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
reel BE % AS AD)
oe +,
;
< Pens Bear APU Le iG
rays -*e ws
LAT
2 (Q
‘e a 7,8, ? VR
%
~ >.
|
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
; Q on Ne Ww RENE S To¥6
\) Ss
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
See ne ‘ai @ RASS Av 3 ne abst ae
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)
» 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
_ = = aos
S
—
<a
= eH
=a =S
aa —
SES SSE ‘
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
¥
Hl | IT} | {|
|
I
Smee
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
rh.
VA
a
€.
s ¢, Ge
@Q
~
i
Orr INA
Cam toma
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
awe
x
t} uN
»/
.
we \ yy:
A
ny
!
¢.
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)