenter & Sherwood, 12 August 1976. Isotypi:
COL, FH.

Etymology: crystalligerus (L), forming crystals, referring to the
epithecium.

Apothecia subepidermal, deeply immersed in leaves and rachides
of an unidentified polypodiaceous fern, 0.7—1.2 mm diam, dark brown
when closed, opening irregularly by teeth, without a preformed
dehiscence mechanism. Hymenium pale translucent yellow when wet,
drying white-pruinose, dotted with discrete clumps of colorless crys-
tals. Covering stroma ca. 20 pm thick, of carbonized cells 3-5 pm
diam, without periphysoids. Basal stroma complete, ca. 20 pm thick,
of carbonized cells 3-5 wm diam. Subhymenium colorless, 25 jm
thick. Proper excipulum well-developed, 50 um thick, of closely septate
hyphae 2.5 ym diam, the cells 8-10 ym long, brown above. Paraphyses
filiform, 1.0 jm broad below, barely enlarged above, richly branched
above, colorless, crystalliferous, forming an epithecium. Asci cylindri-
cal, short-stalked, J—, 180-225 x 10-11 wm, 8-spored. Ascospores
150-180 x 1.5-2.0 um, narrowly but distinctly sheathed.

44 MARTHA A. SHERWOOD

Fic. 17. Coccomyces crystalligerus;—a. cross section of margin, *300.—b. detail of asci, pa-
raphyses, and spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

On fern leaves and rachides, known only from a single collection
from the Colombian highlands. A very distinctive species because
of its unusual host, very long asci, and crystalline epithecium.

Specimen examined: see holotype, above.

28. Coccomyces delta (Kunze) Sacc., Flor. Myc. Lusit. 13 (1893).
Figure 18
= Phacidium delta Kunze, Linnaea 5: 551 (1830).
Phacidium quercinum Desm., Pl. Crypt. France 1644 (1847).
= Coccomycella quercina (Desm.) Hohnel, Ber. Deutscher Bot.
Gesell. 35: 419 (1917).
= Coccomyces quercinus (Desm.) Terrier, Essai Syst. Phacid.

39 (1942).

THE GENUS COCCOMYCES 45

ret
“4 ,
ay y YY }
SBS) ‘ yy
1 / / y Hf ff ff f |
. a (} Y /f Ue My / / WY / y ff |
: ry \ | 4 Y] /; y/ WY If / ) / / /f |
0)
/ ERY tl! cS
Kas, ) ? // j .
ALIN RSA wu Be |
AKL ISL TDR pie ;

Fic. 18. Coccomyces delta:—a. cross section of apothecium, *375.—b. ascus, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from CUP-MM 654

Apothecia intraepidermal, on distinct bleached spots bounded by
and intersected by a delicate line, black, shining, triangular or quadrate
(triradiate or cruciate when immature), with an obvious preformed
dehiscence mechanism of light colored cells. opening by 3-4 teeth.
Covering layer 30 wm thick, of isodiametric cells 4-10 wm diam,
heavily carbonized at the center, less so at the periphery. Periphysoids
absent. Edges of teeth bordered by a fringe of periphysis-like lip
cells 1.5-2.5 ym diam, immersed in a gel (as in Lophodermium).
Lower stroma 5 yum thick, heavily carbonized, separated from the
upper stroma by a large-celled pseudoparenchymatous matrix. Subhy-
menium 25 wm thick, colorless. Paraphyses filiform, slightly enlarged
to 2.5-3 wm at the apex, simple, not immersed in a gel. Asci 100-150
x 8-10 um, cylindrical, short-stalked, thin-walled. J—, 8-spored. Asco-

46 MARTHA A. SHERWOOD

spores 80-100 xX 2.0 um, nonseptate, narrowly sheathed. Pycnidia
absent.

On dead coriaceous leaves of Lauraceae and evergreen Quercus,
throughout the winter rainy season, Mediterranean area and Atlantic
Islands, common. Coccomyces dentatus is similar, but differs in having
shorter asci and ascospores, lacks lip cells, and usually has associated
pycnidia. The species was originally described from Madeira. I could
locate no type material, but have little doubt that this is the correct
application of the name. De Thuemen, Mycotheca Universalis 1367,
Fl. Lusit. Exs. 1208, and Rabenhorst, Fungi Europaei 1208, cited
below, were distributed as Phacidium trigonum, all misidentifications.
California fungi 430, distributed as C. delta, is Stictis emarginata
Cke. & Massee.

Specimens examined: EuROPE. ITALY: on Laurus nobilis, July, 1906 (pap). FRANCE:
Desm., Pl. Crypt. France sér. 1: 1694, in fol. sicc. Querc. coccif., autun, isotype of
Phacidium quercinum (FH). PORTUGAL: de Thuemen, Mycoth. Univ. 1367, Coimbra,
1879, on L. nobilis (FH, zt); Rabenh., F. Eur. 3363, Coimbra, 1885 (Fx, z1); Flor.
Lusit. Exs. 1208, Coimbra, 1893 (FH, zT). ATLANTIC ISLANDS. MADEIRA: Ribeiro Frio, Korf
et al., 13.1.1977 (cup-mM 1486); Faja da Noguera, 21.11I.1978, Korf et al. (cup-MM-2291);
Chao dos Louros, 26.11I.1978, Korf et al. (cup-mMM 2455). CANARY ISLANDS: Tenerife,
Korf et al., January, 1976 (cup-mM 1, 217, 11, 76, 207); La Palma, Korf et al., January,
1976 (cup-mMM 817, 647, 654). Azores: Sao Miguel, 5.IIT.1978, Korf et al. (cup-MM-1845;
Phacid. Exs. #16).

29. Coccomyces dentatus ([Schm. & Kunze] ex Fr.) Sacc., Michelia
1: 59 (1882). Figure 19
= [Phacidium dentatum Schm. & Kunze, Deutschlands

Schwamme 2 (1816)] ex Fr., Syst. Mycol. 2(2): 577 (1823).
?= Coccomyces bromeliacearum Theiss., Bot. Centralbl. 27(2):
406 (1910).
= Coccomyces dentatus var. lauri Rehm in Theiss., Bot. Cen-
tralbl. 27(2): 406 (1910).
?= Coccomyces filicicola Speg., Bol. Acad. Nac. Ci. 23: 152
(1919).
= Coccomyces pentagonus Kirschst., Ann. Mycol. 34: 208 (1936).
Apothecia scattered in prominent bleached spots bounded by a black
line, intraepidermal, usually accompanied by pycnidia, 0.5-1.0 mm
diam, quadrate to hexagonal, black, shining, with a distinct stellate
preformed dehiscence mechanism of lighter colored cells, opening
by teeth to expose the dull yellow hymenium. Covering layer ca.
30 ym thick, of carbonized cells 5-6 ym diam; periphysoids and
lip cells absent. Basal stroma ca. 5 ym thick, carbonized. Excipulum
absent. Paraphyses simple, filiform, gradually enlarged to 2.0 pm
at the apex, the contents rather granular. Asci cylindric-clavate, short-
stalked, 70-105 x 8-10 pm, thin-walled, J—, 8-spored. Ascospores

THE GENUS COCCOMYCES 47

Fic. 19. Coccomyces dentatus:—a. cross section of apothecium (slightly immature), X375.—b.
ascus, paraphyses, and spores, X750.—c. habit sketch, approximately natural size.—d. habit sketch,
7.5, Drawn from FH-Parks 4084.

45-65 X 2.0 um, narrowly but distinctly sheathed, nonseptate. Pycnidia
appearing before the apothecia mature, intraepidermal, lenticular in
cross section, 0.1-0.3 mm diam, with a dark brown covering layer;
phialides in a basal layer, borne on short conidiophores, subulate,
without a collarette, 5-10 X 2-2.5 um; conidia bacilliform, nonseptate,
colorless, 4-5 x 1.0 pm.

On dead (rarely living) leaves of a wide variety of angiosperms,
notably on Fagaceae and Ericaceae, widespread and common, chiefly
in warm temperate areas, summer and fall in the northern part of
its range, throughout the year in subtropical areas. Owing to its
abundance, wide geographical distribution, and conspicuousness, this
is the most frequently collected species of Coccomyces. Coccomyces

48 MARTHA A. SHERWOOD

coronatus, with which it is frequently confused, has inflated para-
physes, longer asci and ascospores, less regular apothecia, and rarely
occurs on leaves of evergreens.

Coccomyces bromeliacearum and C. filicicola, cited as tentative
synonyms above, cannot be completely characterized from their re-
spective type specimens. The type of the former is old, and the type
of the latter completely immature. The specimens and original descrip-
tions suggest that they are very close to, if not identical with, C.
dentatus. Coccomyces dentatus var. lauri does not differ in any
recognizable morphological detail from typical C. dentatus. Cocco-
myces pentagonus likewise appears to be typical C. dentatus; infections
on Berberis aquifolium in the western U.S. agree exactly with infections
which appeared when the host was introduced into Europe.

Specimens examined:” EUROPE. GERMANY: auf Mahonia aquifolium, Brandenberg,
Tamsler Baumhiiln, V.1936, leg. P. Vogel, isotype? of C. pentagonus (zr); on M.
aquifolium, Brandenberg, Sydow Mycoth. 3136 (FH); Schmidt & Kunze, Deutschlands
Schwimme 106, on Quercus robur, isotype of Phacidium dentatum (Fu, Bri); Cassel,
on Quercus, C. Grabe 1872 (Bet); Sydow, Mycoth. Germ. 1249, on Q. pedunculata,
Lothringen (FH); Sydow, Mycoth. Germ. 2367, on Mahonia, (as C. coronatus) (FH);
Herb. Barb. Boiss. 1011 & F. Rhen. 1090, Nassau, auf Castanea & Quercus, Fuckel
(FH, BPI); Bayreuth, 4.1875, on Q. rubra (FH). CZECHOSLOVAKIA: De Thuem., F. Austriaci
973, Bohemia, on Fagus (FH). FRANCE: Contremoulins, Nov. 1872, on Castanea vesca
(rH-Patouillard). corsica: on Quercus, Terrier, 13.4.1938 (zr); on Q. ilex, Terrier, 13.4.38
(zr). BELGIUM: Libert, Pl. Crypt. Ard. IV 368, on Quercus (z1); Liege, Kinkempois,
on Quercus, leg. E. Morren (LG). 1raLy: Saccardo, Mycoth. Ital. 682, on Quercus, Novarra,
111.1899 (FH); on Castanea, Montignoso, Arx, 25.5.48 (ZT). AFRICA. TUNISIA: El Feidja,
on Q. morbeckii, I. 1893 (FH-Patouillard), ATLANTIC ISLANDS. CANARY ISLANDS: La Palma,
on Myrica faya, Korf et al., 18.1.1976 (cup-MM 888). NORTH AMERICA. CANADA. British
Columbia: Vancouver, on Berberis, Bandoni, 30.111.1960 (Bp1). usA. Pennsylvania: Ell.
& Ev., N. Am. F. 2057, on Quercus, 1X.1884 (FH). Virginia: Mansanutten, 26.1X.1932,
Davidson (pr); Airmont, 18.VII.1903, on Quercus alba, Shear (spt). North Carolina:
on Quercus, Whitewater falls, Shear, 19.VIII.1923 (spi); on Q. nigra, new Bern,
Hedgecock, 23.VI.1938 (pPi); on Q. virens, Smith Island, 23.VIII.1931, Cash (ppl).
Tennessee: on Q. michauxii, La Follette, Sharp & Hessler, 7-11-1934 (pri). Louisiana:
on Quercus, Langlois, 27.V.1895 & 30.VII.1889 (FH). Florida: Highlands Hammock,
on Quercus, Shear 118 & 132, 1937 (spi). Arizona: on Q. reticulata, Pima Co., Lindsey
153 (ariz). Washington: on Berberis, Marysville, Grant, XII.1927 (sri); on B. aquifolium,
1895, Ell. & Ev., N. Am. F. 3336 (spi, FH); on Gaultheria shallon, Orcas Isl., VI1.1906
(F). Oregon: on B. nervosa, Siskiyou Mts., Haines, 20.X.1968 (Nys); on Q. vaccinifolia,
Curry Co., Sherwood (FH); on B. nervosa, Umatilla Co., Cooke, Mycob. N. Am. 310
(BPI, FH); on Castanopsis chrysophylla, Lane Co., Sherwood, 1978 (FH); on Rhododendron
macrophyllum, Lane Co., Sherwood, 13.VIII.1975 (FH, cup). Idaho: on Mahonia repens,
Moscow, IX.1955 (FH). California: on Lithocarpus densiflora, Santa Cruz, Bonar,
24.V1.1955 (Bri); on Q. agrifolia, Solano Co. Bonar, 7.1I.1964 (spi); on Q. kelloggii,
Lake Co., Bonar, 15.V.1943 (FH); on Arbutus menziesii, Marin Co. (Bpi, Cup 47382);
on R. californicum, Parks, 11.1943, Calif. F. 1011 (FH); on Gaultheria shallon, Trinidad,
Shear, 7.VI.1939 (BPI). CENTRAL AMERICA, PANAMA: Prov. Chiriqui, Dumont & Carpenter,
2.V1I.1978 (Ny-Pa 1729). SOUTH AMERICA. VENEZUELA: Edo. Merida, on Clusia? Dumont
et al., 23.VII.1971 (nvy-Ve 2792); Edo. Merida, Dumont et al., 20.VII.1971 (Ny-Ve
2563). COLOMBIA: Putumayo, Dumont et al., 26.1.1976 (Ny-Co 3646); on Macleania,

5 . :
When more than one specimen on the same host from the same general locality was examined,
only one has been cited.

THE GENUS COCCOMYCES 49

Cundinamarca, Dumont et al. (Ny-Co 283). BRasiL: S40 Leopoldo, Rio Grande do Sul,
Theiss., Dec. Fung. Brasil. 135, Rick, 1908, presumed isotype of Coccomyces bromelia-
cearum (FH); Apiahy, s/Pteridium?, 1890 leg. J. Puiggari no. 2107, holotype of C.
filicicola (Les 28187).

30. Coccomyces dentatus (Kunze & Schm. ex Fr.) Sacc. var. hexagonus
Penz. & Sacc., Malphigia 15: 222 (1901).

No specimen of this variety is deposited in Saccardo’s herbarium
(pap), and Penzig’s herbarium was destroyed during World War II.
The disposition of the taxon cannot be determined from the original
description. The name has been applied to large specimens of Cocco-
myces dentatus with regular, hexagonal apothecia from the western
LS.

31. Coccomyces dentatus var. lauri Rehm. See C. dentatus.

32. Coccomyces dianthi (Fuckel) Rehm, Rabenh., Krypt.-Fl. ed. 2,
1(3): 79 (1888).

Phacidium dianthi Fuckel, Jahrb. Nassauischen Veriens Na-

turk. 23-24: 262 (1870).

This species is not a Coccomyces and appears to be dermateaceous
rather than rhytismataceous. It is characterized by short broad asci
65 < 14 pm, with a pronounced J+ apical ring, and 8 unicellular
or l-septate inequilateral ascospores 15 X 5 ym. It may be a member
of the Naevioideae, but does not correspond to any of the taxa discussed
by Hein (1976). The specimen of this species in the Farlow Herbarium
contains no asci, but rather fruitbodies similar in external appearance

the species.

Specimens examined: EUROPE. GERMANY: Herbier Barbey-Boissier 1012 (= Fungi
Rhenani 1091), an durren Stengeln von Dianthus carthusiana, selten, in Fruhling,
Budenbremenwalk, Oestrich (Nassau) leg. Fuckel, isotype of Phacidium dianthi (Fn,
BPI); P. Vogel, Tamsel, 24.9.1937 (FH).

33. Coccomyces dubius Rehm, Leafl. Philipp. Bot. 8: 2926 (1916).
Figure 20
= Schizochora elmeri Sydow, Ann. Mycol. 11: 265 (1913).

The type specimen of Coccomyces dubius consists of flattened,
subcuticular stromata containing a few small perithecia. The asci are
clavate, stalked, J—, uniformly thin-walled, 8-spored, and occur inter-
mingled with paraphyses. The ascospores are hyaline, ca. 30 x 3
um, nonseptate, and bear a filiform appendage at each end.

This is surely not a Coccomyces and seems to be a synonym of
Schizochora elmeri (Polystigmataceae) which also occurs on leaves
of Ficus in the Philippines, and was redescribed by von Arx and

Muller (1954).

50 MARTHA A. SHERWOOD

Fic, 20. Schizochora elmeri (Coccomyces dubius):—a. ascus, paraphyses, and spores, X750,.—b.
cross section of ascocarp, X225. Drawn from Baker, Fungi Malayani 128 (s).

Specimen examined: Asia. PHILIPPINES: Baker, Fungi Malayana 128, on Ficus minahas-
sae, Mt. Maquiling, Los Banos, May, 1914 (s, herb. Rehm, holotype; pc, FH).

34. Coccomyces duplicarioides Sherwood, spec. nov. Figure 21

Ascocarpi primo immersi, orbiculati, 0.2-0.3 mm diam, per lacinias irregulariter
aperientes, in macula pallida stromate non obvallato insidentes. Margo superior stro-
matica 40 ym crassa, ex hyphis intertextis brunneis constata. Periphysoidei nulli. Margo
inferior stromatica 10-15 ym crassa, ex hyphis carbonaceis constata. Excipulum nullum.
Paraphyses filiformes, circinatae, apice non incrassatae, in epithecio gelatinoso non
inclusae. Asci 150-200 x 12-15 yum, clavati, pedicellati, 8-spori, in iodo non caerule-
scentes; sporis filiformibus, continuis, in tunica gelatinosa inclusis, 45-63 X 1.5 um.
In foliis emortuis Andromedae polifoliae, Magadan region, ussr.

Holotypus: Taa, on Andromeda polifolia, Magadan Region, distr.
Tenkinski, Kontakt Field Station, USSR, 5.IV.1975, leg. Lar. Vasiljeva.
Isotypus: FH.

Etymology: from Duplicaria, a genus which this species resembles
in its external appearance and apothecial structure.

Apothecia subcuticular in bleached spots not surrounded by a line
on dead leaves, orbicular, black, shining, 0.2-0.3 mm diam, without
a preformed dehiscence mechanism, opening irregularly by teeth.
Covering layer 40 ym thick, the outer surface consisting of interwoven
brown noncarbonized hyphae 5 pm diam, lined on the inner face
by a matrix of interwoven, nearly colorless hyphae with a cell lumen
1.5 4m diam and very thick gelatinizing walls. Basal stroma 10-15
ym thick, of nongelatinous carbonized hyphae 3-5 um diam, separated
from the subhymenium by 75 pm of colorless, vertically oriented
hyphae widely spaced in a gel. Excipulum absent. Subhymenium
colorless, 15 ym thick. Asci 150-200 x 12-15 um, long-stalked, clavate,

THE GENUS COCCOMYCES 51

ir)

Fic. 21. Coccomyces duplicarioides:—a. cross section of margin, X375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, 7.5. Drawn from the holotype.

J—, 8-spored. Paraphyses colorless, 1-1.5 jm broad, circinate, not
enlarged above or immersed in a gel. Ascospores 52-63 X 1.5 wm,
filiform, nonseptate, tapering slightly below, prominently sheathed.
Pycnidia not seen.

On dead leaves of Andromeda polifolia, Magadan Region, eastern
USSR. The distribution of this inconspicuous species is unknown;
it should be sought elsewhere on its widely distributed host. Duplicaria
empetri (Fr.) Fuckel is similar in external appearance and ascocarp
structure, but has bifusiform ascospores.

35. Coccomyces epiphyllus Velen. See C. coronatus.

36. Coccomyces ericae Dennis & Spooner, Kew Bull. 32: 113 (1977).
Figure 22

52 MARTHA A. SHERWOOD

LLP gaat
Fic. 22. Coccomyces ericae:—a. cross section of apothecium, X375.—b. ascus, paraphysis, and

spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

Apothecia scattered on decorticated wood, not forming confluent
crusts, black, orbicular or somewhat elongate, 0.5-1.0 mm diam,
without a preformed dehiscence mechanism, splitting open irregularly
to expose the yellowish-orange disc. Covering stroma ca. 20 pm thick,
heavily carbonized, of vertically oriented hyphae. Lower stroma 10-15
wm thick, heavily carbonized, of globose cells 4-6 1m diam, separated
from the hymenium by a hyaline subiculum interspersed with a few
crystals. Subhymenium colorless, 25 wm thick. Proper exciple well-
developed, ca. 40 ym thick, the hyphae 2.5 um diam, closely septate,
carbonized above. Asci clavate, rather long-stalked, 125-150 x 8-10
wm, J—, 8-spored. Paraphyses filiform, enlarged to 2.5 um at the
apex, interspersed with brownish amorphous material. Ascospores
60-100 x 2.0 im, nonseptate, not prominently sheathed.

UU
ie)

THE GENUS COCCOMYCES

On decorticated wood of Erica azorica, Azores, known only from
the type specimen.

Specimen examined: ATLANTIC ISLANDS. AZORES: on Erica azorica, Caldesia road, above
Flamengos, Faial, 23.3.1975, leg. R. W. G. Dennis, holotype of Coccomyces ericae
(K).

37. Coccomyces farlowii Sherwood, spec. nov. Figure 23

Ascocarpi primo immersi, dein erumpentes, tetragoni vel hexagoni vel irregulariter
orbiculati, 0.5-1.0 mm diam, per lacinias 46 aperientes, in macula pallida stromate
obvallato insidentes. Margo superior stromatica 5-10 jm crassa, ex hyphis carbonaceis
constata. Periphysoidaei nulli. Margo inferior stromatica 30 jm _ crassa, ex hyphis
intertextis carbonaceis constata. Excipulum brunneum. Paraphyses filiformes, apice
ad 3-5 ym incrassatae, achromae, in epithecio gelatinoso non inclusae. Asci 115—130
x 10-12.5 pm, cylindrici-clavati, haud pedicellati, 8-spori, in iodo non caerulescentes:
sporis filiformibus, continuis, in tunica gelatinosa inclusis, 60-75 x 2.0 4m. In foliis
dejectis Quercus rubrae, Massachusetts, USA,

oe: —

Fic. 23. Coccomyces farlowii:—a. cross section of apothecium, X375.—b. ascus, paraphysis,
and spores, X750.—c. habit sketch, *7.5. Drawn from the holotype.

54 MARTHA A. SHERWOOD

Holotypus: FH, on fallen leaves of Quercus, Estabrook Woods,
Concord, Mass., M. A. Sherwood 9/25/1977.

Etymology: farlowii, after W. G. Farlow, cryptogamic botanist at
Harvard University.

Apothecia intraepidermal, at first immersed, dark brown, shining,
orbicular to polygonal or irregular in outline, in bleached spots
bounded by a line stroma on dead leaves, 0.5-1.0 mm diam, without
a preformed dehiscence mechanism, opening by 3-6 teeth to expose
the olivaceous hymenium, the teeth becoming reflexed, the hymenium
remaining exposed when dry. Covering stroma 5-10 ym thick, heavily
carbonized, devoid of internal differentiation; periphysoids absent.
Lower stroma 30 pm thick, the basal part heavily carbonized, merging
above into a layer of lighter brown cells 3-5 pm diam. Excipulum
well-developed, up to 75 ym thick, of closely septate hyphae 3-4
wm diam, faintly brownish above. Subhymenium colorless, 40 4m
thick. Paraphyses filiform, gradually expanded to 3-5 pm at the apex,
not agglutinated at the apex. Asci cylindric-clavate, short-stalked, J—,
8-spored. Ascospores 60-75 X 2.0 zm, nonseptate, narrowly sheathed.

On fallen leaves of Quercus rubra, autumn, on leaves of the previous
year, Concord, Massachusetts, USA. The dimensions of the hymenial
elements match Coccomyces coronatus, but the external aspect of
the two species is quite different. Coccomyces coronatus has larger
apothecia which open very irregularly, an orange disc, somewhat longer
asci and spores, and characteristic agglutinated paraphyses which form
an epithecium. Coccomyces farlowii can further be distinguished from
C. coronatus and C. dentatus by the presence of an excipulum and
a hymenium which remains exposed when dry.

Specimen examined (see also holotype, above): North America. USA: Concord, Mass.,
on Quercus rubra, D. Pfister, 17.1X.1978.

38. Coccomyces filicicola Speg. See C. dentatus.

39. Coccomyces fimbriatus (Schm. ex Fr.) Quelet, Euchirid. Fung.
338 (1886).
= [Phacidium fimbriatum Schm., Myc. Heft 1: 39 (1817)] ex
Fr., Syst. Mycol. 2(2): 578 (1823).

According to the original description the species occurred on leaves
of Populus and resembled C. coronatus, but with smaller fruitbodies
and a white (turning grey) hymenium. I am unable to locate any
original material of this species, and cannot characterize it from the
original description. It may be a synonym of C. tumidus, which occurs
on this host.

THE GENUS COCCOMYCES a0

40. Coccomyces foliicola (Dennis & Spooner) Sherwood, comb.
nov. Figure 24
= Coccomyces boydii A. L. Smith f. foliicola Dennis & Spooner,

Kew Bull. 32: 111 (1977).

Apothecia intraepidermal in bleached spots bounded by a line stroma
on dead leaves, circular or quadrangular, 0.7-1.0 mm diam, opening
along indistinct, paler preformed lines of dehiscence to expose the
yellowish hymenium. Covering stroma 25 um thick, of slightly carbon-
ized cells 3-5 zm diam, paler toward the periphery of the stroma,
without periphysoids. Basal stroma 10-15 jm thick, the cells 5-8
wm diam, slightly carbonized. Excipulum absent. Subhymenium
colorless, 15 jm thick. Paraphyses filiform, enlarged to 4.0 pm at

ff
/;

yf /
i! ff

/
/

A /}
ye

Fic. 24. Coccomyces foliicola:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, approximately natural size. Drawn from the holotype.

56 MARTHA A. SHERWOOD

the apex, cemented in a gel, forming an epithecium. Asci cylindrical,
short-stalked, 135-170 <x 6.5 um, J—, 8-spored. Ascospores 90-120
x 1.0 pm, nonseptate, not obviously sheathed.

On fallen leaves of Myrica faya, Azores. Dennis and Spooner (l.c.)
give somewhat shorter dimensions for the asci. The material which
I examined appeared mostly immature. In their comments they remark
‘““Coccomyces boydii was described on bark and twigs of Myrica gale
in Perthshire, with rather shorter ascospores. Until there is a critical
revision of Coccomyces it seems best to treat the Azores collection
as a foliicolous form of this.”” Coccomyces foliicola differs from C.
boydii not only in ascospore dimensions, but also in the preformed
dehiscence mechanism of the stroma and the inflated paraphyses
forming an epithecium. It is close to C. leptosporus, but lacks an
excipulum. In general, a species of Coccomyces is less likely to grow
on leaves and twigs of the same host than it is to grow on leaves
of ecologically similar but taxonomically unrelated hosts.

Specimen Examined: ATLANTIC ISLANDS. AZORES: on Myrica faya, Terceira de Macela,
Terceira, 26.3.1975, holotype of C. boydii f. foliicola (x).

41. Coccomyces heterophyllae Funk, Canad. J. Bot. 45: 2263 (1967)
[1968]. Figure 25
Apothecia intracortical, becoming erumpent, orbicular, polyhedral,
or slightly elongate, 0.5-1.1 mm diam, black, pulvinate, without an
obvious preformed dehiscence mechanism, splitting open irregularly
to expose the pale yellow hymenium, solitary or becoming confluent
but not immersed in swarms under a common black stromatic crust.
Covering layer 40-50 ym thick, of globose dark brown cells 3 pm
diam, barely carbonized on the exterior, without a defined periphysoi-
dal layer. Excipulum absent. Basal stroma poorly developed. Subhy-
menium 75-120 pm thick, of loosely consolidated somewhat gelatin-
ous, nodulose hyphae. Paraphyses filiform, 1.0 pm broad, circinate,
sometimes branched. Asci clavate, long-stalked, 75-140 x 8-10 pm,
8-spored; ascospores 19-27 * 1.5-2 4m (but see comments, below).
Pycnidia not produced.

On twigs of Tsuga heterophylla in the coastal forests of British
Columbia and Washington, spring and summer, possibly weakly
parasitic but more common as a secondary invader on branches killed
by other fungi, according to Funk (1967).

Funk (1967) reported that he was unable to find the species in
central British Columbia. I have collected two specimens in the
Cascade Mountains in Oregon which may represent this species but
differ significantly in their ascospore characters. One, on Tsuga
heterophylla from Wildcat Mt. (elev. 4000 ft) has ascospores 17.5-22
x 4.5-5.5 um. The other, on Tsuga mertensiana from the same area

THE GENUS COCCOMYCES 57

Aa
Ce:
dead:

6,

Ve.
os ht |
YAPX b

Wi gi

t"

Ham |,
i al J

Fic. 25. Coccomyces heterophyllae:—a. cross section of margin, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype

(elev. 5200 ft) has ascospores 25-35 & 2.5-3 um. Additional collections
and cultural studies would be desirable to determine whether or not
these represent distinct species, infraspecific taxa, or ecological
variants.

Specimens examined: NORTH AMERICA. CANADA. British Columbia: on T'suga heterophyl-
la, Courtenay, Victoria Island, A. T. Foster, May 1, 1962, holotype of C. heterophyllae

(DavFP 17300). usa. Oregon: Linn Co., Wildcat Mt., on Tsuga heterophylla, Sherwood
& Pike, 18.VIII.1978 (FH); on T. mertensiana, ibid. (rn).

42. Coccomyces hiemalis Higgins, Science n.s. 37: 637-638 (1913).
= Higginsia hiemalis (Higgins) Nannf., Nova Acta Regiae Soc.

Sci. Upsal. IV, 8(2): 175 (1932).

Blumeriella jaapii (Rehm) Arx, Phytopathol. Z. 42: 164 (1961).

58 MARTHA A. SHERWOOD

According to Nannfeldt (1932), Arx (1961) and Korf (1973), this
is a member of the Dermateaceae unrelated to Coccomyces.

Specimen examined: EUROPE. GERMANY: Rehm, Ascomyceten 1727, auf Blattern von
Prunus padus, Oldesloe, Schleswig-Holstein, V/1907, leg. O. Jaap, isotype of
Pseudopeziza jaapii (FH).

43. Coccomyces insignis Karst., Meddel. Soc. Flora Fauna Fennica
16: 16 (1888).

Apothecia scattered, oblong-ellipsoid, at first closed, subcuticular,
black, fleshy, opening by a longitudinal slit, 1 mm diam, the disc
pallid. Asci 215-240 x 21-30 pm, clavate, long-stalked, uniformly
thin-walled; ascospores 8, 180 xX 1.5-2.0 (—3.0 fide Karsten) pm,
colorless, very prominently sheathed, nonseptate. Paraphyses filiform,
branched, noncircinate, 1.5-2.0 1m diam.

On leaves of Carex pauciflora, Mustiala, Finland. As the species
is known only from a fragmentary collection containing a single
apothecium I am unable to supply microanatomical details of the
construction of the apothecial stroma. The species is clearly rhytis-
mataceous, but its generic placement is unclear. The subcuticular,
fleshy ascocarps opening by a longitudinal slit are reminiscent of
certain forms of Coccomyces tumidus. It would be desirable to recollect
C. insignis.

Specimen examined: EUROPE. FINLAND: Tavastia australis, Tammela, Salois (= Mustiala
N.) in Caric. paucifl., 1X.1887, leg. & det. P. A. Karsten, authentic (H, herb. Karsten).

44. Coccomyces irretitus Sherwood, spec. nov. Figure 26

Ascocarpi primo immersi, dein erumpentes, orbiculati vel irregulariter elongati, 1-1.8
um diam, per lacinias irregulariter aperientes, in macula pallida non insidentes. Margo
superior stromatica 50 wm crassa, ex hyphis intertextis carbonaceis constata. Periphysoi-
dei hyalini, reticulati, 200 1m longi. Margo inferior stromatica 20 1m crassa, ex hyphis
brunneis constata. Excipulum nullum. Paraphyses filiformes, haud circinatae, apice
non incrassatae. Asci 130-140 x 11 wm, clavati, pedicellati, 8-spori, in iodo non
caerulescentes; sporis filliformibus, continuis, in tunica gelatinosa inclusis, 45-55 x
1.0 pm. In ramis emortuis Pini, Abietis, Piceae et Laricis, Amer. Bor.

Holotypus: FH-Phacidiales Exsiccati 29, on bark of Picea sp., Seal
Cove Road, Acadia National Park, Mt. Desert Island, Maine,
16.VI.1979, leg. M. A. Sherwood.

Etymology: irretitus (L), surrounded by a net, referring to the
distinctive reticulate periphysoids.

Apothecia at first immersed, intracortical, becoming erumpent and
at length nearly superficial, orbicular to irregularly elongate, 1-1.8
mm diam, black, pulvinate, without a preformed dehiscence mecha-
nism, splitting open irregularly by teeth, not surrounded by a bleached
area of bark or developing beneath a common stromatic crust. Covering
layer 50 wm thick, of heavily carbonized, disintegrating cells mixed

THE GENUS COCCOMYCES 59

. 26. Coccomyces irretitus:—a. cross section of apothecium, *225.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, 7.5. Drawn from rH-Thaxter 252.

with host tissue, lined on the inside with 200 1m of branched and
netlike interwoven colorless periphysoids immersed in a gel, merging
at the periphery of the stroma with vertically oriented, nodulose hyphae
2-3 wm diam, intermingled with a few scattered crystals. Lower stroma
20 pm thick, of brown, noncarbonized cells 2-3 wm diam. Subhymen-
ium colorless, 20 pm thick. Asci clavate, long-stalked, 130-140 x
11 wm, thin-walled, J—, 8-spored. Paraphyses filiform, weakly circinate,
not enlarged above, colorless, not forming an epithecium. Ascospores
45-55 * 1.0 wm, distinctly sheathed.

On corticate branches of a variety of conifers, eastern North America,
frequent. Not to be confused with Therrya fuckelii, which occurs
on the same hosts, but has septate ascospores and a brown epithecium.
The hymenial elements of Coccomyces pseudotsugae are similar, but
the structure of the margin is entirely different. This is almost certainly

60 MARTHA A. SHERWOOD

the same fungus to which Reid and Cain (1961) refer when they
mention a specimen (#2082) collected by Wehmeyer in Colchester
Co., Nova Scotia, identified as C. cembrae but representing a distinct,
undescribed species of Coccomyces which they declined to describe
from the scanty material they had on hand.

Specimens examined (see also holotype, above): NORTH AMERICA. CANADA. Nova Scotia:
Wehmeyer 1004, on Larix laricina, upper Brookside, Colchester Co., 13.VII. 1931 (mMicw).
usa. New Hampshire: on Pinus resinosa?, Shelburne, VI.1894, Thaxter (FH); ibid.,
VI.1899. R.T. #252 (FH); on Abies nigra, Lake Pennigewasset, VII.1895, Farlow (Fn);
on Picea negra vel Pinus resinosa, Shelburne, VI.1894 (FH); on Picea, Tuckerman’s
ravine, Sherwood & Kneiper, VII.1978 (FH). North Carolina: Indian Gap, 10.VI.1927,
on Picea mariana (osc 12012).

45. Coccomyces javanicus Sacc. & Sydow, Syll. Fung. 16: 789 (1902).

= Coccomyces rhododendri Rac., Paras. Algen Pilze Javas 3: 19

(1900), non C. rhododendri (Schw.) Sacc. nec C. rhododendri
Rehm.

I could locate no type or authentic material of this species, which
occurred on leaves of Rhododendron javanicum in Java, causing
bleached lesions bordered by a reddish discolored zone, and having
asci 125-140 X 6-7 um, with spores 1 wm broad. It may be an older
name for C. vilis. The question of the synonymy and host range
of parasitic Asian species of Coccomyces needs further investigation.

46. Coccomyces juniperi [Karst.] Karst., Bidrag Kannedom Finlands
Natur Folk 19: 254 (1871). Figure 27
[= Phacidium pini (Alb. & Schw. ex Fr.) Fr. f. juniperi Karst.,
Fungi Fenniae 339 (1866), nom. nud.]
[= Phacidium juniperi [Karst.] Karst., Not. Soc. Fauna Flora Fenn.
11: 257 (1870), nom. nud.].
= Clithris juniperi ([{Karst.] ex Karst.) Rehm in Rabenh., Krypt.-
Fl. ed. 2, 1(3): 102 (1888).
= Colpoma juniperi ([Karst.] ex Karst.) Dennis, Kew Bull.
1957: 401 (1957).
= Godronia juniperi Rostr., Meddel. Grénland 3: 611 (1891).°
Apothecia intracortical, becoming erumpent, 1-3 mm broad, orbicu-
lar to irregular in outline, never linear, at most 1.5 times as long
as broad, brownish-black, without a preformed dehiscence mechanism,
splitting open irregularly to expose the hymenium. Covering layer
90-100 pm thick, with a heavy, black, carbonized crust and internal
layer of loosely interwoven brown hyphae 5 wm diam, oriented
perpendicular to the surface. Lower stroma absent, replaced by a
subiculum of dark hyphae 3 wm diam, merging above into a compact

6p : ° ‘ a
For other synonyms reported in the literature, see discussion below.

THE GENUS COCCOMYCES 61

Fic. 27. Coccomyces juniperi:—a. cross section of apothecium, *225.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, *7.5. (a) and (b) drawn from Rehm, Ascomyceten 272b (MICH);
(c) drawn from Poelt specimen (GRZ).

matrix of colorless hyphae 3-5 wm diam. Subhymenium colorless,
40-70 wm thick. Paraphyses filiform, colorless, circinate, often
branched, 1.5 wm diam. Asci stalked, clavate, 75-120 xX 7.5-9 wm,
uniformly thin-walled, J—, 8-spored. Ascospores 40-50 x 1-1.5 (—2)
um, tapering slightly below, nonseptate, distinctly sheathed.

On corticate twigs of Juniperus communis subsp. nana, Europe.
Dennis (1958) noted that the American material to which this name
had been applied had larger ascospores than the European material,
and questioned the synonymy, widely accepted in the literature, of
Colpoma juniperinum Cke. and Pk. with Coccomyces juniperi. Reports
of C. juniperi from North America (Ellis & Everhart, 1892; U.S.D.A.
Host Index, 1960) appear to be based either on C. petersii (q.v.) or
Colpoma species. Coccomyces juniperi is distinguished from C.

62 MARTHA A. SHERWOOD

petersii by its consistently larger apothecia, thicker covering layer,
and smaller ascospores, and from Colpoma spp., which occur on
Juniperus (probably Colpoma crispum (Pers. ex Fr.) Sacc.), by the
lack of a preformed longitudinal slit bordered by distinct lip cells.
Notwithstanding that the asci, ascospores, and paraphyses of Colpoma
crispum and Coccomyces juniperi are similar, there is ample evidence
against synonymizing the two species, as was done by Holm and
Holm (1977). The synonymy with Godronia juniperi, also suggested
by Holm and Holm is more plausible. I have not examined the type
of that species.

Specimens examined: EUROPE. FINLAND: Karsten, Fungi Fenniae 339, Merimasku, a
pa enbarr, isotype of Phacidium pini forma juniperi (FH); Inari-Lappland, Kevo, 196-,
leg. J. Poelt, (GRz). COUNTRY UNCERTAIN: Petrak, Mycotheca Carpatica 211, Wetin, VIII.1923
(rH). aAusTRIA: Rehm, Ascomyceten 272, on Juniperus nana, Tyrol, 8/1874 (FH, MICH).
iraLy: Mycotheca Italica 880, Riva-Valdobbia (Novara), Carestia, 1901 (FH).

47. Coccomyces kerriae Stewart, Phytopathology 7: 405 (1917).
= Higginsia kerriae (Stewart) Nannf., Nova Acta Regiae Soc.
Sci. Upsal. ser. 4, 8(2): 175 (1932).
= Blumeriella kerriae (Stewart) Korf, Phytologia 21: 202
(1971).
This is a dermateaceous fungus unrelated to Coccomyces.

48. Coccomyces laciniatus (Alb. & Schw. ex Fr.) Quélet, Enchirid.
Fung. 337 (1886).
= [Peziza laciniata Alb. & Schw., Consp. Fung. Nisk. 333 (1805)]
ex Fr., Syst. Myc. 2(1): 113 (1822).
= Heterosphaeria laciniata (Alb. & Schw. ex Fr.) Fr., Summa
Veg. Scand. pars post. 365 (1849).
= Phacidium laciniatum (Alb. & Schw. ex Fr.) Fr., Elench.
Fung. 2: 132 (1832).
= Coccomyces coronatus (Schum. ex Fr.) de Not.* laciniatus
(Alb. & Schw. ex Fr.) Karst., Bidrag Kannedom Finlands
Natur Folk 19: 257 (1871).
= Discocainia laciniata (Alb. & Schw. ex Fr.) Torkels. &
Eckbl., Norw. J. Bot. 24: 137 (1977).

This species has been widely cited as a synonym of Coccomyces
coronatus. The original illustration, however, shows a stalked apothe-
cium, a character not found in Coccomyces. Recently Torkelsen and
Eckblad (1977) have suggested a different interpretation based on
specimens collected in Norway. Their specimens are a substipitate
discomycete on leaves and herbaceous debris, initially entirely en-
closed in a black carbonaceous matrix, opening at maturity by irregular
teeth, correctly referred to Discocainia J. Reid and Funk. This inter-

THE GENUS COCCOMYCES 63

pretation appears to be in accord with all the available information
about the original authors’ concept of the species, and should be
adopted.

The genus Discocainia has generally been considered (J. Reid &
Funk, 1966; Korf, 1973; Torkelsen and Eckblad, 1977) to be a member
of the Helotiales. Anatomically, however, both the type species, D.
treleasei (Sacc.) J. Reid and Funk, and D. laciniata resemble some
of the wood-inhabiting species of Coccomyces. Points of similarity
include the carbonized, stromatic covering layer splitting radially by
means of teeth to expose the hymenium, stalked, clavate, pointed,
J— asci of the type characteristic of the Rhytismataceae, and filiform-
clavate, sheathed, colorless spores. Tryblidiopsis Karst. is similar and
probably congeneric. A paper dealing with these fungi is in preparation.

Specimens examined: EUROPE. NORWAY: Rana, Lien N. for Rundfjell, Dissing &
Sivertsen, 7/9/1973; Rana, Lasken, Rundfjell, 7/9/1973, Dissing & Sivertsen (TRH).

49. Coccomyces ledi Rehm, Ann. Mycol. 11: 153 (1913). Figure 28

Apothecia intraepidermal, scattered on bleached areas not bounded
by a line, on living twigs, 0.4-0.6 mm diam, quadrate to hexagonal,
black, shining, opening along light-colored preformed lines of dehi-
scence. Cover layer 40 ym thick, carbonized above, lined with nearly
colorless pseudoparenchyma, with a layer of colorless cells flanking
the torn margin of the teeth. Basal stroma 10 jm thick, dark brown,
not heavily carbonized, the cells 5 wm diam. Subhymenium colorless,
10-20 ym thick. Paraphyses filiform, enlarged to 2 um at the apex,
slightly bent, colorless. Asci 100-130 x 13-17 wm, clavate, short-
stalked, J—, 8-spored. Ascospores 30-38 x 2-2.5 wm, tapered below,
with a broad, conspicuous sheath.

On living twigs of Ledum groenlandicum and L. palustre, late
summer and early fall, common in arctic and alpine localities. B.
Eriksson (1970) reports it from Finland, Norway, Sweden, and the
USSR. Judging from the material in GH it is commoner in the northern
part of its range.

Specimens examined: EUROPE. SWEDEN: Vasterbotten, leg. Sylvén, isotype of C. ledi
(s); Micromycetes Rariores selecti, Vasterbotten, Kuhlbachsliden, Aug. 1909, leg. Sylvén,
isotype of C. ledi (s); Vasterbotten, Vindeln, B. Eriksson, 24.X.1970 (UME). NORTH AMERICA.
CANADA. Manitoba: Nueltin L., 19.VII.1951 (Gu); Duck L., 10.VIII.1950 (Gu). Northwest
Territories: Hay River, Lewis 665 (GH); Athabaska R., Raup 1157 (cu). British Columbia:
Ft. Nelson, Raup & Correll 10719 (FH). usa. New Hampshire: King’s Ravine, White
Mts., VIII.1882 (FH); Mt. Chocoru, 3.1X. 1909 (FH); Shelburne, Farlow, LX.1885 (Fu):
Tuckerman’s Ravine, Sherwood, VII.1978 (FH). Oregon: Lane Co., II1.14.1979, Sherwood

(FH). ALASKA: Sitka, Bischoff 1865-66 (GH); Kuskowim R., Drury 2206, 22. VII.1949 (GH).
ASIA. JAPAN: Mt. Hakkoda, Honda, 25.VI.1950 (GH).

50. Coccomyces leptideus ([Fr. ex Fr.]) B. Erikss., Symb. Bot. Upsal.
19(4): 18 (1970). Figure 29

64 MARTHA A. SHERWOOD

Fic. 28. Coccomyces ledi:—a. cross section of apothecium, X375.—b. ascus, paraphysis, and
spores, ¥750.—c. habit sketch, 10. Drawn from Thaxter, Shelburne, NH, specimen (FH).

[Phacidium leptideum Fr., Obs. Mycol. 2: 312 (1818)] ex Fr.,
Syst. [Mycol. 2(2): 576 (1823).
= [Phacidium quadratum Schm. & Kunze, Myc. Hefte I: 32
(1817)].
= Coccomyces quadratus [Schm. & Kunze] Karst., Bidrag
Kannedom Finlands Natur Folk 19: 255 (1871).
= Lophodermium quadratum [Schm. & Kunze] Ces., Erb.
Crittogam. Ital. 885 (1862).

Apothecia scattered in bleached spots on living or recently killed
twigs, quadrangular to hexagonal, black, shining, 0.5-1.0 mm diam,
opening regularly by teeth along preformed lines of thin-walled, lightly
pigmented cells. Covering layer 20-25 jm thick, heavily carbonized,
the cells globose, 5-7 wm diam; lower stroma 20-25 pm thick,

THE GENUS COCCOMYCES 65

Fic. 29. Coccomyces leptideus:—a. cross section of apothecium, X375.—b. asci, paraphysis, and
spores, X750.—c. habit sketch, X7.5. Drawn from Muller 19.9.1973 (21)

constructed like the covering layer. Subhymenium colorless, 20 pm
thick. Asci clavate, long-stalked, 115-140 x 12-15 ym, either 4-spored,
with spores 60-90 X 4(—5) pm, or 8-spored, with spores 60-90 x
3.0-3.5 jm; spores prominently sheathed, nonseptate. Paraphyses
filiform, enlarged gradually to 3.0-3.5 ym at the apex, curved but
not circinate. Pycnidia absent.

On living and recently killed twigs of Vaccinium myrtillus, V.
vitisidaei, Rhododendron macrophyllum, and Gaultheria shallon, sum-
mer and fall, Europe and western North America. Common. The
ascocarps persist for a long time after sporulation has ceased and
are often collected in an effete condition. Eriksson (1970) described
the asci as being 4-spored; the material I examined had predominantly
8-spored asci. A collection on stems of Oxalis from Colombia (Ny-Co

66 MARTHA A. SHERWOOD

4190) agrees in all morphological particulars with typical C. leptideus,
but may be distinct, as the species is normally quite host-specific.
Specimens with the characters of C. leptideus on leaves from tropical
highland areas (TNs-F-225572, Mt. Wilhelm, New Guinea; Ny-Co 501,
on petiole of Ericaceae, Colombia, Cundinamarca, Dumont et al.)
may represent large-spored forms of C. dentatus.

There is another species of Coccomyces, Phacidium gaultheriae
Dearn. which occurs on living twigs of Gaultheria shallon in North
America. It is distinguished from C. leptideus by its shorter, broader
ascospores 19-22 x 6-6.5 um. Frank Di Cosmo, who called this taxon
to my attention, is preparing to publish a rediscription of it.

Specimens examined: EUROPE. SWEDEN: Fries, Scleromycetes Sueciae 98, isotype of
Phacidium leptideum (Fu); Uppland, Vaksala, B. Eriksson 691 (ups); on V. myrtillus,
Vasterbotten, 11.VII.1974, Eriksson 403c (ups); Smaland, Femsjo (UME 25386); Eriksson
2282 (ups); Lycksele Lappmark, B. Eriksson 780 (ups); Vasterbotten, Umea, Laestadius,
11.VI.1901 (UME 25385); Vasterbotten, Bygdea (umME 25381); Dalarna, Avesta, Terrier
416 (zr). swirzERLAND: Ct. Valais, Terrier, 21.5.1944 (zr); Ct. Wallis, Aletschreservat,
Muller, 19.9.1973 (zr); Ct. Wallis, Zermatt, Terrier, 31.VII.1938 (zr); Ct. Graubiinden,
18.9.1963, on V. uliginosum (zr). tracy: ? Val Marobia, Alp. Giumello, 15.VII.1937,
Terrier (zt); Erbar. Crittog. Ital. 885, Valdobbia, Valsesia, Carestia, 1861 (FH). NORTH
AMERICA, USA. Oregon: Yachats, on Gaultheria shallon, Sherwood, 10.VIII.1978 (Fx);
Lane Co., on R. macrophyllum, Sherwood, III.1979 (FH); on Rhododendron sp.,

Rhododendron, 22.V.1930 (osc 27,997). Washington: Langley, on G. shallon, I. 1923
(osc 9962).

51. Coccomyces leptosporus Speg., Bol. Soc. Cien. Argentina 19:
188 (1885). Figure 30
Apothecia intraepidermal, scattered in bleached spots bounded by
a delicate black line on dead coriaceous leaves, immersed, becoming
erumpent, orbicular, 0.3-0.5 mm diam, opening irregularly by 3-5
teeth, without a preformed dehiscence mechanism, the disc greyish-
yellow when fresh, drying yellow, remaining exposed when dry.
Covering layer 10 pm thick, devoid of internal differentiation, heavily
carbonized. Lower stroma 10 ym thick, heavily carbonized. Proper
exciple well-developed, 35 ym thick, of closely septate hyphae 3 um
diam, inflated and dark brown above, visible from above in dried
material as a narrow black rim around the hymenium. Subhymenium
colorless, 25 wm thick. Paraphyses filiform, abruptly enlarged to 2.5-4
wm at the apex, cemented in a gel and forming an obvious epithecium.
Asci cylindrical, short-stalked, 90-105 * 5-6 1m, 8-spored; ascospores
nearly as long as the asci, 0.8-1.0 1m broad, nonseptate, not obviously
sheathed.

On dead coriaceous leaves at low and middle elevations, tropical
America, common. The identity of the host is unfortunately unknown
for most of the specimens but it appears that most of them are on
the same host plant. A number of species of Coccomyces with
cylindrical asci and ascospores less than 1 4m diam occur on leaves

THE GENUS COCCOMYCES 67

Fic. 30. Coccomyces leptosporus:—a. cross section of apothecium, X375.—b. asci, paraphyses,
and spores, X750,—c. habit sketch, 7.5. Drawn from Ny-Co 5246,

in tropical America, including C. spegazzinii, C. tesselatus, C. antil-
larum, C. concolor, C. limitatus, and C. venezuelae. I have chosen
to separate them on rather fine morphological grounds partly because
it appears that the morphological distinctions are correlated with host

preference.

Specimens examined: CENTRAL AMERICA, PANAMA: Prov. Panama, Dumont et al.,
11.VI1.1975 (Ny-Pa 14); ibid., 30.VI.1975 (Ny-Pa 1582). CARIBBEAN. PUERTO RICO: Luquillo
Exp. Forest, on Dacryodes excelsa, Haines & Korf, 6.V1I.1970 (Nys). GUADELOUPE: Grand
Etang, 18.VII.1973, Pfister 527 (rH). SouUTH AMERICA. VENEZUELA: Puerto Zamuro, Gaillard
26 (ec); Sydow, Fungi Venezuelani 295, Fk] Limon, D.F. (z1, ru); Petrak, Mycotheca
Generalis 725, El Limon (zr, rH); Edo. Aragua, Dumont et al. (Ny-Ve 1208, 1186,
1145); Colonia Tovar, D.F., Dumont et al. (Ny-Ve 6609); Edo. Merida, Dumont et
al. (Ny-Ve 2776); Edo. Sucre, ibid. (Nv-Ve 4429): Edo. Bolivar, ibid. (Ny-Ve 7085). COLOMBIA:
Cundinamarca, Dumont et al. (Ny-Co 4427, 5246, 2270); Antioquia, ibid. (Ny-Co 6086,
1775, 6204, 1630); Boyaca, ibid. (Ny-Co 7969). peru: Dpto. Junin, Dumont et al. (Ny-Pe
308); Dpto., Loreto, ibid. (Ny-Pe 996). BoLivia: Prov. Nov. Jungos, Dpto. La Paz, R.

68 MARTHA A. SHERWOOD

Singer B1431 (k). Brasit: S Lauracea, Guarapi, leg. Balansa 3861, holotype of C.
leptosporus (LPs 28182).

52. Coccomyces limitatus (Berk. & Curt.) Sacc., Syll. Fung. 8: 747
(1889). Figure 31
= Phacidium limitatum Berk. & Curt., J. Linnean Soc. (London)

10: 371 (1868).

Apothecia intraepidermal on bleached spots bounded by a black
line on dead leaves, triangular or quadrangular, black, shining, with
a preformed dehiscence mechanism consisting of faint lines of light
colored, thin-walled cells, opening by 3-4 teeth to expose the yellow
hymenium, 0.5-0.8 mm diam. Covering layer and basal stroma 10-15
ym thick, of heavily carbonized cells 5 um diam. Periphysoids and
excipulum absent. Subhymenium colorless, 30 1m thick, separated

Fic. 31. Coccomyces limitatus:—a. cross section of apothecium, *375.—b. asci, paraphysis, and
spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

THE GENUS COCCOMYCES 69

from the basal stroma by a loose subiculum of colorless hyphae
intermingled with crystals. Asci cylindrical, short-stalked, 90-110 x
5.0-5.6 wm, 8-spored. Ascospores nearly as long as the asci, 0.8-1.0
wm diam, obscurely sheathed. Paraphyses 2-4 1m broad along their
entire length, not agglutinated or forming an epithecium.

On dead leaves of Clusia and unidentified plants, at low and middle
elevations, tropical America, generally on material in a more advanced
state of decay than Coccomyces clusiae and C. leptosporus. A specimen
collected on Medinilla speciosa in Java (Cup-Sa 344) appeared identical
morphologically to the South American specimens.

Specimens examined: NORTH AMERICA, MEXICO: Oaxaca, Dumont, 25.VIHI.1967 (cup-Me
317). CENTRAL AMERICA. PANAMA: Altos de Pacora, Prov. Panama, Dumont et al., 30.VI.1975
(NY-Pa 1595). CARIBBEAN. CUBA: “Cuba, C. Wright” holotype of Phacidium limitatum
(kK); Fungi Cubensis Wrightiani 714, Cobre, Dec. (FH). SOUTH AMERICA, VENEZUELA: on
Clusia, Rancho Grande, Aragua, Chardon & Whetzel (cup-vz 2798); Rancho Grande.

Aragua, Dumont (Ccup-vz 4271, 4337); Maracay, Edo. Aragua, Dumont et al., 13.VII.1971
(Ny-Ve 2112). cotomBia: Valle de Cauca, Dumont & Molina, 30.VIII.1976 (Ny-Co 7552).

53. Coccomyces lutescens Higgins, Amer. J. Bot. 1; 166 (1914),
= Blumeriella jaapii (Rehm) Arx.
For a discussion of this species see under C. hiemalis.

54. Coccomyces macer Karst. See Karstenia macer.

55. Coccomyces maritimus (Rolland) E. Muller, Rev. Mycol. N.S.
27: 71 (1962).
= Stictis maritima Rolland, Bull. Soc. Mycol. France 14: 84
(1898).
= Lasiostictis fimbriata (Schw.) Baumler, Ann. K. K. Naturh.
Hofmus. Wien 16: 67 (1901).
The morphology and taxonomic position of this species are discussed

by Sherwood (1974) and DiCosmo (1979).

56. Coccomyces martinae Hansf., Proc. Linn. Soc. N.S.W. 81: 40
(1956).
= Stictis emarginata Cooke & Massee, Grevillea 18: 7 (1889).
= Propolis emarginata (Cooke & Massee) Sherw., Mycotaxon
5: 323 (1977).

This species, which is widespread and common on fallen leaves
of Eucalyptus, is a member of the Rhytismataceae and closely allied
to Coccomyces, differing principally in having subepidermal ascocarps
with a very reduced covering layer. Sherwood (1977b), who provided
a redescription and illustration of S. emarginata, referred the species
to Propolis (Fr.) Cda for nomenclatural reasons; the majority of authors
use Naemacyclus Fckl for subepidermal filiform-spored rhytismata-

70 MARTHA A. SHERWOOD

ceous fungi without a dark stromatized covering layer. California Fungi
430 was distributed as C. delta, a misidentification.

Specimens examined: NORTH AMERICA. USA. California: California Fungi 430 (FH). souUTH
AMERICA. BRASIL: Minas Geraes, Hodges, 28.2.1974 (im1 186648B). asia. INDIA: Thekkaty,
on Eucalyptus grandis, Hodges, 17.4.1975 (imi 194560c). PACIFIC ISLANDS. HAWAII: Maui,

10.11.1976, Hodges (im1 209218c). AUSTRALASIA. AUSTRALIA: with Phoma eucalyptidea,
Brisbane, n.d., leg. Martin, holotype of Coccomyces martinae (k).

57. Coccomyces maydis Rehm. See Karstenia maydis.
58. Coccomyces memecycli Sydow. See C. vilis.

59. Coccomyces minutissimum (Auersw.) Sacc. ex Gola, Atti Acad.
Sci. Veneto Trent. Istriana 21, suppl. 1: 129 (1930).
= Naevia minutissima (Auersw.) Rehm in Rabenh., Krypt.-FI.
ed. 2, 1(3): 138 (1888).
The above combination was published, probably inadvertently, in
the catalogue of the Saccardo Herbarium. The species is dermateaceous.

60. Coccomyces monticola Sherwood, spec. nov. Figure 32

Ascocarpi primo immersi, dein erumpentes, orbiculati, 0.5-0.8 mm diam, per lacinias
4-5 irregulariter aperientes, in macula pallida stromate obvallato insidentes. Margo
superior stromatica 20 ym crassa, ex hyphis intertextis carbonaceis constata. Periphysoi-
dei nulli. Margo inferior stromatica 20 jm crassa, ex hyphis intertextis carbonaceis
constata. Excipulum brunneum. Paraphyses filiformes, apice ad 2.5 jm incrassatae,
achromae, in epithecio gelatinoso non inclusae. Asci 150-175 x 8-9 wm, cylindrici,
haud pedicellati, 8-spori, in iodo non caerulescentes; sporis filiformibus, continuis,
in tunica gelatinosa inclusis, 90-100 x 1.5-2 ym. In foliis dejectis Ericaceae, Amer.
Austral.

Holotypus: Ny-Co 5065b, on Ericaceae leaves, along the Sogamoso-
Aguazul road, at a point ca. 38 km from the intersection with the
Sogamoso-Aquitania rd., Dpto. Boyaca, Colombia, elev. ca. 9000 ft,
K. P. Dumont, S. E. Carpenter, M. A. Sherwood & L. A. Molina,
13.V1I.1976. Isotypus: COL.

Etymology: monticola (L), inhabitor of mountains, as the species
grows at high elevations.

Apothecia at first immersed, intraepidermal, orbicular, black, shin-
ing, 0.5-0.8 mm diam, opening by 4-5 irregular teeth to expose the
orange disc, remaining open when dry, scattered in bleached spots
bounded by a black line on dead leaves. Covering layer and lower
stroma ca. 20 wm thick, black, carbonized, devoid of internal differen-
tiation. Proper excipulum well-developed, consisting of 30 pm of
closely agglutinated septate colorless hyphae 3-4 zm diam, short-
celled, with brown apices. Subhymenium colorless, 30 jm _ thick,
separated from the basal stroma by a subiculum of colorless hyphae

THE GENUS COCCOMYCES vial

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Fic. 32. Coccomyces monticola:—a. cross section of apothecium, *375.—b. detail of asci,
paraphyses, and spores, *750.—c. habit sketch, X7.5. Drawn from NyY-Co 5065b.

2.0 wm diam. Paraphyses filiform, colorless, enlarged to 2.5 um at
the apex, bent but not circinate. Asci cylindrical, short-stalked, 150-175
x 8-9 um, J—, 8-spored. Ascospores 90-100 xX 1.5-2.0 um, with a
distinct, narrow gelatinous sheath.

On dead leaves at high elevations in Colombia. The host of the
holotype is a species of Vaccinium; the hosts of the other specimens
are unknown, but may well be ericaceous. The long asci and rather
broader ascospores distinguish Coccomyces monticola from members
of the C. leptosporus complex.

Specimens examined (see also holotype, above): SOUTH AMERICA, COLOMBIA: ca. 62 km
from Ansermanuevo on the Ansermanuevo-San José de Palomar road, boundary Valle
de Cauca and Choco Dptos., elev. 2100 m, Dumont, Molina & Forrero, 25. VIII.

1976 (Nny-Co 7297); km 16-17 from Mosquera on the Mosquera-La Mesa rd., Dpto.
Cundinamarca, elev. 9300 ft, Dumont, Carpenter & Sherwood, 5.V1I.1976 (Ny-Co 4189);

72 MARTHA A. SHERWOOD

km 112 from Pasto, Pasto-Mocoa rd., Intendencia Putumayo, elev. 8000 ft, Dumont,
Buritica, Molina & Luteyn, 28.1.1976 (Ny-Co 3822, 3832).

61. Coccomyces musae (Lév.) Sacc. See C. clusiae.
62. Coccomyces mussaendae Sawada. See Biostictis tjibodensis.

63. Coccomyces palmicola Sherwood, spec. nov. Figure 33

Ascocarpi primo immersi, trigoni vel tetragoni, 0.3-0.6 mm diam, per lacinias 3-4
aperientes, in macula pallida stromate obvallato insidentes. Margo superior stromatica
50 ym crassa, ex hyphis intertextis carbonaceis constata. Margo inferior stromatica
15 um crassa, ex hyphis intertextis carbonaceis constata. Excipulum nullum. Paraphyses
filiformes, simplices vel ramosae, apice ad 2.0-2.5 jum incrassatae, circinatae, in epithecio
gelatinoso non inclusae. Asci 110-130 x 7-9 ym, cylindrici, haud pedicellati, 8-spori,
in iodo non caerulescentes; sporis filiformibus, continuis, in tunica gelatinosa inclusis,
75-100 X 1.5 pm. In foliis dejectis palmae, Panama.

Y PAWS

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Fic. 33. Coccomyces palmicola:—a. cross section of apothecium, X375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

THE GENUS COCCOMYCES 73

Holotypus: Ny, Fungi of Panama 49, on frond of indet. palm, summit
of Cerro Jefe, ca. 23 km N. of Pan American Hwy., Prov. Panama.
Elev. ca. 3000 ft, K. P. Dumont, S. E. & S. M. Carpenter, S. Mori,
LL. VEA9TS.

Etymology: palmicola (L), growing on palms, from the host.

Apothecia intraepidermal, at first immersed, black, shining, triangu-
lar or quadrate, 0.3-0.6 mm diam, without an obvious preformed
dehiscence mechanism, opening by 3-4 teeth to expose the greyish
(when dry) hymenium, scattered in prominent bleached spots on dead
palm leaves. Covering stroma 50 um thick, heavily carbonized. Basal
stroma 15 wm thick, heavily carbonized, the cells 4-7 jm diam.
Subhymenium colorless, 30 wm thick. Excipulum absent. Paraphyses
colorless, enlarged to 2.0-2.5 wm at the apex, circinate, sometimes
branched, not immersed in a gel. Asci cylindrical, short-stalked,
110-130 x 7-9 wm, somewhat truncate, J—, 8-spored. Ascospores
75-100 x 1.5 wm, distinctly sheathed, the sheath appearing as a
refractive cap at the apex of the spore.

On palm fronds, Panama. Coccomyces cocoes, which occurs on
palm fronds in Africa, differs in having a distinct excipulum and
inflated, noncircinate paraphyses.

Specimens examined: see holotype, above.

64. Coccomyces pampeanus Speg., Contrib. Estud. Fl. Sierra Ventana
85 (1896). Figure 34
= Coccomyces pampeanus Speg. var. chilensis Speg., Fungi

Chilensis 132 (1910).

Apothecia intraepidermal, immersed, triangular to quadrate, black,
shining, 0.3-0.6 mm diam, without an obvious preformed dehiscence
mechanism, opening by 3-4 teeth, scattered on inconspicuous bleached
areas not bounded by a line on dead leaves. Covering layer 10-15
um thick, undifferentiated, of carbonized cells 2.5 tm diam. Basal
stroma consisting of | layer of carbonized cells 5 wm diam. Subhyme-
nium colorless, 5 wm thick. Excipulum absent. Paraphyses filiform,
enlarged to 2.5 um at the apex, not imbedded in a gel. Asci clavate,
short-stalked, 80-110 * 11-14 pm, 4- or 8-spored, J—. Ascospores 50-60
x 2-3 pm, tapered below, distinctly sheathed, becoming 1|-septate.

On dead leaves of Eryngium paniculatum, Argentina and Chile,
Musa, and an unidentified bromeliad. The type of Coccomyces
pampeanus is immature. Spegazzini erected his var. chilensis on the
grounds that the Chilean material had longer, broader spores. As the
two types are on the same host and do not differ in any other respect
I consider that the Chilean material represents the species in its fully
mature state. The specimens on Musa from Guadeloupe agree in all
morphological particulars with the specimens on Eryngium (Umbelli-

74 MARTHA A. SHERWOOD

Fic. 34. Coccomyces pampeanus:—a. cross section of apothecium, X375.—b. ascus, paraphysis,
and spores, X750.—c. habit sketch, X7.5. Drawn from Gaudeloupe 956 (FH).

ferae). The specimen on Bromeliaceae from Colombia differs in having
4-spored asci. If these indeed all represent the same species, this
is an interesting example of a Coccomyces occurring on ecologically
similar but taxonomically unrelated hosts, Eryngium having the texture,
growth form, and appearance of a terrestrial bromeliad.

Specimens examined: CARIBBEAN. GUADELOUPE: Morne a Louis des Mamelles, Pfister
et al., 5.1.1974 (FH); Camp Jacob, Pfister et al., 7.1.1974 (FH). SOUTH AMERICA, VENEZUELA:
on Heliconia, Edo. Sucre, Dumont et al., 13. VII.1972 (Ny-Ve 4987). coLoMBIA: Mosquera-
La Mesa rd., Dpto. Cundinamarca, Dumont et al., 5.V1I.1976, on indet. bromeliad (Ny-Co
4188). ARGENTINA: S/ Eryngium paniculatum, Buenos Aires, Sierra de la Ventana, XI.1895,
leg. Spegazzini, holotype of C. pampeanus (Lips 12655). CHILE: Cerro Caracol de
Concepcion, 1.1909, leg. C. Spegazzini, holotype of C. pampeanus var. chilensis (LPs
28178).

THE GENUS COCCOMYCES TS

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Fic. 35. Coccomyces papillatus:—a. cross section of apothecium, *150.—b. asci, paraphyses,
and spores, X750.—c. detail of hyphae from interior matrix of stroma, X750.—d. habit sketch.
7.5. Drawn from the holotype.

65. Coccomyces papillatus Sherwood in Ahmad, Ascom. Pakistan
216 (1978). Figure 35
Ascocarps at first immersed, raising the overlying bark into promi-
nent pustules with a depressed center and small central papilla, at
length splitting open irregularly by teeth to expose the hymenium,
without any preformed dehiscence mechanism, not accompanied by
stromatic lines or bleached discoloration of the surrounding substrate.
Covering layer 50 ym thick, of heavily carbonized cells 5-8 pm diam,
devoid of internal differentiation. Lower stroma ca. 20 wm thick,
heavily carbonized, separated from the subhymenium by a very thick
(up to 500 pm) matrix of thick-walled, gelatinous hyphae 10 wm diam,
which eventually break down, leaving a mass of crystals. Subhymenium

76 MARTHA A. SHERWOOD

colorless, 30 4m thick. Paraphyses filiform, circinate, not cemented
in a gel. Asci clavate, long-stalked, 95-120 x 9-11 um, J—, 8-spored.
Ascospores cylindrical, tapering slightly below, 29-41 x 2.0 wm,
narrowly sheathed.

On smooth bark of twigs and small branches of Pinus wallichiana
(= P. excelsa), Pakistan, common, according to S. Ahmad (personal
communication). Very similar to Coccomyces strobi on Pinus strobus
in eastern North America, differing principally in having larger
apothecia with a much thicker matrix of hyphae beneath the subhy-
menium. I have collected C. strobi on cultivated P. wallichiana in
Boston; the apothecia were of the same dimensions as those of this
species on P. strobus. Since neither of these two species of Coccomyces
is known to occur in Europe and their respective ranges are conse-
quently quite isolated from one another, I have chosen to regard
them as distinct species, although the morphological differences are
slight.

Specimens examined: Asia. PAKISTAN: Ahmad 19341, Loon Bagla, Muzaffarabad,
26.7.1963 (LAH); Ahmad 27504, Swat, Kalam, 20.8.1952 (Lan); Dunga Gali, 22.7.1965

(LAH); ex herb. S. Ahmad 4356, Kaghan Valley, Shogran, holotype of C. papillatus
(BPI).

66. Coccomyces parvulus Sherwood, spec. nov. Figure 36

Ascocarpi primo immersi, orbiculati vel irregulariter elongati, 0.3-0.8 mm diam,
per lacinias irregulariter aperientes, in macula pallida non insidentes. Margo superior
stromatica 75-100 jm crassa, ex hyphis intertextis carbonacea constata. Periphysoidei
nulli. Margo inferior stromatica nulla. Excipulum nullum. Paraphyses filiformes,
ramosae, Circinatae, in epithecio gelatinoso non insidentes. Asci 120-140 x 8-9 um,
clavati, pedicellati, in iodo non caerulescentes, 8-spori; sporis cylindricis, continuis,
in tunica gelatinosa inclusis, 25-30 = 1.0-1.5 ym. In ligno coniferarum, Amer. Bor.
Occident.

Holotypus: FH, Reliquiae Suksdorfiana II] (unnumbered, sub Clith-
ris crispa), on old conifer wood, Washington, Yakima Co., Mt. Adams,
Tune 27, 1883, W. N. S. 55.

Etymology: parvulus (L), small, insignificant, referring to the size
of the ascocarps (relative to other wood-inhabiting species of Cocco-
myces) and their inconspicuousness.

Ascocarps immersed in swarms beneath a common, coal black
stromatic crust on decorticated conifer wood, orbicular to elongate
but not linear, at most twice as long as broad, without a preformed
dehiscence mechanism, splitting open irregularly by teeth. Covering
stroma 75-100 wm thick, with a carbonized outer crust 20 itm diam,
and an inner portion of fleshy brown globose cells 5-8 um diam.
Lower stroma absent, replaced by a layer 100 ym thick of loosely
consolidated gelatinous hyphae 3-5 wm diam, resting directly on
disintegrating wood. Subhymenium 15 um thick, faintly brown.

THE GENUS COCCOMYCES

Fic. 36. Coccomyces parvulus:
spores, X750.—c. habit sketch, *7.5. Drawn from the holotype

a. cross section of margin, X225,—b. asci, paraphyses, and

Excipulum absent. Paraphyses 1.5 tm diam, branched, circinate,
forming an epithecium. Asci 120-140 x 8-9 wm, clavate, long-stalked,
8-spored; ascospores nonseptate, 25-30 * 1.0-1.5 jm, prominently
sheathed.

On decorticated wood of various conifers, probably common. The
fungus is easily overlooked in the field because the colonies resemble
charred wood and the tiny apothecia are extremely inconspicuous.
The specimens cited were found deposited in herbaria under the names
Clithris crispa, C. graphis, and C. morbida, all species which have
linear ascocarps opening by a definite longitudinal slit and usually
occur on corticate twigs.

Coccomyces parvulus is closely related to C. atactus, C. bipartitus,
and C. cembrae, differing from all of them in paraphysis and ascospore

78 MARTHA A. SHERWOOD

characters. Colpoma deusta Sherw., also occurring on conifer wood
in Oregon, is similar but has long-linear ascocarps.

Specimens examined (see also holotype, above): NORTH AMERICA. USA. Washington:
on Pinus sp., Crystal Ridge, Olympic Mts., elev. 4800 ft, A. H. Smith 14422, 14423,
17.VI.1939 (Micu). Oregon: Takilma, on Douglas fir, C. H. Kauffman, Dec. 1925 (micu);
on Abies, Mary’s Peak, 21.VIII.1975 (Herb. Sherwood 2020).

67. Coccomyces peltiformis (Preuss) Sacc. See C. tumidus.
68. Coccomyces pentagonus Kirschst. See C. dentatus.
69. Coccomyces petersii (Berk. & Curt.) Sherwood, comb. nov.

Figure 37
= Hysterium petersii Berk. & Curt., Grevillea 4: 13 (1875).

y,

ORI)

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JO
a

QA

ND

Pa SS

Fic. 37. Coccomyces petersii:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch, X7.5. Drawn from FH-Thaxter X.1902.

THE GENUS COCCOMYCES 79

= Lophodermium petersii (Berk. & Curt.) Sacc., Syll. Fung.
2: 795 (1883).
= Colpoma juniperinum Cke. & Pk., Bull. Buffalo Soc. Nat.
Sci. 3(2): 36 (1876).

Apothecia gregarious, at first immersed in nondiscolored bark,
orbicular to oblong, 0.8-1.5 mm diam, black, shining, without a
preformed dehiscence mechanism, splitting open irregularly by teeth
to expose the yellowish hymenium. Covering layer 100 ym thick,
without internal differentiation, of globose carbonized cells 3-7 xm
diam. Basal stroma reduced to a subiculum of dark brown hyphae
2.5-3.5 4m diam. Subhymenium colorless, 30 wm thick. Excipulum
absent. Paraphyses filiform, circinate, 1.5-2 wm broad. Asci clavate,
long-stalked, 85-110 x 8-10 ym, J—, 8-spored. Ascospores 45-60 x
1.5-2 wm, narrowly sheathed, nonseptate.

On thin bark covering large limbs of Juniperus virginiana, eastern
North America. This species has generally been considered in the
literature to by synonymous with the European Coccomyces juniperi
(Ellis and Everhart, 1892), although Dennis (1957) mentioned that
the North American type of Colpoma juniperinum had larger spores
than European specimens of Coccomyces juniperi. In addition to
differences in spore dimensions, non-overlapping geographical ranges,
and different host species, C. juniperi and C. petersii exhibit a number
of small but consistent morphological differences which suggest that
they are distinct taxa. Coccomyces petersii typically has smaller, flatter
apothecia less deeply immersed in the substrate, is more gregarious,
and occurs on trunks and large branches rather than on small twigs.

Specimens examined: NORTH AMERICA. USA: On Juniperus virginiana, F. A. Wolf, 1936
(FH); Waverley, Massachusetts, Oct. 1902, R. Thaxter (FH); Buffalo, New York, Clinton,
on J. virginiana (FH); Curtis Herb. 4016, Alabama, Peters 376, isotype of Hysterium
petersii (FH).

70. Coccomyces philippinus (Rehm) Sherwood, comb. nov.
Figure 38

= Coccomyces quadratus (Schm. & Kunze) Sacc. var. philippinus
Rehm, Leafl. Philipp. Bot. 8: 2926 (1916).

Apothecia immersed in bleached spots bounded by a black line
on dead leaves, intraepidermal, 0.5-1.0 mm diam, triangular to hexa-
gonal, black, shining, with a distinct preformed dehiscence mechanism
of pale colored, thin-walled cells, opening by teeth. Covering stroma
20 pm thick, heavily carbonized, bearing a few short periphysoids
on its inner face. Basal stroma 10 wm thick, heavily carbonized.
Subhymenium 20 ym thick, colorless, the cells 5 im diam. Excipulum
absent. Paraphyses filiform, inflated to 5.0 1m at the apex, not cemented
in a gel. Asci cylindrical, short-stalked, 100-125 x 5-6 itm, 8-spored;

80 MARTHA A. SHERWOOD

Fite taow

| Ld NY pin

Fic. 38. Coccomyces philippinus:—a. cross section of apothecium, X375.—b. ascus, paraphyses,
and spores, X750.—c. pyenidium, x750,—d. habit sketch, 7.5. Drawn from the holotype.

ascospores nearly as long as the asci, 1.0 »m diam, nonseptate, not
obviously sheathed.

Pycnidia numerous, occurring with the apothecia, 0.1-0.2 mm diam,
appearing as minute concolorous pustules bounded by a thin black
line. Covering layer absent; basal stroma incomplete, forming a ring
as seen from above. Conidiophores in a compact basal layer, 10-12
x 1.5 pm, simple. Conidia colorless, unicellular, 3-4 x 1.0 wm.

Ascospore dimensions, paraphyses type, and presence of pycnidia
all separate Coccomyces philippinus from C. quadratus (= C. lepti-
deus). It is rather close morphologically to C. limitatus, but that species
also does not produce pycnidia.

Specimen examined: ASIA. PHILIPPINES: Baker, Fungi Malayani 129, on Neolitsea, Mt.

Maquiling, near Los Banos, Prov. Laguna, June, 1914, isotype of C. philippinus (pc,
FH).

THE GENUS COCCOMYCES 81

71. Coccomyces piceae (Fuckel) Sacc., Syll. Fung. 8: 746 (1889).
= Phacidium piceae Fuckel, Jahrb. Nassauischen Vereins Naturk.
28-29: 51 (1875).
= Lophodermium piceae (Fuckel) Hohnel, Sitzungsber. Kai-
serl. Akad. Wiss., Math.-Naturwiss. Kl. Abt. 1, 126: 296
(1917).
This is a species of Lophodermium, according to Darker (1967).

72. Coccomyces pini ({[Alb. & Schw.] Schm. ex Fr.) Karst., Bidrag
Kannedom Finlands Natur Folk 19: 254 (1871).
= Phacidium pini [Alb. & Schw.] Schm. ex Fr., Syst. Mycol.
8(2): 573: (1823).
= Therrya pini ({Alb. & Schw.] Schm. ex Fr.) Hohnel, Sit-
zungber. Kaiserl. Akad. Wiss., Math.-Naturwiss. KI. Abt.
1, 121: 62 (1912).
For complete synonymy and a discussion of this and the following
species, see Reid and Cain (1961).

73. Coccomyces pini ([Alb. & Schw.] Schm. ex Fr.) Karst. var. fuckelii
Rehm in Rabenh., Krypt.-Fl. ed. 2, 1(3): 99 (1888).
= Therrya fuckelii (Rehm) Kujala, Comm. Inst. Forestry Inst.
38: 4 (1950).

74. Coccomyces pluridens (Berk. & Curt.) Sacc. See C. clusiae.

75. Coccomyces prunophorae Higgins, Amer. J. Bot. 1: 165 (1914).
= Blumeriella jaapii (Rehm) Arx.
For a discussion and references, see C. hiemalis.

76. Coccomyces pseudotsugae Funk, Canad. J. Bot. 53: 2297 (1975).
Figure 39
Apothecia intracortical, immersed, becoming erumpent, occurring
on undiscolored host tissue, orbicular to polygonal, 0.6-1.2 mm diam,
black, without a preformed dehiscence mechanism, splitting open
irregularly. Covering stroma consisting of a crust of disintegrating
carbonized hyphae 35-50 um thick, lined on its inner face with 40-50
wm of brown noncarbonized pseudoparenchymatous cells 5 wm diam.
Basal stroma black, carbonized, 20 ym thick, separated from the
subhymenium by 50 pm of pale brown interwoven hyphae 2-4 pm
diam. Subhymenium 40 ym thick, pale brown, the cells 2-4 jm diam.
Asci (113-)125-168(-200) x 8-10 pm, clavate, long-stalked, 8-spored;
ascospores nonseptate, 35-43 x 2-3 ym, rather narrowly sheathed.
Paraphyses septate, partly recurved at the tips and nodose. Pycnidia
absent.

82 MARTHA A. SHERWOOD

SSSR

NS &
iN

Fic. 39, Coccomyces pseudotsugae:—a. cross section of apothecium, X375.—b. ascus, paraphysis,
and spores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

On bark of Pseudotsuga menziesii and Abies grandis, west coast
of North America, summer and fall. Funk (1975) gives information
on the cultural characteristics of the species.

Specimens examined: NORTH AMERICA. CANADA: on Pseudotsuga menziesii, Shawnigan
Lake, Victoria Island, British Columbia, 8.X1I.1974, holotype of Coccomyces pseudotsu-
gae (pDAvFp 20863). usa: On conifer twigs, Lake Crescent, Olympic National Park,
Washington, 15.VI.1939, A. H. Smith (mic); on Abies grandis, Trinidad, California,
111.1935, Parks 5520, distributed as Clithris juniperi (micu, FH, OSC 13452).

77. Coccomyces puiggarii Speg., Bol. Acad. Nac. Ci. Cérdoba 23:
517 (1919). Figure 40
Apothecia subepidermal in prominent bleached spots bounded by

a raised, reddish discolored line, on dead leaves still attached to the
host, 0.2-0.5 mm diam, dull brown with a darker rim when seen

THE GENUS COCCOMYCES 83

Fic. 40. Coccomyces puiggarii:—a. cross section of apothecium, X375.—b. habit sketch, approxi
mately natural size. Drawn from the holotype

from above, opening irregularly by 3-5 teeth, without a preformed
dehiscence mechanism. Covering layer 25-30 jm thick, dark colored,
slightly carbonized; lower stroma 15 jm thick, composed of carbonized
cells 5 pm diam. Subhymenium colorless, 15 wm thick. Excipulum
absent. Paraphyses filiform, enlarged to 2.0-2.5 tm above, cemented
in a gel. Asci 110-120 x 8-10 um, 8-spored. Ascospores 60-75 xX
2.5 wm.

On dead leaves of Myrtaceae, Brazil. All of the apothecia remaining
in type collection are immature; dimensions of asci and ascospores
are taken from Spegazzini’s original description. The species appears
very close to Coccomyces vilis. As well as can be determined from
the scanty type specimen it lacks the inner periphysoidal layer of
C. consocians, an otherwise similar subepidermal foliicolous species
of Coccomyces known from South America. Both C. vilis and C.
puiggarii are imperfectly known. In view of the parasitic habit of
the former and widely separated geographical areas it would seem
premature to synonymize them.

Specimen examined: SOUTH AMERICA, BRAZIL: Apiahy, 1879°, leg. J. Puiggari nro. 278,
holotype of Coccomyces puiggarii (ps 28188).

78. Coccomyces quadratus [Schm. & Kunze] Sacc. See C. leptideus.

84

79.

80.

81.

82.

MARTHA A. SHERWOOD

Coccomyces quadratus [Schm. & Kunze] Sacc. var. arctostaphyli
Rehm. See Coccomyces arctostaphyli.

Coccomyces quadratus [Schm. & Kunze] Sacc. var. philippinus
Rehm. See Coccomyces philippinus.

Coccomyces quercinus (Desm.) Terrier. See Coccomyces delta.

Coccomyces radiatus Sherwood, spec. nov. Figure 41

Ascocarpi primo immersi, tetragoni vel hexagoni, 1.0-1.5 mm diam, per lacinias
4-6 aperientes, in macula pallida stromate non obvallato insidentes. Margo superior
stromatica 30 im crassa, ex hyphis intertextis brunneis constata. Periphysoidei nulli.
Margo inferior stromatica 10 jm crassa, ex hyphis carbonaceis constata. Excipulum
hyalinum. Paraphyses filiformes, apice ad 4-5 ym incrassatae, achromae, in epithecio

UZ

Fic. 41. Coccomyces radiatus:—a. cross section of apothecium, *375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch of rehydrated material, 7.5. Drawn from Shear 16.X.1925 (pt).

THE GENUS COCCOMYCES 85

gelatinoso inclusae. Asci 90-120 x 5-6 wm, cylindrici, haud pedicellati, in iodo non
caerulescentes, 8-spori; sporis filiformibus, in tunica gelatinosa inclusis, continuis,
60-75 X 1.0 um. In foliis emortuis Rhododendri, Amer. Bor.

Holotypus: Bri, on Rhododendron sp., Dismal Creek, Virginia, Oc-
tober 16, 1935, leg. C. L. Shear (sub. C. rhododendri)

Etymology: radiatus (L), radiata, referring to the stellate dehiscence
mechanism.

Ascocarps at first immersed, subepidermal, epiphyllous and hypo-
phyllous in bleached spots not bounded by a line stroma, 1.0-1.5
mm diam, mostly hexagonal, black, when closed exhibiting a distinct
preformed pale stellate dehiscence mechanism, opening by 4-6 regular
teeth to expose the clear pale orange disc. Covering stroma ca. 30
wm thick, consisting of globose brown noncarbonized cells 5-8 um
diam, without periphysoids; basal stroma 10 wm thick, carbonized,
separated from the subhymenium by a layer 15 jm _ thick of
pseudoparenchyma constructed like the covering layer. Subhymenium
colorless, 15 im thick. Proper exciple colorless, 20 1m thick, consisting
of closely-septate agglutinated hyphae 2.0 um diam. Asci cylindrical,
short-stalked, 90-120 xX 5-6 wm, J—, 8-spored. Paraphyses filiform,
abruptly inflated to 4-5 um at the apex, cemented in a gel and forming
an epithecium. Ascospores 60-75 xX 1.0 um, nonseptate, obscurely
sheathed.

On Rhododendron, Appalachian Mountains. There is an overmature
specimen on Epigaea repens (White 2526) in FH which may be this
species. The tropical American specimens are similar morphologically
and have been tentatively referred here. The species differs from C.
coronatus in having a well-defined dehiscence mechanism and para-
physes firmly cemented in an epithecial gel. Members of the tropical
C. leptosporus complex differ in having narrower spores, smaller
apothecia, and lacking the combination of a hyaline proper exciple
and paraphyses cemented in a gel.

Specimens examined (see also holotype, above): NORTH AMERICA. USA: on Rhododendron,
Rugby, Tennessee, 7.VIII.1949, Shear (Bp1); on Rhododendron, Occoquan, Va., Shear,
X1.1922 (pp); Whitewater Falls, N.C., Shear, 19.VIII.1933 (pp1); on Rhododendron
maximum, Mt. Lake, Virginia, 3.1X.1936 (BPI), SOUTH AMERICA. VENEZUELA: E] Avila,
Dpto. Federal, 24.VII.1972, leg. Dumont et al. (Ny-Ve 5863). CoLoMBIA: Medellin-Pto.

Valdivia rd., Dpto. Antioquia, 12.VIII.1976, Dumont et al. (Ny-Co 6230); Valle de
Cauca-Choco, Dumont et al., 25. VIII.1976 (ny-Co 7272).

83. Coccomyces rehmii Sacc. See Coccomyces rhododendri.

84. Coccomyces repandus ([Fr.] ex Fr.) Quélet, Enchirid. Fung.
338 (1886).
= [ Phacidium repandum Fr., K. Vet. Akad. Hand. 40: 108 (1819) ]
ex Fr. Syst, Mycol. 2(2): 578 (1823).

86 MARTHA A. SHERWOOD

= Pseudopeziza repanda ({Fr.] ex Fr.) Karst., Acta Soc. Flora
Fauna Fennica 2: 161 (1885).

85. Coccomyces rhododendri Rehm, Hedwigia 21: 117 (1882).
= Clithris rhododendri (Rehm) Rehm in Rabenh., Krypt.-FI. ed.
2, 1(3): 104 (1888).
= Coccomyces rehmii Sacc., Syll. Fung. 8: 751 (1889).

The staff of the Stockholm herbarium were unable to locate any
type or authentic specimens of the above species in Rehm’s Herbarium.
The description suggests that it may be allied to Coccomyces bipartitus
and related species, but it cannot be characterized completely from
the description alone. Rehm’s name, C. rhododendri, has priority over
Saccardo’s combination, C. rhododendri (Schw.) Sacc.; hence the name
C. rehmii is superfluous.

86. Coccomyces rhododendri (Schw.) Sacc., Syll. Fung. 8: 748 (1889).
= Hysterium rhododendri Schw., Trans. Amer. Philos. Soc. n.s.
4; 246 (1832).
= Lophodermium rhododendri (Schw.) Ell. & Ev., N. Am.
Pyrenomycet. 717 (1892), nom. inval.
= Lophodermium schweinitzii M. Wils. & Roberts, Trans. Brit.
Mycol. Soc. 61: 527 (1947), a name change.
Modern authors consider this to be a Lophodermium rather than
a Coccomyces.

87. Coccomyces rhododendri Racib. See Coccomyces javanicus.

88. Coccomyces rubi (Fr.) Karst., Bidrag Kannedom Finlands Natur
Folk 19: 258 (1871).
Phacidium rubi Fr., Syst. Mycol. 2(2): 578 (1823).
Coleroa chaetomium (Kunze ex Fr.) Rabenh., Herb. Mycol.
1456 (1850).
According to Muller and von Arx (1962), this is a Loculoascomycete
in the Venturiaceae.

89. Coccomyces rubicola Ell. & Dearn. See Karstenia rubicola.
Specimens filed under this name in American herbaria may also
be C. tumidus, which occasionally occurs on Rubus canes.

90. Coccomyces salicinus (Ell. & Ev.) Sacc., Syll. Fung. I1: 432
(1895).
= Coccophacidium salicinum Ell. & Ev., Proc. Philadelphia
Acad. Sci. 46: 151 (1893) [1894].
= Stictis schizoxyloides Ell. & Ev., l.c. p. 150.

THE GENUS COCCOMYCES 87

The two names for the taxon were published simultaneously. Stictis
schizoxyloides is preferred, since it is more familiar and reflects the
correct taxonomic position of the species. For a redescription, see

Sherwood (1977a).

Specimen examined: NORTH AMERICA. USA: Helena, Montana, Kelsey #5, 13.1.1889,
isotype of Coccophacidium salicinum (cup-b 2008).

91. Coccomyces spegazzinii Sacc., Syll. Fung. 8: 747 (1889).
Figure 42
= Coccomyces brasiliensis Speg., Bol. Acad. Nac. Ci. Cérdoba
11: 593 (1889) non C. brasiliensis Karst.
Apothecia intraepidermal, scattered in bleached areas bounded by
a black line stroma, 0.5-0.8 mm diam, triangular to hexagonal, with

I MH Lf | : |
HH! I f | sy / | ff |
/ L/f Hy LY Maal
} / f ff / I fit | /
LEA ff LHF LE

aoe =

I

Fic. 42. Coccomyces spegazzinii:—a. cross section of apothecium, X375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, approximately natural size. Drawn from Theissen 212 (FH).

88 MARTHA A. SHERWOOD

a distinct preformed dehiscence mechanism consisting of thin-walled
colorless cells, splitting open by teeth. Covering layer 20 ym thick,
of dark, moderately carbonized cells 3-7 pm diam, without periphy-
soids; lower stroma complete, carbonized, 10 ym thick. Subhymenium
20 um thick, colorless. Proper exciple well-developed, 30 »m thick,
of closely septate hyphae 3 jm diam, brown above. Paraphyses filiform,
enlarged above to 3-4 ym, not imbedded in a gel. Asci cylindrical,
short-stalked, 100-120 x 6-6.5 ym, J—, 8-spored. Ascospores nearly
as long as the asci, 1.5 pm broad, nonseptate, obscurely sheathed.

On coriaceous leaves, Brazil. Spegazzini’s type no longer contains
mature apothecia; the above description is taken from Theissen’s
exsiccata material. Coccomyces spegazzinii is similar to C. leptosporus,
differing in having somewhat broader spores and no gelatinous
epithecium. The well-developed exciple will distinguish it from the
otherwise similar C. dentatus.

Specimens examined: SOUTH AMERICA, BRAZIL: Apiahy, VII.1882, leg. J. Puiggari 2928,
holotype of Coccomyces brasiliensis Speg. (Lps 28183); Apiahy, 1889, leg. Puiggari
(Les 34990); Theissen, Decades Fungorum Brasiliensis 212, Sao Paulo, Rick, 1911 (Fx).

VENEZUELA: trail from Rincon E. along Rio Media, Edo. Sucre, Dumont et al., 15. VII.1972
(NY-Ve 5083).

9la. Coccomyces striatus (Phill. & Plowr.) Massee. See Coccomyces
tumidus.

92. Coccomyces strobi Reid & Cain, Canad. J. Bot. 39: 1127 (1961).
Figure 43
= Phacidium crustaceus Curt. in Ellis, N. Am. Fungi 455 (1880)
nom. nud.
= Coccomyces crustaceum Curt. in Reid & Cain, Canad. J.
Bot. 39: 1118 (1961) pro synon.
Coccophacidium crustaceum (Curt.) Durand, in Seymour,
Host Index 39 (1929) nom. nud.

Apothecia at first immersed, intracortical, raising the substrate into
prominent, flattened pustules, without a preformed dehiscence mech-
anism, splitting open by numerous teeth to expose the dull yellow
hymenium, 1-2 mm diam. Covering layer 50-75 ym thick, heavily
carbonized, the cells 5-8 im diam, with a distinct vertical orientation.
Basal stroma reduced to a subiculum of interwoven brown hyphae
2-3 um diam, separated from the hymenium by a matrix of hyaline
thick-walled hyphae 100 um thick which soon breaks down leaving
a mass of colorless crystals. Subhymenium colorless, 25 jzm thick.
Asci clavate, rather long-stalked, 70-120(-125) x 6-9(-10.5) ym, J-,
8-spored. Paraphyses filiform, unbranched, circinate, 1.5-2 1m diam.
Ascospores 27-40.5(-48) 1-2 ym, nonseptate, narrowly sheathed.

On recently killed branches on Pinus strobus and introduced 5-needle

THE GENUS COCCOMYCES 89

Fic. 43. Coccomyces strobi:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
spores, X750.—c. habit sketch of rehydrated material, x7.5.—c. habit sketch of rehydrated material,
7.5. (a) and (b) from FH ex DAOM 20761, (c) from rH-Pfister 9.VI.1977

pines, eastern North America, maturing in late spring, very common.
Reid and Cain (1961) cite numerous specimens and discuss the name
Coccophacidium crustaceum, which has been widely used for this
species but was never validly published. Since the fungus is nearly
ubiquitous and appears. soon after the death of the portions of the
host on which it occurs, it is tempting to brand it as parasitic and
speculate on its importance, but nothing definite is known about
the role of C. strobi in the ecology of its host.

Although introduced western North American pines planted in
Boston are susceptible to C. strobi, no natural occurrences of this
species west of the Rocky Mountains have been encountered either
in the field or in herbaria.

90 MARTHA A. SHERWOOD

Specimens examined:’ NORTH AMERICA. CANADA: Midhurst, Simcoe County, Ontario,
4.V1.1953, Reid & Cain, isotype of C. strobi (pri); New Brunswick, Marysville, Miller,
VI.1916 (spi). usa. Maine: Kittery Point, Thaxter, 1886 (rH). New Hampshire: Gorham,
14.VI.1915 (Bri). Massachusetts: Concord, Pfister, 9.VI.77 (FH); Boston, on Pinus
wallichiana, 28.V.1977, Sherwood (FH); on P. monticola, ibid. (rH); on P. avacahuite,
ibid. (FH). New York: Ithaca, Durand, 26.11.1904 (sp1). Ohio: Oxford, Fink, 2.VI.1912
(FH). Virginia: Arcadia, Hedgecock, 6.111.1928 (spt). Illinois: 2.VI.1915, Dearness (Bp).
Georgia: Talulah Falls, 12.1X.1910, Dearness (pi). Pennsylvania: State College,
30.111.1933, White (rH). Minnesota: St. Louis Co., M. Palm, 6.VI.1978 (FH). Michigan:
Ishpenning, 17.VI.1916, Hedgecock (spt).

93. Coccomyces strobilinus Grelet, Rev. Mycol. (Paris) 24: 88 (1959).
I was unable to locate type or other authentic or reliably identified
specimens of this species. Grelet’s description of elongate apothecia
opening by a longitudinal slit rather than by teeth suggests a Colpoma
rather than a Coccomyces.

94. Coccomyces tesselatus Sherwood, spec. nov. Figure 44

Ascocarpi primo immersi, tetragoni vel hexagoni, 0.5-0.8 mm diam, per lacinias
aperientes, in macula pallida stromate obvallato insidentes. Margo superior stromatica
10 pm crassa, ex hyphis intertextis carbonaceis constata. Margo inferior stromatica
5 xm crassa, ex hyphis intertextis carbonaceis constata. Periphysoidei nulli. Excipulum
hyalinum. Paraphyses filiformes, apice ad 4-5 ym incrassatae, achromae, in epithecio
gelatinoso non inclusae. Asci cylindrici, haud pedicellati, 90-120 x 5.0-5.6 ym, 8-spori,
in iodo non caerulescentes; sporis filiformibus, continuis, in tunica gelatinosa non
inclusis, 80-90 x 0.8 ym. In foliis dejectis Ingae, tropical America.

Holotypus: cup-PrR 4050, along rte 105 at km 23.8, Maricao Forest
Reserve, Puerto Rico, elev. 510 m, June 12, 1970, leg. R. P. Korf,
J. Benson, D. H. Pfister, A. Y. Rossman & L. Skog. Isotypi: FH, MAPR,
NYS.

Etymology: tesselatus (L), tiled, referring to the appearance of the
much-dissected bleached spots on leaves.

Apothecia scattered in prominent bleached spots bounded by a black
line stroma on dead coriaceous leaves, 4-6 sided, 0.5-0.8 mm diam,
without a preformed dehiscence mechanism, black, shining, splitting
open by irregular teeth to expose the bright yellow hymenium, drying
orange, diffusing a soluble yellow pigment in water. Covering layer
10 ,.m thick, carbonized, the cells ca. 5 tm diam; periphysoids absent.
Basal stroma 5 wm thick, carbonized. Proper exciple consisting of
3-4 rows of hyaline closely-septate hyphae 4-5 1m diam. Subhyme-
nium colorless, 25 ym thick, separated from the basal stroma by a
hyaline subiculum interspersed with a few colorless crystals. Pa-
raphyses filiform, enlarged to 4-5 wm at the apex, colorless, not
cemented in a gel. Asci cylindrical, short-stalked, J—, 90-120 x 5.5-6.0

‘Only selected specimens of this common and frequently collected species are listed. Host is
Pinus strobus unless otherwise indicated.

THE GENUS COCCOMYCES 91

/ Yf WY A
Yi), | I) DN afininnan

/
|

HAMEL A

Fic. 44. Coccomyces tesselatus:—a. cross section of apothecium, *375.—b. ascus, paraphyses,
and spores, X750.—c. habit sketch, *7.5. Drawn from Cup pR 4050.

zm, 8-spored; ascospores 80-90 x 0.8 wm, not obviously sheathed.

On Inga and unidentified coriaceous leaves, tropical America,
apparently fairly common. Basically Coccomyces tesselatus is distin-
guished from allied species by the intensely pigmented hymenium
which diffuses a soluble yellow pigment in KOH.

Specimens examined (see also holotype, above): CARIBBEAN. PUERTO RICO: Woods near
Maricao Fish Hatchery, on Inga laurina, Korf et al., 15.VI.1970 (Cup-pk 4119); Mayaguez,
Britton, 18-27.11.1914 (Ny). GUADELOUPE: Transverse Road, Pfister et al., 4.1.1974 (FH-
GUADELOUPE 844). GRENADA: Grand Etang, Thaxter 1912-13, 2612 (FH). pomiINnica: Roseau,
Korf et al., 20.VI.1970 (cup-po 13, ny).

95. Coccomyces tetracerae (Rudolph in Fr.) Sacc., Sylloge Fungor-
um 8: 748 (1889).

= Phacidium tetracerae Rudolph in Fr., Linnea 5: 551 (1830),

92 MARTHA A. SHERWOOD

= Hysterostomella tetracerae (Rudolph) Hohnel, (Sitzungsber.

Acad. Wiss., Math.-Naturwiss. Kl. Abt. 1, 118: 1541 (1909).

The type specimen in herb. Fries agrees well with the illustration

and description provided by von Hohnel (1l.c.) of a crustlike, multi-

loculate ascostroma which becomes apothecioid by cracking open

irregularly, bitunicate asci, hyaline, 2-celled spores, and dark conidia
with a hyaline equatorial band.

Specimen examined: CARIBBEAN. CUBA: on Tetracera volubilis, Poeppig, holotype of
Phacidium tetracerae (ups).

96. Coccomyces tjibodensis Racib. See Biostictis tjibodensis.

97. Coccomyces triangularis (Schw.) Sacc., Syll. Fung. 8: 750

(1889). Figure 45
= Peziza triangularis Schw., Schriften Naturf. Gesell. Leipzig
1: 122 (1822).
= Cenangium triangularis (Schw.) Fr., Syst. Mycol. 2(1): 182
(1822).
= Colpoma triangularis (Schw.) Petr., Sydowia 11: 346 (1957)
[1958].

Apothecia at first immersed, subcortical, becoming erumpent, re-
maining for a long time closed, covered by a thick, buff-colored,
pruinose covering layer, triangular, square, or oblong in face view,
never linear, without a preformed dehiscence mechanism, cracking
open irregularly, usually by 3 teeth. Covering layer up to 400 pm
thick, externally crystalliferous, with a thick, black, carbonized crust
and an internal layer of thin-walled brown hyphae 2-4 wm diam
running parallel to the surface, becoming thick-walled and carbonized
on the interior face. Periphysoids abundant, 150 * 2-3 wm, branched
but not netlike, imbedded in a gel. Basal stroma massive, up to 300
um thick, the lower portion carbonized, separated from the subhyme-
nium by a matrix of vertically oriented pale brown hyphae 5-8 pm
diam, widely spaced in a gel. Subhymenium colorless, 40-50 jm
thick. Asci clavate, long-stalked, 130-160 x 9-11 wm, 8-spored;
ascospores 80-100 X 1.5 ym, nonseptate, narrowly sheathed. Paraphy-
ses enlarged to 2.5 1m above, weakly circinate.

On dead twigs of Quercus alba, maturing in late summer and fall,
eastern North America, common, apparently host specific. The generic
position of the species is unclear. As Petrak (1958) indicated, it is
scarcely a typical Coccomyces. The type of Colpoma, C. quercinum
(Pers. ex Fr.) Wallr., is a hysterothecial fungus which invariably opens
along a preformed longitudinal slit flanked by lip cells. I have chosen
to exclude from Colpoma and include in Coccomyces those wood-
inhabiting Rhytismataceae which lack a preformed longitudinal slit

THE GENUS COCCOMYCES 93

UN TAT Tail

Fic. 45. Coccomyces triangularis:—a. cross section of apothecium, X75.—b. ascus, paraphysis,
and spores, X750.—c. habit sketch, X7.5. Drawn from Shear 5907 (FH).

and lip cells. Coccomyces triangularis is most closely related to C.
castanopsidis, which occurs on Fagaceae in western North America.
The morphological distinctions between these two species and typical
leaf-inhabiting species of Coccomyces may well be sufficient to warrant
erection of a separate genus to accomodate them; it would be premature,
however, to propose generic segregates in Coccomyces without revising
Colpoma as well.

Specimens examined:” NORTH AMERICA. CANADA: Komoka, Ontario, 8.V.1915, Dearness
(BPI). Usa. Massachusetts: Farlow, Newton, 4.VII.1891 (FH). Connecticut: Lakeville,
Hansbrough, 7.VI.1933 (pei). New York: Ithaca, 16.V.1933 (cup 22, 561). New Jersey:

Fungi Columbiani 410, Newfield, Ellis, VI.1894 (pp, rH); Newfield, 1874, de Thuemen,
Mycotheca Universalis 570 (FH). Pennsylvania?: Herb. Curtis, leg. Schweinitz, authentic

8 P - ° FS .
Only selected specimens, indicating the geographical range, are cited.

94 MARTHA A. SHERWOOD

(FH). Maryland: Takoma Park, Shear 5907, 1899 (FH). Washington, D.C.: 7.V.1899 (ppl).
South Carolina: Ravenel, Fungi Caroliniani 2: 42 (FH). Georgia: Winterville, Miller,
26.1V.1926 (cup 18266). Louisiana: Palmetto, Shear, 16.11I.1932 (ppt). Ohio: Hocking
Co., Fink, 6-4-1918 (pi). Illinois: Kewaunee, Harper, V.1899 (FH).

98. Coccomyces tridentatus (Lév.) Sacc. See Myriophacidium tri-
dentatum.

99. Coccomyces trigonus [Schm. & Kunze] Quel. See Coccomyces
tumidus.

100. Coccomyces tumidus (Fr.) de Not., Giorn. Bot. Ital. 2 (7-8): 14,
38 (1847). Figure 46
= Hysterium tumidum Fr., Syst. Mycol. 2(2): 591 (1823).

Y jeer sera
& LIP net |
Z ‘ae, x) eR b

as =, ‘
NG S727 rr
\ RB NE ce

Fic. 46. Coccomyces tumidus:—a. cross section of apothecium, X225.—b. detail of asci, paraph-
yses, and spores, X750.—c. habit sketch, X7.5. Drawn from Allerscher & Schnabl 548 (FH).

THE GENUS COCCOMYCES 95

= Lophodermium tumidum (Fr.) Lamb., Fl. Myc. Belg. 2: 452
(1880).
[= Peziza comitialis Batsch, Elench. Fung. cont. 1: 217, t. 27 f. 152
(1786)].
= Coccomyces comitialis [Batsch] Dearn. & House, Bull. N.Y.
State Mus. 266: 65 (1925),
[= Phacidium trigonum Schm. & Kunze, Myc. Hefte 1: 40 (1817)].
= Coccomyces coronatus °** trigonus [Schm. & Kunze] Karst.,
Bidrag Kannedom Finlands Natur Folk 19: 257 (1871).
= Coccomyces trigonus [Schm. & Kunze] Quélet, Enchirid.
Fung. 337 (1886).
= Phacidium peltiforme Preuss, Linnaea 26: 724 (1853).
= Coccomyces peltiformis (Preuss) Sacc., Syll. Fung. 8: 745

(1889).

Phacidium striatum Phill. & Plowr., Grevillea 13: 75 (1885).
= Coccomyces striatus (Phill. & Plowr.) Massee, Brit. Fung.

Fl. 4:51 (1895).

= Henriquesia quercina Grove, J. Bot. (London) 66: 136 (1935).

Apothecia subcuticular, widely scattered on long-dead leaves, on
bleached areas usually bounded by a black line, 1.0-2.0 mm diam,
orbicular or elongate, dark brown, wrinkled, when immature appearing
hat-shaped from above, with a flat brim, raised margin, and depressed
center, fleshy when rehydrated, splitting open irregularly by teeth,
or when hysterothecial by a longitudinal slit, to expose the dull yellow
to yellowish-tan disc. Covering layer 120 um thick, consisting of an
outer layer of dark brown, but not heavily carbonized hyphae 3-5
wm diam, running parallel to the surface of the substrate, and a central
matrix of fleshy brown globose pseudoparenchymatous cells 3-5 wm
diam, separated from the subhymenium by a matrix of closely-packed,
pale brown hyphae 2-5 pm diam. Subhymenium colorless, 45 wm
thick. Asci clavate, long-stalked, 160-220 x 10-15 wm diam, J-,
8-spored. Paraphyses filiform, mostly unbranched, circinate, 1.5 um
broad. Ascospores 32-45 Xx 3.0-4.5 4m, nonseptate, but sometimes
appearing pseudoseptate due to the presence of large guttules, narrowly
sheathed.

On fallen leaves of a variety of plants, commonest on Quercus
and Fagus, but reported on Betula, Acer, Populus, and Gaultheria,
rarely on Rubus canes, late summer and fall, very common and
widespread in Europe and North America. The hysterothecial form
and orbicular form agree in all details of stroma construction and
hymenial elements; I do not doubt that they are the same species.
The two forms do not, however, commonly co-occur in the same
lesion; hence, it is possible that there is some genetic distinction
between them. One can learn to recognize this species on macroscopic

96 MARTHA A. SHERWOOD

characters alone. The large, fleshy, rugose apothecia are unmistakable.

Considerable confusion exists in the literature concerning Cocco-
myces tumidus. The name Lophodermium tumidum, based on the
hysterothecial form, was used by Rehm (1887-96) and others (for
example, Romell in Fungi Scandinaviae Exs. 84 and Fuckel, Fungi
Rhenani 746) for a bona fide and entirely distinct Lophodermium
on Sorbus aucuparia. Nannfeldt (1932) realized that two taxa were
involved, and used L. tumidum (ascribed to Rehm) for the Lophoder-
mium on Sorbus, synonymizing Fries’s species with C. coronatus.
The epithet tumidum cannot legitimately be used in this sense, and
now this species is known as Lophodermium aucupariae (Schleich.
ex Schleich.) Darker.

The discocarpous form has widely been considered to be a synonym
of Coccomyces coronatus and is distributed under that name in
numerous exsiccatae. A partial list of such misidentifications is includ-
ed in the list of specimens cited. Since the collections from which
the exsiccatae specimens were drawn may well have been mixed,
the presence of the species in the examples I examined does not
guarantee that it was present in all sets of an exsiccata issued.

No type or authentic specimens of Peziza comitialis, Phacidium
trigonum, or Phacidium peltiforme could be located; the synonymy
is based on descriptions alone. The isotype specimen of Hysterium
tumidum in FH is immature, but aggrees well in stromatal characters
and appearance with better-developed specimens of the species.

Specimens examined:” EUROPE. SWEDEN: Fries, Scleromycetes Sueciae 166, on Fagus,
isotype of Hysterium tumidum (FH). DENMARK: Sordé, on Quercus, Pfister, 9.X.1978 (FH).
BELGIUM: Libert, Pl. Crypt. Ard. 175, on Fagus, sub Phacidium coronatum (BPI, LG,
FH); Fungi Gallici 2279, on Betula, Brussels (FH). GERMANY: Sydow, Mycoth. Germ.
2714 p.p., Brandenberg, sub C. coronatus (spi); Rehm, Ascomyceten 1803, on Quercus,
sub C. coronatus (ppl, FH-Hohnel); Schmidt & Kunze, Deutschlands Schwamme 82
(BPI, not FH). CZECHOSLOVAKIA: Rudics, on Populus tremula, Svrcek (pRM 149920); on
Betula, Ulehle, Svréek (prM 756277); on Quercus, Bohemia (PRM 620799); on Fagus,
Brno (pRM 659122). POLAND: on Quercus, Kalmien, Kubicka, 2.1X.1966 (pRM 624897).
ITALy: Erbar. Crittog. Ital. 780, on Castanea, Locarno, 1860 (FH). NORTH AMERICA, CANADA,
Ontario: Dorset, on Quercus rubra, Cain, 11.1X.1963 (Bri). Nova Scotia: on Populus
tremuloides, Wehmeyer 1198 (mMicH); on Fagus and Acer, Wehmeyer 1297 (MicH). USA.
Maine: Enfield, Shear, 23.VIII.1940 (spi). New York: McClean, on Rubus setosus (cup
14944); on Betula, Adirondack Mts., Kauffman & Mains (micu). Massachusetts: Concord,
Pfister, [X.1978 (FH). New Hamphire: Mt. Pacmonadnock, on Quercus rubra, Sherwood,
1X.1978 (FH). Virginia: Mt. Lake on Quercus, 4.1X.1936, Cash (ppt). Washington: on

Gaultheria shallon, Seattle, Piper, IX.1892 (Bri). Oregon: Woodburn, on Quercus
garryana, Clemens (bri); Blue River, on Gaultheria, Sherwood, VIII.1978 (FH).

101. Coccomyces tympanidiosporus Sherwood, spec. nov. Figure 47

Ascocarpi primo immersi, orbiculari vel irregulariter elongati, 0.5-1.0 mm diam,
per lacinias irregulariter aperientes, in macula pallida insidentes. Margo superior

9 : . . «< ye P A :
Only one specimen/host/locality has been included. “p.p.” indicates that the collection involved
included more than one species of Coccomyces.

THE GENUS COCCOMYCES 97

c——— J
soa

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aos

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Fic. 47. Coccomyces tympanidiosporus:—a. cross section of apothecium, *375.—b. asci,
paraphysis, spores, and ascoconidia, X750.—c. habit sketch, X7.5. Drawn from the holotype.

stromatica 90 wm crassa, ex hyphis intertextis carbonaceis constata. Periphysoidei nulli.
Margo inferior stromatica 20 ym crassa, ex hyphis carbonaceis constata. Excipulum
nullum. Paraphyses filiformes, apice ad 3.0-3.5 ym incrassatae, in epithecio gelatinoso
non inclusae. Asci 140-165 x 12.5-14.0 pm, clavati, pedicellati, 8-spori, in iodo non
caerulescentes; sporis filiformibus, septatis, in tunica gelatinosa inclusis, 65-90 x 2.5
um. In ligno decorticato, Ins. Canariensis.

Holotypus: cup-Mycoflora Macaronesia 807, on pine? wood, La
Palma, Canary Islands, along road in hills above la Giratte, Jan. 16,
1976, leg. W. C. Denison and M. A. Sherwood. Isotypus:FH.

Etymology: tympanidiosporus, (L), having spores like a Tympanis,
i.e., producing ascoconidia by budding.

Apothecia at first immersed in old, bleached, decorticated wood,
becoming more or less superficial by erosion of the overlying substrate,

98 MARTHA A. SHERWOOD

gregarious and tending to form confluent crusts, 0.5-1.0 mm diam,
orbicular to irregularly elongate but not linear, black, rugose, splitting
open irregularly by 4-6 teeth to expose the clear orange hymenium.
Covering layer 90 pm thick, heavily carbonized above, merging below
into pale brown pseudoparenchyma of globose cells 8 1m diam, without
periphysoids. Basal stroma carbonized, 20 jm thick, separated from
the hymenium by a layer of small crystals and hyaline hyphae.
Subhymenium colorless, 20 jm thick. Paraphyses filiform, un-
branched, enlarged to 3.0-3.5 wm at the apex, not imbedded in a
gelatinous epithecium, slightly agglutinated near the periphery of the
ascocarp but not sufficiently differentiated to constitute a proper
excipulum. Asci clavate, long-stalked, 140-165 x 12.5-14.0 pm, 8-
spored; ascospores 65-90 X 2.5 ym, filiform, 3-septate, prominently
sheathed, eventually budding off rod-shaped ascoconidia 5.0 xX 1.0
Lm.

Coccomyces ericae (q.v.) is similar, but the spores lack the conspic-
uous sheath, are not septate, and do not bud off ascoconidia.
Coccomyces ericae also has a prominent proper exciple lacking in
the present species.

Specimens examined (see also holotype, above): ATLANTIC ISLANDS. CANARY ISLANDS: on

Cistus monspeliensis 17 km SW of San Sebastian de la Gomera, towards Tejiade,
Gomera, 4.1.1977, Korf et al. (cup-mm 1375).

102. Coccomyces umbonatus (Preuss) Sacc., Syll. Fung. 8: 750 (1889).
= Phacidium umbonatum Preuss, Linnaea 26: 724 (1853).
I was unable to locate any type or authentic material of this species,
which was described from bark of Pinus in Europe. It may be a
synonym of Therrya pini.

103. Coccomyces urceolus Sherwood, spec. nov. Figure 48

Ascocarpi primo immersi, tetragoni, 0.3-0.6 mm diam, per lacinias aperientes, en
macula pallida stromate obvallati insidentes. Margo superior stromatica 30 Lm crassa,
ex hyphis intertextis brunneis constata. Periphysoidei nulli. Margo inferior stromatica
8-10 ,m crassa, ex hyphis intertextis carbonaceis constata. Excipulum brunneum.
Paraphyses filiformes, apice ad 3 ym incrassatae, in epithecio gelatinoso non inclusae.
Asci 75-100 x 4.0 um, cylindrici, haud pedicellati, 8-spori, in iodo non caerulescentes;
sporis filiformibus, continuis, in tunica gelatinosa non inclusis, 50-75 xX 0.8 um. In
foliis dejectis ignotis, Venezuela.

Holotypus: Ny-Fungi of Venezuela 2053, on unidentified leaf ca.
14 km above Maracay, on the Maracay-Choroni road, Parq. Nac. Henry
Pittier, Edo. Aragua, K. P. Dumont, J. H. Haines, & G. J. Samuels,
12 July 1971.

Etymology: urceolus (L), urceolate, referring to the deeply immersed
fruitbodies.

THE GENUS COCCOMYCES 99

Fic. 48. Coccomyces urceolus:—a. cross section of apothecium, X375.—b. asci, paraphyses, and
ascospores, X750.—c. habit sketch, X7.5. Drawn from the holotype.

Ascocarps subepidermal, scattered in slightly bleached areas bound-
ed by a black line on decaying coriaceous leaves, at first appearing
as concolorous pustules bounded by a black line, later opening by
4-5 irregular teeth, without a preformed dehiscence mechanism,
0.3-0.6 mm diam. Covering stroma 30 wm thick, of brown, noncarbon-
ized hyphae, without periphysoids. Basal stroma 8-10 pm thick,
carbonized, the cells 5-7 wm diam. Proper exciple 30 wm thick,
well-developed, of closely septate hyphae 2-3 1m diam, brown above.
Subhymenium colorless, 15 wm thick. Asci cylindrical, short-stalked,
75-100 x 4.0 wm, 8-spored, J—. Paraphyses filiform, enlarged to 3.0
ym at the apex, the apex solid and refractive. Ascospores 50-75 xX
0.8 wm, nonseptate, not obviously sheathed.

On leaves, tropical America. The thickened, solid, refractive apices

100 MARTHA A. SHERWOOD

of the paraphyses are distinctive and distinguish this taxon from
Coccomyces leptosporus, which is also intraepidermal rather then
subepidermal.

Specimens examined (see also holotype, above): CARIBBEAN. GUADELOUPE: Grand Etang,

Pfister & Sarreira 527 (FH). SOUTH AMERICA, VENEZUELA: Rancho Grande, Parque Nac.
Henry Pittier, Edo. Aragua, Dumont, Haines, & Samuels, 3.VIL.1971 (Ny-Ve 1213).

104. Coccomyces ursinus (Sacc. & Paol.) Petr., Ann. Mycol. 32: 322
(1936).
= Coccomyces quadratus [Schm. & Kunze] Sacc. subsp. ursinus
Sacc. & Paol., Bull. Soc. Bot. Belg. 28: 96 (1889).
According to B. Eriksson (1970), no type specimen of this taxon
remains, but it is probably a synonym of Naemacyclus phacidioides.

105. Coccomyces venezuelae Sherwood, spec. nov. Figure 49

Ascocarpi primo immersi, trigoni vel tetragoni, 0.2-0.4 mm diam, per lacinias
aperientes, in macula pallida non insidentes. Margo superior stromatica 20 jm crassa,
ex hyphis intertextis brunneis constata. Periphysoidei nulli. Margo stromatica inferior

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Fic. 49. Coccomyces venezuelae:—a. cross section of apothecium, X375.—b. asci, paraphyses,
and spores, X750.—c. habit sketch, 7.5. Drawn from the holotype.

THE GENUS COCCOMYCES 101

10 »m crassa, ex hyphis intertextis carbonaceis constata. Excipulum hyalinum. Para-
physes filiformes, ramosae, apice non incrassatae, achromae, in epithecio gelatinoso
non inclusae. Asci 75 X 4.5-5.5 ym, cylindrici, haud pedicellati, 8-spori, in iodo non
caerulescentes; sporis filiformibus, continuis, in tunica gelatinosa non inclusis, 50-60
x 0.8 wm. In foliis dejectis ignotis, Venezuela, Amer. Austral.

Holotypus: Ny-Fungi of Venezuela 221, on unidentified leaf, El
Eden, Parq. Nac. El Avila, Edo. Miranda, K. P. Dumont, J. H. Haines
& B. Manara, 19 June 1971.

Etymology: venezuelae (L), from the country where the species
occurs.

Apothecia intraepidermal, not accompanied by obvious bleaching
ora black line, on well-rotted leaves, 0.2-0.4 mm diam, when immature
appearing as nearly concolorous pustules bounded by a faint black
line, splitting open to expose the pale yellow disc, the reflexed lobes
of the covering layer translucent. Covering layer 20 wm thick, of
pale brown hyphae 3-5 pm diam, carbonized along the lines of
dehiscence; basal stroma 10 um thick, heavily carbonized. Subhyme-
nium colorless, 15 ym thick, separated from the basal stroma by a
matrix of colorless hyphae and crystals. Excipulum 20 pm thick, of
closely-septate colorless hyphae 2.5-3.0 wm diam. Asci cylindrical,
short-stalked, 75 X 4.5-5.5 um, J—, 8-spored. Paraphyses filiform,
colorless, not enlarged apically, branched. Ascospores 50-60 x 0.8
44m, nonseptate, not obviously sheathed.

On well-rotted leaves, Venezuela. Short asci, branched paraphyses,
a reduced covering layer, and lack of bleached spots bounded by
a black line distinguish this species from Coccomyces leptosporus
and its allies.

Specimen examined (see also holotype, above); SOUTH AMERICA. VENEZUELA: Parq. Nac.
El Avila, Edo. Miranda, Dumont et al., 30.VI.1972 (Ny-Ve-3816).

106. Coccomyces vilis Sydow & Butler, Ann. Mycol. 9: 377 (1911).
Figure 50
= Coccomyces memecycli Sydow, Ann. Mycol. 12: 570 (1914).
?= Naemacyclus korfii Rao, Ullasa & Patil, J. Shivaji Univ.
5: 131 (1972).

Apothecia imbedded in distinct bleached necrotic spots bounded
by raised reddish discolored veins, 0.2-0.5 mm diam, orbicular or
triangular, when closed with a dark rim and dark radiating lines
marking the zones of dehiscence, otherwise scarcely darker than the
surrounding leaf tissue. Apothecia intraepidermal, hypophyllous, the
covering layer nearly obsolete, with 15 wm of dark carbonized cells
surrounding the zone of dehiscence; basal stroma ca. 20 pm thick,
carbonized, separated from the subhymenium by acolorless subiculum.
Subhymenium colorless, 20 tm thick. Asci 70-90 x 5.0-6.5 (fide

102 MARTHA A. SHERWOOD

Fic. 50. Coccomyces vilis:—a. cross section of apothecium, X375.—b. habit sketch, X7.5, Drawn
from the holotype.

Sydow) p.m, 8-spored; ascospores filiform, colorless, nonseptate, nearly
as long as the asci. Paraphyses colorless, unbranched, scarcely enlarged
above.

On living leaves of Mangifera indica, Malda, India, and possibly
on other hosts throughout tropical Asia. The holotype of the species,
cited below, is extremely immature. The type specimen of Coccomyces
memecycli is in poor condition but agrees externally with C. vilis;
the asci of C. memecycli were, however, originally described as being
8.0-9.5 1m broad. Coccomyces javanicus (q.v.) may be an older name,
but the synonymy is open to question. Another possible synonym,
judging from the description, is Naemacyclus korfii, on Carissa from
India. Coccomyces puiggarii, occurring in Brazil, is a very similar

THE GENUS COCCOMYCES 103

species, known only from a depauperate type specimen. Insufficient
information precludes either synonymizing these two species or
separating them definitively on morphological grounds.

Specimens examined: asia. INDIA: Herb. Crypt. Orient 1372, on Mangifera indica, Malda,
5-8-1905, coll. E. Butler, holotype of C. vilis (s). pHiLippINES: auf Memecyclon lanceola-
tum, Luzon, Antipolo, Prov. Rizal, leg. Ramos, 19.1.1914, holotype of C. memecyli
(s); Elmer, Phillipine Island Plants 16236, Luzon, Mt. Bulusan, June, 1916 (FH).

107. Coccomyces villae-vicosae Torrend, Bull. Soc. Portugaise Sci.
Nat. 3: 7 (1919).

I was unable to locate any type or authentic material of this species,
described on pine wood from Portugal. It is impossible to determine
from the original description whether it is a species of Coccomyces,
Colpoma, or Therrya.

108. Coccomyces viridis (Richon) Sacc., Syll. Fung. 10: 51 (1892).
= Phacidium viride Richon, Cat. Champ. Marne 223 (1889).

I was unable to locate any material, type or otherwise, of this species.
The original publication describes a fungus 1-2 mm diam, opening
by teeth, with a brown epithecium and spores 40 um long, occurring
on bark of Populus. The description might possibly pertain to a fungus
similar to Propolidium fuscocinereum Ell. & Ev., which occurs on
Populus in North America; P. fuscocinereum, however, has spores
only 20 um long.

109. Coccomyces wagnerianus (Rehm) Rehm, Ber. Bayerische Bot.

Gesellsch. 13:131 (1912). Figure 51

= Belonidium wagnerianum Rehm in Rabenh., Krypt.-Fl. ed. 2,
1(3): 1230 (1896).

Apothecia gregarious, at first immersed, 0.5-1.5 mm diam, often
elongate, black, opening along preformed lines of dehiscence, either
by teeth or by a longitudinal slit to expose the yellowish disc. Covering
stroma 100 um thick, carbonized without, fleshy within, of brown
globose cells 5-8 um diam, the margins provided with unbranched
lip cells 30 * 3.0 um diam. Lower stroma 20 ym thick, carbonized.
Subhymenium colorless, 50 jm thick. Asci clavate, long-stalked,
100-120 x 12 pm, J-—, 8-spored; ascospores 25-35 xX 3.0-3.5 um,
nonseptate, narrowly sheathed. Paraphyses circinate.

On Opuntia rafinesquii, Germany. The species is undeniably rhytis-
mataceous, but its correct generic placement is uncertain. It may be
an unusual growth form of a Hypoderma.

104 MARTHA A. SHERWOOD

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Fic. 51. Coccomyces wagnerianus:—a. cross section of apothecium, X375.—b. habit sketch, X7.5.
Drawn from the holotype.

Specimen examined: EUROPE. GERMANY: a faul. Opuntia rafinesquii Engelm., 8.1893,
in hort. meo leg. E. Wagner, holotype of Belonidium wagnerianum (s).

110. Coccomyces yerbae Speg., Ann. Mus. Nac. Buenos Aires 17:
132 (1908).

The type specimen contains three minute apothecia of a Coccomyces
occurring on bleached spots on dead leaves of Ilex paraguayensis.
According to the original description the asci measured 50 x 5-6
wm and the ascospores were | pm broad. Without additional material
on this host from South America the species cannot be characterized
entirely; it appears to be closely related to C. spegazzinii.

Specimen examined: SOUTH AMERICA. ARGENTINA: S/ Ilex paraguayensis, Misiones, San
Pedro, II.1906, leg. C. Spegazzini, holotype of C. yerbae (Lps 28184).

THE GENUS COCCOMYCES 105

ALLIED GENERA AND EXCLUDED SPECIES

Biostictis Petr., Sydowia 4: 357 (1950).

Biostictis, a genus of the Stictidaceae, is characterized by a parasitic
habit, immersed apothecia framed by a toothed, crystalline margin,
cylindrical capitate asci, and cylindrical or filiform, septate ascospores.
For further information and a formal description of the genus, see
Sherwood (1977a, 1978). Coccomyces tjibodensis Racib. is a Biostictis.
It may be distinguished from the other species in the genus as follows:

KEY TO BIOSTICTIS SPECIES

1. Ascospores filiform, multiseptate, nearly as long as the asci............, 2:
1’. Ascospores cylindrical or cylindric-fusiform, much shorter than the asci, 3-5 sep-
rr Sid Ci I aren OTT pri nunrane: 7
2(1). Marginal hyphae in part pigmented. Ascospores 2.0 4m broad. On La-
Santhus EUCTIO RICO \ 15-05 19.8- 6 a ey ed Re Aor B. portoricensis Sherw.
2'(1). Marginal hyphae colorless. Ascospores 1.5 ym broad. On various Rubiaceae,
South America. .........---20 5200058 B. psychotriae (Mont.) Sherw.

3(1’). Ascospores 3-septate; paraphyses branched. On undetermined leaves, Trinidad.
eg EC NMS 6, ori Sic ios Bie de & oy Ae Lah a gh B. chroodiscoides Sherw.
3’(1’). Ascospores 5-septate; paraphyses unbranched. On Lasianthus spp., southeast

SiGe ee Nee ols SRS. 8, ache, 4 Al ayia wed, aod lay arg od a ee nn B. tjibodensis.

Biostictis tjibodensis (Racib.) Sherw., comb. nov. Figure 52

= Coccomyces tjibodensis Racib., Paras. Alg. Pilze Javas 3: 19
(1900).

= Coccomyces mussaendae Sawada, Rept. Gov. Res. Inst. Formosa
87: 2 (1944) nom. inval.

Apothecia hypophyllous, gregarious in reddish discolored patches
on living leaves, at first immersed, opening by splitting the overlying
substrate irregularly, 0.3-0.6 mm diam, the margin brown, not crystal-
line, the disc moderately deeply immersed, dark brown. The reflexed
epidermal lobes tend to break off. Margin consisting of disintegrating
host tissue and hyphae, lined on the inner face with a compact layer
of colorless, branched, periphysoids 15 * 1.0 pm. Subhymenium
colorless. Asci cylindrical, thick-walled when young, with an indistinct
apical thickening when mature, J—, 65-85 x 10-15 pm, 8-spored.
Ascospores 24-30 X 2.5-3.0 ym, 5-septate, constricted at the septa,
not sheathed. Paraphyses filiform, unbranched, colorless, 1.0 wm diam.

On Lasianthus and Mussaenda (Rubiaceae), Java, Philippine Is-
lands, Taiwan, and China, common, according to Raciborski (l.c.).
No conidial stage was found associated with any of the specimens
examined, although conidial stages have been reported for other species
of Biostictis (Petrak, 1950; Sherwood, 1978). Sawada (l.c.) described
Coccomyces mussaendae as causing reddish-brown lesions, having
ascocarps 0.3-0.8 mm diam, 8-spored asci 60-65 x 7-10 ym, and

106 MARTHA A. SHERWOOD

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Fic. 52. Biostictis tjibodensis:—a. cross section of ascocarp, X750.—b. asci, ascospores, and
paraphyses, 750.—c. habit sketch, X7.5. Microanatomical details from zr-Raciborski-Salak; habit
sketch from zt-Philippine Fungi 5200.

5-septate spores 31-38 X 2.0-2.5 um. I have not seen type material,
but suspect it isa synonym of B. tjibodensis. The name C. mussaendae
is invalid since Sawada did not provide a Latin diagnosis for it.

Specimens examined: asia. JAVA: on Lasianthus latifolius, Gedeh, leg. Raciborski, isotype
(?) of Coccomyces tjibodensis (rH-Hohnel 4526); on Lasianthus spec., Salak, leg.

Raciborski, isotype? (ZT). PHILIPPINE ISLANDS: on Lasianthus tashiroi, Mindanao, M. S.
Clemens 5200 (zr). CHINA: on Lasianthus, Yunnan, Tsai 5267, 25.X11.1932 (Fx).

Cerion Massee & Rodway, Bull. Misc. Inform. 1901: 159 (1901).

Stromata erumpent from woody plant parts, eventually appearing
nearly superficial, massive, heavily carbonized, when mature opening
by splitting irregularly to expose the hymenium and remaining as
a thick, rugose margin mimicking a true excipulum, internally
composed of pseudoparenchymatous hyphae arranged nearly perpen-
dicular to the surface. Asci cylindrical, short-stalked, uniformly thin-
walled, J—. Paraphyses filiform, colorless, simple or branched. Asco-
spores long-filiform, obscurely sheathed, simple or 1-septate.

THE GENUS COCCOMYCES 107

Cerion was formerly included in the Stictidaceae; its rhytismataceous
affinities are discussed by Dennis (1958) and Sherwood (1977a). The
genus includes two species, C. coccineum Massee & Rodway, from
Australasia, and C. leucophaeum (Speg.) Dennis, from tropical Ameri-
ca. The involved synonymy of the latter includes species of Gorgoni-
ceps, Coccomyces, and Godronia.

Cerion leucophaeum (Speg.) Dennis, Kew Bull. 13: 465 (1958).
Figure 53
= Gorgoniceps leucophaeus Speg., Bol. Acad. Nac. Ci. 11: 589
(1889).
= Godronia splendida Speg., l.c.

Fic. 53. Cerion leucophaeum:—a. cross section of apothecium, X225.—b. detail of asci, paraphy-
ses, and ascospores, *750.—c. habit sketch, <7.5. Drawn from FH-Thaxter, Grenada 1912-13.

108 MARTHA A. SHERWOOD

= Coccomyces brasiliensis Karst., Hedwigia 28: 193 (1889) non.
Speg.
= Godronia jamaicensis Seaver, Mycologia 37: 356 (1945).

Apothecia initially intracortical, becoming erumpent and appearing
nearly superficial at maturity, 1-4 mm diam, orbicular to slightly
elongate or irregular in outline, the margin black, shining, rugose,
the disc barely immersed, pale pinkish buff to bright red, conspicuously
pruinose. Margin in cross section ca. 100 wm thick, of radiating,
tightly-packed, slightly thick-walled hyphae 4-8 wm diam (narrower
near the hymenium) lying at a high angle to the surface, becoming
pseudoparenchymatous and heavily carbonized near the surface.
Subhymenium colorless, 30 wm thick, resting on ca. 100 wm of loosely
interwoven colorless hyphae 2 1m diam, subtended by a thick, heavily
carbonized basal stroma. Paraphyses filiform, 1.5 jm broad, repeatedly
dichotomously branched near the apex (propoloid), J—, colorless,
forming a distinct epithecium. Asci 200-230 x 8-10 ym, cylindrical,
short-stalked, the apex blunt and slightly thickened, J—, 8-spored.
Ascospores 100-150 X 1.5 im, nonseptate, obscurely sheathed.

On decaying twigs and bark, plurivorous, at moderate elevations
throughout tropical and subtropical America, common and conspicu-
ous. Cerion coccineum is virtually identical but has unbranched or
once-branched paraphyses and 1-septate spores.

The names Coccomyces brasiliensis Karst. and C. brasiliensis Speg.
(q.v.) were both published in 1889. I can find no clear indication
which has priority. Spegazzini (1919) stated that the publications
appeared within a few days of each other and that he could not
resolve which had priority. Saccardo (1892), providing a new name,
C. spegazzinii, for Coccomyces brasiliensis Speg., considered Karsten’s
publication to be earlier.

Although I have been unable to locate a name in the pre-1889
literature, I suspect that one will eventually be found for this common
and conspicuous species, and therefore have adopted Dennis’s com-
bination rather than introduce another combination into the literature.

A specimen from Colombia (Ny-co 5611), from bark of a living
tree, differed from typical Cerion leucophaeum in having once-
branched paraphyses 2.0-2.5 wm diam, small apothecia, and asci 175
um long. It may be distinct, but does not differ from typical C.
leucophaeum sufficiently to be described as new on the strength of
a single specimen.

Specimens examined; NORTH AMERICA, MEXICO: Chiapas, Dumont, 19. VIII. 1967 (CUup-ME
285). CARRIBBEAN, JAMAICA: Cincohona, Murrill 495, holotype of Godronia jamaicensis
(NY). CENTRAL AMERICA. COSTA RICA: Prov. San Jose, 10.VII.1936, Dodge and Gregaer (Ny).
PANAMA: Prov. Chocle, Dumont et al., 14.VI.1975 (Ny-Pa 277); Prov. Chiriqui, Dumont

et al., 2.VII.1975 (ny-Pa 1632, 1689). sourH AMERICA. VENEZUELA: Edo. Merida, Dumont
et al., 1971 (Ny-Ve 3426, 2352); Edo. Yaracuy, Dumont et al., 7.VII.1971 (Ny-Ve 1597):

THE GENUS COCCOMYCES 109

Dto. Federal, Dumont et al., 22.VI.1976 (Ny-Ve 456); Edo. Barinas, Dumont et al..
26.VII.1971 (ny-Ve 3151); Edo. Bolivar, Dumont et al., 4.VIII.1972 (ny-Ve 6820).
Cotomsia: Dpto. Cundinamarca, Dumont et al., 1974 & 1976 (Nvy-Co 8105, 4029, 4185.
2195, 959, 941, 439, 1869); Dpto. Boyacd, Dumont et al., 12.1X.1976 (Ny-Co 7728):
Dpto. Narino (Ny-Co 3343); Dpto. Antioquia (Ny-Co 825, 6593); Dpto. Choc6, Dumont
et al. (NY-Co 5611). ECUADOR: Prov. Pastaza, Dumont et al., 24.VII.1975 (Ny-Ec 1446).
BRASIL: Minas Geraes, E. Wainio 1885, holotype of Coccomyces brasiliensis (H, herb.
Karst. 1740); Apiahy, 1888, leg. J. Puiggari, holotype of Gorgoniceps leucophaeus (ps
24437); Sao Paulo, Apiahy, leg. J. Puiggari 2919, holotype of Godronia splendida
(Les 1003).

Karstenia Fr. in Karst., Acta Soc. Fauna Fl. Fenn. 2(6): 166 (1885),

Four species previously included in Coccomyces appear rather to
belong in Karstenia, a genus of immersed discomycetes characterized
by a fleshy pseudoparenchymatous covering layer with vertically
oriented cells, cylindrical or clavate, uniformly thin-walled asci without
a J+ apical pore (the hymenial gel is sometimes J+ blue), true
paraphyses, and cylindrical to filiform, multiseptate spores. The
affinities of Karstenia at the family level are obscure (Sherwood, 1977a),
but it does not appear to be closely related to Coccomyces either
morphologically or ecologically, and would be quite out of place
in the Rhytismataceae.

Karstenia clematidis (Phill.) Sherwood, comb. nov.
= Phacidium clematidis Phill., Grevillea 16: 46 (1888).
= Coccomyces clematidis (Phill.) Sacc., Syll. Fung. 8: 751
(1889).

Apothecia immersed, opening by a pore and eventually splitting
the overlying substrate irregularly, 0.3-0.6 mm _ diam, colorless
throughout. Covering layer 50 wm _ thick, of vertically arranged
pseudoparenchymatous cells ca. 3 jm diam. Paraphyses filiform,
simple, colorless, not enlarged above, J—. Asci cylindric-saccate, ca.
70 X 10 um, 8-spored. Ascospores long-cylindrical, not sheathed, 35-45
x 3-4 um, closely septate, the cells 3-4 um long.

Similar to Karstenia idaei (Fckl.) Sherw., but the asci are shorter.
Species concepts in this genus have yet to be defined; a brief
redescription has been provided here since the species is not an obvious
synonym.

Specimen examined; EUROPE. GREAT BRITAIN: Carlisle, Dr. Carlisle, on stems of Clematis,
holotype of Phacidium clematidis (k).

Karstenia macer (Karst.) Sherwood, comb. nov. Figure 54
= Phacidium macrum Karst., Not. Sallsk. fauna flora fennica 11:
218 (1871).

= Coccomyces macer (Karst.) Karst., Bidrag Kannedom Fin-
lands Natur Folk 19: 255 (1871).

110 MARTHA A. SHERWOOD

ft AN
“SAIKK)
a ZAM

a

Fic. 54. Karstenia macer:—a. cross section of apothecium, *375.—b. ascus, paraphysis, and
spore, X750.—c. habit sketch, X7.5. Drawn from the holotype.

Apothecia immersed, becoming erumpent by splitting the overlying
substrate irregularly, 1.0-1.5 mm diam, orbicular, externally dark grey,
the surrounding host tissue slightly bleached but not exhibiting defined
spots. Covering layer ca. 50 pm thick, of pale brown angular cells
3.5 wm diam, with a distinct vertical orientation, not ending in a
fringe of hyphal filaments. Subhymenium pale brown, ca. 10 pm
thick. Apothecia subepidermal. Asci 65-80 x 8-10 ym, thin-walled,
pointed, J—. Ascospores 8, ca. 60 X 2.5 ym, 7-septate, not sheathed.
Paraphyses filiform, 1.5-2.0 ym broad, a little enlarged at the apex,
closely septate.

On umbelliferous (?) stems, Finland.

Specimen examined: EUROPE. FINLAND: Ad. caul. umbellif?, I.VI?.1866, Mustiala, leg.
P. A. Karsten 1737, holotype of Phacidium macrum (nu).

Karstenia maydis (Rehm) Sherwood, comb. nov. Figure 55
= Phacidium maydis Rehm, Ascom. Lojk. 23 (1883).

= Coccomyces maydis (Rehm) Sacc., Syll. Fung. 8: 752 (1889).
Apothecia immersed, orbicular, inconspicuous, pale brownish,
opening by splitting the overlying substrate irregularly. Covering layer
ca. 50 um thick, pale brown above, not carbonized, of angular, vertically
oriented cells 3-4 pm diam, not ending ina fringe of hyphal filaments.
Asci uniformly thin-walled, J—, 60-80 x 8-10 um, 8-spored. Ascospores

THE GENUS COCCOMYCES 111

\)
ESN AS
ire Pe

Fic. 55. Karstenia maydis:—cross section of apothecium, <150. Drawn from the holotype.

30-55 X 2.5 pm (30-36 x 1.5 fide Rehm), not sheathed, usually
5-septate. Paraphyses filiform, enlarged to 2.0 um at the apex.
On cornstalks, Hungary.

Specimen examined; EUROPE. HUNGARY: Herb. Rehm, Kaposvar, 1872, Lojka 112,
holotype of Phacidium maydis (s).

Karstenia rubicola (Ell. & Dearn.) Sherwood, comb. nov. Figure 56
= Coccomyces rubicola Ell. & Dearn., Proc. Canad. Inst. 1: 90
(1897).

Apothecia intraepidermal, becoming erumpent, scattered, orbicular,
ca. 0.7 mm diam, splitting the overlying substrate irregularly, the
margin dark, not toothed, the disc moderately deeply immersed, pale
ochraceous. Covering layer pale brown, 50 wm thick, of thin-walled
pseudoparenchymatous cells 5-8 m diam with a vertical orientation,
not ending in a fringe of hyphoid hairs. Subhymenium 20 pm thick,
pseudoparenchymatous, colorless. Asci 50-60 X 7 um, short-stalked,
cylindric-clavate, J—, without an obvious apical dehiscence mechan-
ism, 8-spored; paraphyses colorless, filiform, enlarged to 2.5 ym at
the apex, septate. Ascospores 40-50 (50-60 fide Ellis & Dearness)
x 2-2.5 wm, usually 5-septate, not sheathed. The hymenial gel turns
faintly blue in iodine.

On Rubus strigosus, Canada, known only from a fragmentary type
collection. A second specimen in paom (House 829) proved to be
a Pyrenopeziza. The name has been applied in herbaria to Coccomyces
tumidus (q.v.), which occasionally occurs on canes of Rubus.

Specimen examined: NORTH AMERICA, CANADA: on Rubus strigosus, Wm. Grants, Aug.
1895, holotype of Coccomyces rubicola (DAoM).

112 MARTHA A. SHERWOOD

Fic. 56. Karstenia rubicola:—a. cross section of apothecium, X375.—b. ascus, paraphyses, and
spores, X750,—c. habit sketch, X7.5. Drawn from the holotype.

Myriophacidium Sherwood, Mycologia 66: 791 (1974).

One species formerly assigned to Coccomyces, C. tridentatus, be-
longs to the related genus Myriophacidium, distinguished from Cocco-
myces by the possession of oval spores. The type species, in addition,
has an epithecium of paraphyses cemented in a brown gel. This is
lacking in the two species discussed below. Although distinguishing
a genus on the basis of spore characters alone is arbitrary and probably
artificial, it has long been sanctioned in the Rhytismataceae.

Both the holotype and isotype specimens of Phacidium tridentatum,
cited below, are mixtures of two different species of Myriophacidium,
one on leaves of Vaccinium, the other on twigs of an unidentified
host, possibly Cavendishia. Since Léveilleé described the species as
occurring on leaves, I have designated the portion on leaves as the
type of Phacidium tridentatum, and described the portion on twigs
as a distinct species. The three species of Myriophacidium are distin-
guished as follows:

KEY TO MYRIOPHACIDIUM SPECIES

1. Paraphyses cemented in a brown epithecial gel... . . M. aphyophyllicum Sherw.
1’. Paraphyses and epithecium colorless... . 1-6-0 ee 2.
2. Ascospores 17.5-22.5 x 5.0-7.5 pm. On stems of Cavendishia?, Colombia
Ak cs dco ty, aotnrhs bk Oe WODp ee bo eae Gi Ee BO ae es ae te) we A M. corticola.
2’. Ascospores 16-30 x 3.0-5.0 pm. On leaves of Vaccinium, South America
2 gh Ged ee Gb Pad Ha Aral De Ged REO ee Aenea eee Re ee M. tridentatum.

THE GENUS COCCOMYCES 113

1h)

Y),

aa
be Ee

Fic. 57. Myriophacidium corticola:—a. cross section of margin, X375.—b. ascus, paraphysis,
and spores, X750.—c. habit sketch, X7.5. Drawn from the type in Pc.

Myriophacidium corticola Sherwood, spec. nov. Figure 57

Ascocarpi primo immersi, dein erumpentes, trigoni vel tetragoni, 0.8-1.5 mm diam,
per lacinias aperientes, in macula pallida stromate obvallato insidentes. Margo superior
stromatica 60 ym crassa, ex hyphis intertextis carbonaceis constata. Periphysoidei nulli,
Margo inferior stromatica 15-20 ym crassa, ex hyphis intertextis carbonaceis constata.
Excipulum nullum. Paraphyses filiformes, apice ad 1.5 jm incrassatae. Asci 100-105
x 15-18 pm, cylindrici-clavati, pedicellati, in iodo non caerulescentes, 8-spori. Sporis
ellipsoideis, continuis, in tunica gelatinosa inclusis, 17.5-22.5 5.0-7.5 pm. In corticem
Cavendishiae?, Bogota, Montserrat, Colombia.

Holotypus: pc, Herb. Lindig 2790, Bogota, Nova Granata, Montser-
rat, p.p. Isotypus: FH.

Etymology: corticola (L), growing on bark, as opposed to the other
two species, which occur on leaves.

Apothecia immersed in bleached spots bounded by a delicate black

114 MARTHA A. SHERWOOD

line on the smooth cortex of dead twigs, triangular (less commonly
quadrate), 0.8-1.5 mm diam, in face view with a distinct black rim,
lighter colored ‘hinge,’ and black center marked by well-defined
preformed dehiscence lines. Covering layer in cross section 60 xm
thick, heavily carbonized toward the center, becoming colorless and
fleshy toward the periphery, the cells 3-5 im diam, oriented parallel
to the surface of the substrate. Basal stroma 15-20 ym thick, of dark
carbonized cells 5 wm diam. Subhymenium colorless, 20 wm thick.
Asci ca. 100-105 15-18 pm, more or less cylindrical, short-stalked,
blunt-ended, with an obscure nonrefractive apical thickening. Para-
physes filiform, simple, barely enlarged to 1.5 wm at the apex.
Ascospores 8, 17.5-22.5 X 5.0-7.5 wm, prominently sheathed, nonsep-
tate.

On dead twigs of an unidentified woody plant (probably ericaceous,
possibly Cavendishia), Bogota, Montserrat (elev. 9500 ft). The con-
struction of the stroma is quite different from the construction of
the stroma of M. tridentatum.

Fic. 58. Myriophacidium tridentatum:—a. cross section of apothecium, X*375.—b. detail of
construction of covering layer, X750.—c. asci, paraphysis, and spores, X750. Drawn from Ny-Co
5065a.

THE GENUS COCCOMYCES LS

Myriophacidium tridentatum (Lév.) Sherwood, comb. nov. Figure 58
= Phacidium tridentatum Lév., Ann. Sci. Nat. Botanique sér. 4,
20: 291 (1863).
= Coccomyces tridentatus (Leév.) Sacc., Syll. Fung. 8: 748
(1889).
Apothecia intraepidermal, occurring on bleached spots bounded by
a black line, triangular or quadrate, 0.3-1.0 mm diam, black, shining,
opening by 3-4 regular teeth along preformed lines of dehiscence
to expose the dull yellow hymenium. Covering layer 40-50 wm thick,
heavily carbonized. Basal stroma 1 cell thick, heavily carbonized;
subhymenium colorless, 5-10 wm thick. Asci 65-100 11.5-15.0 wm,
cylindric-clavate, stalked, with a slightly thickened blunt apex, J-,
8-spored. Paraphyses colorless, filiform, simple, enlarged to 1.5(-2.5)
wm at the apex; ascospores 16-30 X 3.0-5.0 wm, conspicuously
sheathed.
On dead leaves of Vaccinium sp. at high elevations in the Andes,
South America.

Specimens examined: SOUTH AMERICA. COLOMBIA: Hb. Lindig, Bogota, Montserrat, nr.
2790 p.p. (pc, holotype of P. tridentatum, ru); Dpto. Boyaca, Sogamoso-Aguazul rd.,
13.VI.1976, Dumont et al. (Ny-Co 5065). peru: Dpto. Junin, Carrizales, 9.VII.1976,
Dumont et al. (Ny-Pe 1087).

ACKNOWLEDGEMENTS

The author would like to thank the directors and staff of the numerous herbaria
who made available the specimens on which this study was based. G. Cummins (ariz),
D. Farr (Bet), L. Gomez P. (cr), R. P. Korf (cup), J. Parmelee (pAom), A. Funk (pavrp),
A. D. Rehling (Fr), D. H. Pfister (FH), E. Shaw (cu), J. Poelt (Grz), T. Ahti (), B.
C. Sutton (im1), D. Reid (xk), S. Ahmad (Lan), V. Demoulin (tc), I. Gamundi (cps),
R. Shaffer (Mico), P. Holmgren and K. P. Dumont (ny), J. Haines (Nys), K. L. Chambers
(osc), L. Curti (PAD), Mme Jovet-Ast (pc), M. Svrcek (prM), A. Sunderwell (s), A. Raitviir
(rad), Y. Doi (TNs), S. Sivertsen (TRH), I. Tavares (uc), O. Eriksson (uME), R. Moberg
(ups) and E. Muller (zr) assisted by locating specimens and processing loans. M. S.
Christiansen (0) provided color photographs of Schumacher’s manuscript illustrations.
A. Raitviir, L. Gomez P., S. Ahmad, and F. Candousseau made available valuable
personal collections from unusual localities.

Jack D. Rogers, David L. Hawksworth, D. W. Minter, Donald H. Pfister, Everett
S. Luttrell, Richard P. Korf, Linda M. Kohn, Frank DiCosmo, Ove Eriksson, Richard
Hanlin, and Margaret E. Barr-Bigelow reviewed the manuscript and made many helpful
and perceptive suggestions for its improvement. The author was able to collect fungi
in the French West Indies under National Science Foundation grant GH-36162 to
D. H. Pfister, in the Canary Islands under N.S.F. Grant DEB 75-23557 to R. P. Korf,
and in South America under N.S.F. grant GB-28593 to K. P. Dumont. Toshiyuki Aoki
provided assistance with Japanese literature, and Elizabeth Shaw (GH) corrected the
Latin diagnoses.

116 MARTHA A. SHERWOOD

HOST INDEX

The listing below includes the recorded hosts for all species of
Coccomyces treated in the text, including those which are imperfectly
known and those which have been transferred to other genera. The
fungus name given is the one under which the species is listed in
the text.

Host names conform to currently accepted nomenclature. Host names
given in ‘specimens examined’ at the end of each description were
copied directly from herbarium labels and may include synonyms
and other obsolete names.

The symbols L(leaves), S(stems), B(bark), and W(wood) refer to
the portions of the host on which the fungus usually occurs. Cocco-
myces species are quite specific in the portion of the host on which
they grow.

Host indices should be used with caution in identifying members
of the Rhytismataceae. Some species are quite host specific, but the
host range of common pluriverous species is very broad.

ACERACEAE
Acer sp.
Coccomyces tumidus (1)
ANACARDIACEAE
Mangifera indica L.
Coccomyces vilis (L)
APOCYNACEAE
Carissa sp.
Coccomyces vilis (L)
AQUIFOLIACEAE
Ilex paraguayensis A. St. Hil.
Coccomyces yerbae (t)
BERBERIDACEAE
Berberis aquifolium Pursh
Coccomyces dentatus (L)
Berberis nervosa Pursh
Coccomyces dentatus (t.)
Berberis repens Lindl.
Coccomyces dentatus (t)
BETULACEAE
Betula sp.
Coccomyces coronatus (L)
Coccomyces laciniatus (1)
Coccomyces tumidus (t)
BROMELIACEAE
Coccomyces dentatus (t)
Coccomyces pampeanus (L)
BURSERACEAE
Canarium sp.
Coccomyces canarii (L)
Dacryodes excelsa Vahl
Coccomyces leptosporus (L)

CACTACEAE

Opuntia rafinesquii Engelm.
Coccomyces wagnerianus (Ss)

CARYOPHYLLACEAE

Dianthus carthusianorum L.
Coccomyces dianthi (s)

CISTACEAE

Cistus monspeliensis L.
Coccomyces tympanidiosporus (Ww)

CUPRESSACEAE

Juniperus communis L. subsp. nana

(Willd.)

Coccomyces juniperi (B)

Juniperus virginiana L.
Coccomyces petersii (B)

CYPERACEAE

Carex pauciflora Light.
Coccomyces insignis (L)

DILLENIACEAE

Tetracera volubilis H. B. K.
Coccomyces tetracerae (L)

ERICACEAE

Andromeda polifolia L.
Coccomyces duplicarioides (.)

Arbutus menziesii Pursh
Coccomyces arbutifolius (L)
Coccomyces dentatus (L)

Arctostaphylos uva-ursi (L) Spreng.
Coccomyces arctostaphyli (1)

Cavendishia sp.
Coccomyces consociatus (L)
Myriophacidium corticola (s)

THE GENUS COCCOMYCES

Epigaea repens L.
Coccomyces radiatus (.)

Erica azorica Hochst.
Coccomyces ericae (Ww)

Gaultheria shallon Pursh
Coccomyces dentatus (L)
Coccomyces leptideus (s)
Coccomyces tumidus (.)

Ledum groenlandicum Oeder
Coccomyces ledi (s)

Ledum palustre L.

Coccomyces ledi (s)

Macleania sp.

Coccomyces dentatus (1)

Rhododendron macrophyllum G. Don.
Coccomyces dentatus (.)
Coccomyces leptideus (s)

Rhododendron maximum L.
Coccomyces radiatus (1)

Vaccinium sp.

Coccomyces monticola (1)
Myriophacidium tridentatum (.)

Vaccinium myrtillus L.
Coccomyces leptideus (s)

Vaccinium ovatum Pursh
Coccomyces albidus (1)

Vaccinium vitis-idaei L.
Coccomyces leptideus (s)

FAGACEAE

Castanea sativa L.

Coccomyces dentatus (t.)
Coccomyces tumidus (L)
Castanopsis chrysophylla (Dougl. ex
Hook.) DC
Coccomyces castanopsidis (B, w)
Coccomyces dentatus (1)

Fagus grandifolia Ehrh.
Coccomyces coronatus (t,)
Coccomyces tumidus (1)

Fagus sylvatica L.

Coccomyces coronatus ()
Coccomyces dentatus ()
Coccomyces tumidus (1)
Lithocarpus densiflora (Hook. & Arn
Rehd.
Coccomyces dentatus (1)

Quercus sp.

Coccomyces coronatus (.)
Coccomyces dentatus (.)
Coccomyces tumidus 1.)

Quercus alba L.

Coccomyces dentatus (1)
Coccomyces triangularis (B)

Quercus chrysolepis Liebm.
Coccomyces castanopsidis (8)

Quercus coccifera L.

Coccomyces delta (L)
Quercus garryana Dougl. ex Hook.

Lit

Coccomyces dentatus (1)
Quercus ilex L.
Coccomyces dentatus (\)
Quercus kelloggii Newb.
Coccomyces dentatus (\.)
Quercus michauxii Nutt.
Coccomyces dentatus (1)
Quercus nigra L.
Coccomyces dentatus (1)
Quercus reticulata Humb. & Bonpl.
Coccomyces dentatus (1)
Quercus robur L.
Coccomyces dentatus (1)
Quercus rubra L.
Coccomyces farlowii (1)
Coccomyces tumidus (1)
Quercus vaccinifolia Kellogg
Coccomyces dentatus (1)
Quercus virginiana L.
Coccomyces dentatus ()
GRAMINEAE
Zea mays L.
Karstenia maydis (s)
GUTTIFERAE
Clusia sp.
Coccomyces clusiae (t.)
Coccomyces dentatus (.)
Coccomyces limitatus (t)
Clusia rosea Jacq.
Coccomyces clusiae ()
LAURACEAE
Coccomyces leptosporus (t)
Laurus nobilis L.
Coccomyces delta (t)
Laurus canariensis W. B.
Coccomyces delta (t.)
Neolitsea sp.
Coccomyces philippinus (1)
Ocotea foetens Benth. & Hooker
Coccomyces delta (t)
LEGUMINOSAE
Inga laurina (Sw.) Willd.
Coccomyces tesselatus (1)
MELASTOMATACEAE
Memecyclon lanceolatum Cogn.
Coccomyces vilis (1)
MORACEAE
Ficus minahassae Miq.
Coccomyces dubius (1)
MUSACEAE
Musa sp.
Coccomyces pampeanus (L)
Musa paradisica L.
Coccomyces annulatus (1)
MYRICACEAE
Myrica faya Ait.
Coccomyces dentatus (1)
Coccomyces foliicola (t)

118 MARTHA A, SHERWOOD

Myrica gale L. Tsuga heterophylla (Raf.) Sarg.
Coccomyces boydii (B) Coccomyces heterophyllae (8)

MYRTACEAE Tsuga mertensiana (Bong.) Carr.

Eucalyptus sp. Coccomyces cfr. heterophyllae (8)
Coccomyces martinae (L) POLYPODIACEAE

OXALIDACEAE Coccomyces crystalligerus (L,S)

Oxalis sp. Pteridium sp.

Coccomyces cfr. leptideus (s) Coccomyces dentatus (L)

PALMAE RANUNCULACEAE
Coccomyces palmicola (tL) Clematis sp.

Cocos nucifera L. Karstenia clematidis (s)
Coccomyces cocoes (L) ROSACEAE,

Eremospathus macrocarpa Mann & H. _ Kerria japonica DC

Wendl. Coccomyces kerriae (LS)
Coccomyces cocoes (L) Prunus spp.

PINACEAE Coccomyces hiemalis (Lt)
Coccomyces atactus (Ww) Coccomyces lutescens (L)
Coccomyces bipartitus (w) Coccomyces prunophorae (L)

Abies sp. Rubus hispidus L.

Coccomyces parvulus (w) Coccomyces tumidus (s)

Abies grandis (Dougl.) Lindl. Rubus strigosus Michx.
Coccomyces pseudotsugae (B) Kartstenia rubicola (s)

Larix laricina (DuRoi) K. Koch RUBIACEAE
Coccomyces irretitus (B) Lasianthus sp.

Picea mariana (Mill.) B.S.P. Biostictis tjibodensis (L)
Coccomyces bipartitus (w) Lasianthus latifolius Blume
Coccomyces irretitus (B) Biostictis tjibodensis (L)

Pinus sp. Lasianthus tashiroi Mats.
Coccomyces parvulus (w) Biostictis tjibodensis (L)
Coccomyces pini (B) Mussaenda sp.

Coccomyces villae-vicosae (B) Biostictis tjibodensis (L)

Pinus ayacahuite Ehrenb. SALICACEAE
Coccomyces strobi (B) Populus sp.

Pinus cembra L. Coccomyces coronatus (L)
Coccomyces cembrae (w) Coccomyces viridis (B)

Pinus monticola Dougl. Populus tremuloides Michx.
Coccomyces strobi (8) Coccomyces tumidus (L)

Pinus resinosa Ait. Salix sp.

Coccomyces irretitus (B) Coccomyces salicinus (Ww)

Pinus strobus L. SARRACENIACEAE
Coccomyces strobi (B) Sarracenia purpurea L.

Pinus wallichiana A, B. Jackson Coccomyces coronatus (L)
Coccomyces papillatus (8) UMBELLIFERAE
Coccomyces strobi (B) Karstenia macer (s)

Pseudotsuga menziesii (Mirb.) Franco Eryngium paniculatum Cav. & Dombey
Coccomyces parvulus (w) Coccomyces pampeanus (L)

Coccomyces pseudotsugae (B)

THE GENUS COCCOMYCES 119

LITERATURE CITED

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Arx, J. A. von. 1961. Uber Cylindrosporium padi. Phytopathol. Z. 42: 161-165.

& E. MUuLveER. 1954. Die Gattungen der amerosporen Pyrenomyceten. Beitr.
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Carro_., F. E., E. MULLER & B. C. Sutton, 1977. Preliminary studies on the incidence
of needle endophytes in some European conifers. Sydowia 29: 87-103.

Carro_i, G. C. & F. E. Carrour. 1978. Studies on the incidence of coniferous needle
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Darker, G. T. 1932. The Hypodermataceae of conifers. Contrib. Arnold Arboretum
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. 1967. A revision of the genera of the Hypodermataceae. Canad. J. Bot. 45:
1399-1444.

Dearness, J. & H. D. Houser. 1925. New or noteworthy species of fungi. Bull. N. Y.
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Dennis, R. W. G. 1958. Critical notes on some Australian Helotiales and Ostropales.
Kew Bull. 12: 321-358.

. 1977. British Ascomycetes. Vaduz. 585 p., 31 fig., XLIV pl.

DiCosmo, F. 1979. Lasiostictis reassessed. Canad. J. Bot. 57: 1834-1840.

Dusy, M. 1961. Mémoire sur le tribu des Hysterinées. Mém. Soc. Phys. Hist. Nat.
Geneve 16: 15-70, tab. 1-2.

Eriksson, B. 1970. On Ascomycetes on Diapensales and Ericales in Fennoscandia.
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No. 11.

No. 12.
_ Kallymeniaceae (Rhodophyta). A Comparison of the Species of

No. 13.

No. 14.

No. 15.

Robert K. Edgar: An Annotated Bibliography of the American
Microscopists and Diatomist Jacob Whitman Bailey 1811-1857).
Donald H. Pfister: A Note on Phaeofabraea and its Placement in
the Leotiaceae Subfamily Encoelioideae (Discomycetes) (February
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Gayle I. Hansen: Cirrulicarpus Carolinensis, A New Species in the

Cirrulicarpus (Kallymeniaceae, Rhodophyta). Monte G. Manuel:
Studies in Cryphaeaceae Ill. Sphaerotheciella Fleisch. New to the
Americas (April 1977).

Norton G. Miller and Robert R. Ireland: A Floristic Account of the
Bryophtes of Bathurst Island, Arctic Canada. Martha A. Sherwood:
New Ostropales from the Collections of the Farlow Herbarium (July
1978).

Carroll W. Dodge: Edward Angus Burt (1859-1939). Robert K. Edgar:
Jacob W. Bailey and the Diatoms of the Wilkes Exploring Expedition
(1838-1842). Martha A. Sherwood: New Phacidiales and Ostropales
from the Collections of the Farlow Herbarium (July 1979).

Martha A. Sherwood: Taxonomic studies in the Phacidiales: The
genus Coccomyces (Rhytismataceae) (April 1980).

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