PERSOONIA

Published by the Rijksherbarium, Leiden

Volume Part 6, 4, pp. 433-438 (1972)

Differentiationof tribes and genera in

the family Sarcoscyphaceae

William+C. Denison

Department of Botany, Oregon State University, Corvallis

(With Plates 24—25)

used in and Taxonomic criteria differentiating genera are described

illustrated. The three the tribes: genera are assigned to tribes, two existing

Sarcoscypheae and Boedijnopezizeae, and a new tribe Pithyeae.

decade One product of the intensive study of operculate discomycetes in the past

has been of families with defined boundaries recognition new clearly (Eckblad,

1968, Kimbrough, igyo). This paper discusses the internal structure of one of these

Korf describes new families, the Sarcoscyphaceae sensu (igyi). It criteria, common

which delimit the then those which in to all genera, family, proceeds to occur some

genera and not in others, and which are, therefore, useful in delimiting genera.

Finally it discusses the division of the family into three tribes, including a new one,

the Pithyeae.

As presently circumscribed, the family Sarcoscyphaceae no longer includes those

with dark-colored in the tribe Urnuleae. genera apothecia formerly placed They

have been transferred the Sarcosomataceae to a separate family, (Korf, igji).

The family Sarcoscyphaceae is characterized by two groups of taxonomic char-

acters. One consists of characters shared by all members of the family. Some of

these also in the Sarcosomataceae. The other is occur group equally characteristic, but is made up of characters which occur in some genera and not in others. The

described this the second will be included the first group is at point: among criteria for delimiting genera.

The suboperculate ascus (Le Gal, ig46) occurs in all members of the suborder

Sarcoscyphineae. A characteristic organelle, the subapical pad, can be seen as a thickened addition to the inner side of the apex of the ascus wall (Plate 23, Figs.

B, C). In the families Sarcoscyphaceae and Sarcosomataceae the tissues of the medullary excipulum are filamentous, textura intricata, as seen either in vertical section or in

the of the in these crush mounts. As a consequence texture apothecia families is tough, leathery to rubbery.

* Paper read at the Symposium "Taxonomy of operculate Discomycetes" held at the

First International Mycological Congress, Exeter, 1971. 1 This in from the National Science Foundation study was supported part by a grant (GB-6589).

433 Persoonia Vol. Part 434 6, 4, 1972

In the families Sarcoscyphaceae and Sarcosomataceae the ascospores are single-

but contain 16 in celled, they numerous nuclei, or 32 most genera (Berthet, 1964).

In the family Sarcoscyphaceae the cells of the paraphyses are multinucleate,

the uninucleate whereas in the family Sarcosomataceae the cells of paraphyses are (Berthet, 1964).

In the family Sarcoscyphaceae the hymenial pigments are bright colored (red, orange, yellow) caroteinoids (Arpin, 1968), and the exterior of the apothecium is light colored. In rare instances hymenial pigments are absent and the hymenium is white.

In the wood in family Sarcoscyphaceae, apothecia occur on or on foliage early stages of decomposition. In a few genera they are found on soil, although they

arise from buried wood if well- probably or roots. They seldom, ever, occur on very rotted charcoal. wood, on dung or compost, or on

The family Sarcoscyphaceae consists of sixteen genera arranged in three tribes:

Sarcoscypheae, Boedijnopezizeae, and Pithyeae.

DESCRIPTION OF GENERIC CHARACTERS

characters useful in the The following are delimiting genera within family.

Table lists the and summarizes the distributionof these characters 1 genera among the genera.

APOTHECIAL SIZE.—There is a wide range of apothecial size within the family.

Most members of the tribe Sarcoscypheae have medium-sized to large apothecia,

from in diameter.In the tribe ranging 1-5 cm or more Pithyeae, however, apothecia are seldom more than a few millimeters across.

—Most the APOTHECIAL SHAPE. genera in family have shallow, cupshaped to

short discoid apothecia with stipes. Two groups ofgenera depart from this pattern. In Boedijnopeziza, Cockeina, Microstoma, Geodina, and sometimes in Sarcoscypha, the apothecium is deeply cup-shaped or funnel-shaped with a long stipe. In Wynnea and the other the is Aurophora, on hand, apothecium strongly unequal-sided (fan-

with lateral shaped or ear-shaped) a stipe.

SYNCHRONOUS ASCUS DEVELOPMENT.—In the genera Microstoma, Boedijnopeziza, and Cookeina, ascus development is synchronous, so that all of the asci in an apothe-

the of of these cium are at same stage development (Plate 22 Fig. B). Specimens

be collected full in order find In the genera must at maturity to any mature spores.

the asci mature few at that section ofthe remaining genera a a time, so a hymenium shows all stages of development from young asci to mature or discharged ones.

LATERAL —In nine the is eccentric and OPERCULUM. genera operculum strongly the is side of the of the In the sub- opening to one apex ascus. undischarged asci,

be in this The apical pad may seen position (Plate 23 Fig. C.). remaining genera have terminal, or nearly terminal operculi and subapical pads (Plate 23 Fig. B.).

FOUR-SPORED — the Thindia ASCI. In monotypic genus (Korf & Waraitch, 1971); in two of the three species of Nanoscypha (Denison, 1971); and in some species of C. Denison: Differentiation in Sarcoscyphaceae 435

Table 1.-< •

Geodina Pindara Acervus Wynne Pithya Thindia UGenericeneric CharactersCh id ■-id üa Ol the fa rj Cookeina a Aurophora Rickiella Phillipsia of the familyCc >. o Microstoma Boedijnopeziza Sarcoscypha Nanoscypha Pseudopithyella Desmazierella SarcoscyphaceaeSarcoscypXrt 4) 3

+ + ? + B + + + + + + i Ë

+ + + + + + + + + o o» o gg a

+ + + Ol idM o « 0.V ■o >oth

£ i X + + + + + vi g0,0) ecia

T3 i, 0 + + + + + + + + + + + "3 0">3-C U o. 0.

c X uo C o 3 in + + + >• u

"- £ ral

+ ? + + + + + + + rculum ocl 4> asci - £Tiinal + ? + + + + + "o 0) rculum a

■* i W pored + +

X B "2 + U.

D 3g + + + + + + + + + + + + + a w o

0) >- 1Ë a h O ld + + + + + ascospores g 01 3 h«i •O + + + + + O.

a o 3 UI + + + + *.. ￿ j + + +

-O n + + ha

U in a u + irs

id ■ U u3~-< + +

■d 0) ? Oh ? + + 9 +

Uo0 V .s id 1

+ + + ? + ?

FE .2 s00 i ectal

+ + + excipulum s» n.

U-j OO w u aXi«

+ >- 0.

X i a

? ? ? ? ? ? ? ? ? ? + ? ? + S..S xid ment Pig- .2 a c

? ? ? ? ? + ? + ? ? ? ? ? + 5S x c .

u 'S. Uid ? + + + ? ? + + + +

F eM ■ + + o, + ? + + + + + +

o d oo -c + + + + + + + habitat u 0 'S.2 00 4)* 4) + + + + +

0 ■3

+ + + + + Persoonia Vol. Part 436 6, 4, 1972

the asci and the than in Phillipsia, are regularly four-spored ascospores are larger

related eight-spored species. Species which are normally eight-spored produce a

aborted few four-spored asci, but in these one usually finds the remains of four

spores.

ten ASCOSPORE SHAPE.—The species of Pithya have spherical ascospores. In other

the genera the ascospores are symmetrical and ellipsoidal (Plate 23 Fig. B.). In

remaining five genera most ascospores are asymmetrical. They are slightly unequal-

sided curved in and to (Plate 23, Fig. C.). Although Phillipsia Nanoscypha a majority

of have there is in each at least species unequalsided ascospores, genus one species with symmetrical ascospores.

CYANOPHOBIC — the have SCULPTURING. Five genera of family cyanophobic sculp-

the surface of their This consists of folds turing on ascospores. longitudinal ridges or

which do not stain in aniline or cotton blue dye (Plate 23 Fig. C.). In Geodina and

sometimes in Cookeina the connected longitudinal ridges are by cross ridges (Denison,

The have smooth 1965). remaining genera ascospores. PARAPHYSES.— In the tribeBoedijnopezizeae the paraphyses branch freely and the

other branches anastomose laterally to form a network enclosing each ascus. In the

tribes the paraphyses are less frequently branched and form no such network. In

two species, Desmazierella acicola and Cookeina sulcipes, some of the paraphyses have

thick-walled, bristle-like appendages which project above the hymenium. The

adaptive significance of these elements, if any, is unknown.

HAIRS.-— have with hairs which take several Many genera species excipular may

distinctive forms. The is hair most common a flexuous, hypha-like, hyaline (Plate 22

Fig. A.). Cookeina and Geodina have compound hairs composed ofbundles ofparallel,

has scales unbranched, thick-walled hyphae (Plate 23 Fig. A.). Boedijnopeziza com- posed ofexcipular hairs adhering side by side. In Thindia and Desmazierella the hairs

are simple and bristle-like, with thickened, dark brown walls.

— the ECTAL EXCIPULUM. In the tribe Boedijnopezizeae, and in the genus Geodina,

ectal excipulum is sharply differentiatedfrom the medullary excipulum and consists of rows of cells, textura globulosa to textura prismatica, with the rows perpendicular to

the the exterior of the apothecium (Plate 22 Fig. B.). In the tribe Pithyeae, and in

delimited from the genus Nanoscypha, the ectal excipulum is also sharply medullary

but and excipulum, the cells are smaller, textura angularis to textura epidermoidea, not

in In the ectal is iden- arranged rows (Plate 22 Fig. D.). Sarcoscypha excipulum easily

ectal tified, but there is a broad zone of transition to the medullary excipulum. The

is the of the cells the exterior excipulum textura porrecta with long axes parallel to of the In and the ectal apothecium (Plate 22 Fig. A.). Phillipsia, Rickiella, Aurophora

excipulum is poorly differentiated from the medullary excipulum and of textura

intricata to textura epidermoidea (Plate 22 Fig. C.). PIGMENTS.—The family Sarcoscyphaceae has apothecia in which the hymenium

is brightly colored. The pigmentation, localized in the paraphyses, consists, accord-

ing to Arpin ( 1968) of caroteinoids including: beta-carotene, lycopene, torulene,

torula-rhodine, phillipsiaxanthine, and plectaniaxanthine. The latter two are of G. Denison: Differentiation in Sarcoscyphaceae 437

particular interest because of their uneven distributionwithin the family. Plectania-

xanthine in and whereas occurs Pithya, Sarcoscypha, Phillipsia, phillipsiaxanthine occurs

in Phillipsia and Cookeina.

and of SUBSTRATE.—All species of the tribe Pithyeae occur on the foliage twigs

conifers. Nanoscypha occurs on the foliage and twigs of angiosperms. Acervus, Geodina,

Pindara, and Wynnea are found on soil. The remaining genera occur on wood, usually

on wood in early stages of decay.

DISTRIBUTION.—The is divided between and family sharply temperate tropical

genera. When temperate genera, such as Pithya, are found at low latitudes, they occur

elevations where the resemble those of the at high temperatures temperate zone.

The temperate genera are: Desmazierella, Thindia, Pithya, Sarcoscypha, Pindara,

Acervus, Wynnea, and Microstoma. The remaining genera are tropical.

GROUPING INTO TRIBES

There There is a suprageneric structure to the family Sarcoscyphaceae. is a group

of the tribe Korf in which core genera, Sarcoscypheae sensu (igyi), Phillipsia occu-

pies a central position surrounded by smaller, mostly specialized genera. There are

two other small, closely related groups of genera. The tribe Boedijnopezizeae Korf

consists of three in which the asci mature in which (igyi) genera synchronously,

the apothecia are deeply cupulate, and the paraphyses form a reticulum. The

choose of four remaining group I to recognize as a new tribe, Pithyeae, consisting

genera: Pithya, Pseudopithyella, Thindia, and Desmazierella, in which the apothecia

are small, resembling those of inoperculate discomycetes. All occur on foliage of

all have similar and all in distribution. conifers; excipular tissues; are temperate

Pithyeae Denison, trib. nov.

Asci suboperculati, inapothecio singulo deinceps maturescentes; ascospori unicellulares, hyalini,

laeves; apothecia clare colorata, minuta, ad folia gymnospermi, clima temperati vigentes.

Fuckel. Type genus: Pithya

Other included genera: Pseudopithyella Seaver; Thindia Korf & Waraitch; and

Desmazierella Libert.

REFERENCES

ARPIN, N. (1968). Les carotenoides des Discomycetes: essai chimio-taxinomique. These,

in Bull. Univ. Lyon, 169 pp. (& Soc. linn. Lyon 38 (Suppl.): 169 pp. 1969).

P. Essai sur les Univ. 160 BERTHET, (1964). biotaxinomique Discomycetes. These, Lyon, pp.

W. C. Central American Pezizales. I. A the DENISON, (1965). new genus of Sarcoscyphaceae. In Mycologia 57: 649-656.

-—— American and (1971). Central Pezizales. IV. The genera Sarcoscypha, Pithya, Nanoscypha. In Mycologia 64: 609-623.

F.-E. The A of ECKBLAD, (1968). genera of the operculate Discomycetes. re-evaluation their

taxonomy, phylogeny and nomenclature. In Nytt Mag. Bot. 15: 1-191. Persoonia Vol. Part 438 6, 4, 1972

KIMBROUOH, J. W. (1970). Current trends in the classification of Discomycetes. In Bot. Rev. 36: 91-161.

R. P. In KORF, (1971). Some new discomycete names. Phytologia 21: 201-207.

R. P. & K. S. WARAITCH of In KORF, (1971). Thindia, a new genus the Sarcoscyphineae. Mycologia 63: 98-103. LE GAL, M. (1946). Les Discomycetes subopercules. In Bull. Soc. mycol. Fr. 62: 218-240.

EXPLANATION PLATES OF 24, 25

PLATE 24

FIGS. A-D. -—• A. Sarcoscypha coccinea. Vertical section through the ectal excipulum and part of the medullary excipulum. Note the hypha-like excipular hairs. X 200. — B. Cookeina sulcipes. Vertical section showing appendagedparaphysis (upper right), synchronous asci, and excipular tissues. X 200. — C. Phillipsia domingensis. Vertical section through the ectal excip-

ulum — and part of the medullary excipulum. x 200. D. Pithya cupressina. Vertical section

ectal and of through the excipulum part the medullary excipulum. X 400.

PLATE 25

FIGS. — A. section of — A-C. Cookeina tricholoma. Longitudinal a fasciculate hair. X 200. B.

Sarcoscypha coccinea. Apex of an ascus stained with Congo red. Note the nearly terminal

of the — C. position subapical pad. x 3000. Phillipsia domingensis. Apex of an ascus stained with red. Note the of the Congo lateral position subapical pad. X 3000. Persoonia Vol. 6 Plate 24 Persoonia Vol. 6 Plate 25