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FIELDIANA Botany

Published by Field Museum of Natural History

Volume 34, No. 3 August 12, 1971

Revision of the Genus Morganella (Lycoperdaceae)

PATRICIO PONCE DE LEON

ASSISTANT CURATOR, CRYPTOGAMIC HERBARIUM FIELD MUSEUM OP NATURAL HISTORY

Morganella Zeller, Mycologia 40: 650. 1948, emend. Kreisel & Dring, Feddes Rep. 74: 109. 1967. Radiigera Zeller subgenus Endoplectydium Singer, Wright & Horak, Darwiniana 12: 160. 1963.

Fructifications epigeous, 1-3 cm. in diameter, depressed-globose to turbinate; peridium double; exoperidium velutinous, furfuraceous, granular, tuberculose or spinulose, whitish, tan, flesh-colored, or deep red-brown to purplish-black; endo- peridium papery above, thickened below, flaccid, smooth or reticulate, mouth irregular, apical; gleba pulverulent, without a true capillitium, paracapillitium abundant, glebal membranes abundant, pseudocolumella sometimes present, these elements all strongly cyanophilous; subgleba compact or chambered (sometimes almost indiscernible); spores globose to broadly ovoid, smooth, verruculose to spinose, 3.5-4.5 /* in diameter.

Species usually growing on decaying wood.

Type species. Morganella mexicana Zeller, Mycologia 40: 650. 1948.

Type locality. Mexico. Specimens: Macbride, Sonora, Guaymas (NY). The material deposited by Zeller in the Morgan Herbarium at the University of Iowa cannot be found. The material from Newfield, New Jersey, Ellis 5013 (NY), cited as paratype by Zeller, has been identified by Kreisel (1967) as Morganella velutina (Berk, ex Mass.) Kreisel and Dring.

Geographic distribution. Morganella is pantropical, with most of the species, M. fuliginea, M. velutina, M. puiggarii, and M. stercoraria, occurring in tropical America; M. velutina reaches the temperate zone in eastern North America, and M. puiggarii reaches it in South America; M. fuliginea covers a wide area in tropical America, and also occurs in West Africa; M. stercoraria occurs only

Library of Congress Catalog Card Number: 73-156851 Publication 1127 27

I MAY ] 5 1972

I AY 17 1Q7? blOLOGY LIBRAE

28 FIELDIANA: BOTANY, VOLUME 34

in the West Indies; M . afra occurs in tropical Africa; M. purpurascens has a wide distribution in tropical Asia; M. samoense is known only from Samoa; M. compacta and M. subincarnata are the only two extra-tropical species M. compacta occurring only in New Zealand and M. subincarnata in the temperate zone of North America.

It appears possible to me that the Caribbean region may have been the geographical center of development of Morganella, at least of the American species, and perhaps of all of them. Morganella fuliginea and M. velutina are especially abundant in Caribbean and circumcaribbean lands.

Discussion. Morganella is easily distinguished from Lycoperdon, Calvatia, Bovista, and other members of the Lycoperdales, by its small size, habit of growing on wood, absence of capillitium, and the presence of abundant glebal membranes and paracapillitium. The peridium has two layers. The outer one is used to separate the species: it is velvety or velutinous in M. velutina and M. puiggarii; tuberculose in M . fuliginea, M. purpurascens, M. subincarnata, and M. samoense (the tubercules composed of pluricellular hyphae); granulose in M. afra; with large spines in M. compacta. The lignicolous habit is characteristic of all species of Morganella except M . stercoraria, which lives on cow dung. In the remaining Lycoper- dales, only Lycoperdon pyriforme lives on decaying wood and Lycoperdon (Bovista) acuminata in the bark of living trees; both of these species have abundant capillitium in the gleba.

Although Zeller's decision to place Morganella in the Lycoper- daceae instead of the Mesophelliaceae was, according to his own statement, "for convenience" until more was known as to the mem- branes in the gleba, he was correct, not only because of the mem- branes which are also present in the gleba of Lycoperdon, Calvatia, Geastrum, and in the family Mesophelliaecae, but also because, as he later established (Zeller, 1949), the genera of the Mesophelliaceae have three layers in the exoperidium and Morganella has only two. Kreisel (1967) accepted the fact that Morganella falls within the Lycoperdaceae in spite of its lack of capillitium, and separated it from the Mesophelliaceae because of the presence of three layers of exoperidium in the latter.

CONSPECTUS OF THE GENUS MORGANELLA

SECTION MORGANELLA

Subgleba not chambered.

PONCE DE LEON: REVISION OF MORGANELLA 29

Type: Morganella fuliginea Group Fuligineae

Exoperidium spinose or tuberculate, consisting of chains of short cells. M . fuliginea M. purpurascens Group Samoenses

Exoperidium spinose, consisting of elongated cells with irregular contours.

M . samoensis Group Velutinae

Exoperidium velvety, consisting of elongated hyphae of one or a few cells, club-shaped or irregular in contour. M. velutina M. puiggarii

SECTION SUBINCARNATA

Subgleba chambered. Type: Morganella subincarnata Group Subincarnatae

Endoperidium pitted or reticulate-areolate. M. subincarnata M. compacta Group Afrae Endoperidium smooth. M . afra M. stercoraria

KEY TO THE SPECIES

Subgleba not chambered, solid. Exoperidium tuberculate.

Endoperidium smooth M. fuliginea-

Endoperidium pitted.

The tubercles of the exoperidium consisting of

chains of short cells M. purpurascens.

The tubercles of the exoperidium consisting of chains of

elongated and irregular contoured cells M. samoensis.

Exoperidium velvety.

Exoperidium of elongated setose cells M. velutina.

Exoperidium of chains of short, quadrate cells M . puiggarii.

Subgleba chambered. Endoperidium pitted.

Exoperidium of short, subpyramidal spines M. subincarnata.

Exoperidium of elongated, acicular spines M. compacta.

Endoperidium smooth.

Exoperidium granular M. afra.

Exoperidium spinose M. stercoraria.

30 FIELDIANA: BOTANY, VOLUME 34

Morganella afra Kreisel & Bring, Feddes Rep. 74: 116. 1967.

Fructifications 1 cm. diameter, depressed-globose to pyriforme; mycelium white, fibrous; exoperidium fuscous above, lighter below, minutely granular; endoperidium light brown, thin, smooth to very minutely aerolate; gleba light brown, becoming greyish as the spores are shed; pseudocolumella and glebal membranes inconspicuous; subgleba light brown, chambered, the chambers some- times very small and few, or even absent; mouth irregular; paracapillitium hyaline, septate, rarely branched, encrusted, 3.5-6.5 /z diameter; spores light yellowish, globose, 3.5-4.5 n diameter, minutely asperulate, with a short pedicel.

Type.—Dixon 4, Yinahin, Ashanti, Ghana.

Habitat. On decayed wood.

Distribution. Tropical Africa.

Discussion. Kreisel & Dring (1967) said: "This species is quite close to M . subincarnata but is distinguished by the smooth or almost smooth endoperidium and the poorly developed glebal membranes and pseudocolumella."

Material observed. I have not seen any material of this species, but by the description of the authors, it looks like an African repre- sentative of the section Subincarnata.

Morganella compacta (Cunningham) Kreisel & Dring, Feddes Rep. 74: 116. 1967. Lycoperdon compactum Cunningham, Trans. New Zeal. Inst. 57: 195. 1926.

Fructifications 3-4 cm. diameter, subglobose or pyriforme, depressed above, compressed below into a short stem-like base; exoperidium of strong brown spines, :{ 4 mm. long, separate at the base, connivent at the apices, surrounded by a ring

L cm

FIG. 1. M. compacta (Cunningham) Kreisel & Dring. Atkinson s.n. (H.P.D.D.N.Z. 1102) York Bay, New Zealand. Type of Lycoperdon compactum Cunningham (BPI). x 2.

PONCE DE LEON: REVISION OF MORGANELLA 31

1 cm

FIG. 2. M. fuliginea (Berkeley & Curtis) Kreisel & Dring. Wright 705 Valparaiso, Cuba. Type of Lycoperdon fuligineum Berkeley & Curtis (FH). X 2.

of minute brown warts or granules, the spines partly disappearing with age; endo- peridium reticulated, the reticules bordered by the granules of the exoperidium, membranous, ochraceous, becoming brown; mouth plane, torn; gleba olivaceous; pseudo-columella small, elliptical; subgleba ochraceous, minutely cellular, often rudimentary; paracapillitium hyaline, sparsely branched or simple, septate; spores globose, 3.4-4.5 ^ diameter, apedicellate, olivaceous, finely and moderately verrucose.

Type. Atkinson s.n. York Bay, New Zealand.

Habitat. On decaying wood.

Distribution. New Zealand.

Discussion. This species differs from the others in the genus by the pronouncedly areolate surface of the endoperidium and the large exoperidial spines.

Material observed. NEW ZEALAND. York Bay, Atkinson s.n. (ex Herb. PI. Dis. Div. N. Zeal. 10140) Type (BPI) ; Cunningham s.n. (ex Herb. PI. Dis. Div. N. Zeal. 1102) (BPI).

Morganella fuliginea (Berkeley & Curtis) Kreisel & Dring, Feddes Rep. 74 (2): 113. 1967. Lycoperdon fuligineum Berkeley & Curtis, Journ. Linn. Soc. Bot. 10: 345. 1868. L. epixylon Berkeley & Curtis, Journ. Linn. Soc. Bot. 10: 345. 1868. L. astrocaryi Berke- ley & Cooke in Cooke, Journ. Linn. Soc. Bot. 15: 393. 1873. L. cu- bense Berkeley in Massee, Journ. Roy. Microsc. Soc. 1887: 722. 1887. L. confluens Patouillard, Bull. Soc. Myc. Fr. 15: 205. 1899. Borista asterospora Massee, Grevillea 17: 60. 1899. Morganella mexicana Zeller, Mycologia 40: 650. 1948.

32 FIELDIANA: BOTANY, VOLUME 34

Fructifications 1-3 cm. diameter, depressed-globose, sessile, seated on a white cord-like mycelium; exoperidium brown, lighter below, covered by minute conical tubercles or spines consisting of clusters of hyphae composed of chains of more or less isodiametric cells, often with lateral outgrowth reminiscent of a clump connec- tion; endoperidium smooth, cinnamon-buff; mouth small, torn; gleba brown at maturity, membranes present; subgleba tan, compact; pseudo-columella not well developed; paracapillitium branched, smooth, hyaline, septate, 3-4 /* diameter; spores globose, yellowish, echinulate, 3-4 M diameter, with spines of about 1 n long.

Type.— Wright 705, Valparaiso, Cuba.

Habitat. On decaying wood.

Distribution. Mexico, West Indies, Central America, Venezuela, Brazil, Bolivia, West Africa (Dring, 1964).

Discussion. This species, as Kreisel (1967) stated, includes the type of the genus, Morganella mexicana Zeller. It differs from M. velutina in its exoperidium composed of pluricellar hyphae; that of M. velutina is composed of club-shaped setae.

Material observed.— CUBA. Wright 506 (705) Type (FH); El Retire, Wright 881 (FH); Wright 508 (2^9) (type of L. epixylon) (FH); Wright 1053 (as L. cubense) (BPI); Wright s.n. (Lloyd 58724) (as L. epixylon) (BPI). BRITISH WEST INDIES. GRENADA: Thaxter 3259 (FH). GUADALUPE. Gourbevre 91 (ex Hb. Patouil- lar 1664) (FH); Dum 912 (ex Hb. Patouillar 1664) (as L. confluens) (FH); Dum 515 (ex Hb. Patouillar 1664) (FH); Juss 1089 (ex Hb. Patouillar) (FH). PUERTO RICO. Sierra de Naguabo, Britton 3125 (NY); Utuado, Earle 293 (NY); Martinez Peco & Frontera s.n. (BPI); Martinez Peco s.n. (BPI); Aibonito, Fink 1731 (as Lycoper- don ind. (BPI). COSTA RICA. Palmar, Martin & Welden 8277 (as L. subincarnatum) (NY); HEREDIA: Gdmez 3107, 3108, 3108a, (as M. velutina) (F); no location cited: Gomez 3135, (similar to 3086 and 3099) (F). MEXICO. SONORA: Macbride s.n. (type of M. mexicana) (ex Myc. Hb. of Iowa) (NY). PANAMA. Barro Colorado, Martin & Welden 7329 (as L. subincarnata) (FH); Martin & Welden 7923 (as L. subincarnatum) (NY). BRAZIL. Sao Leopoldo: Rick s.n. (FH) ; Bresadola 3 (ex Hb. Patouillar 1649) (as L. globaria) (FH) ; Theissen s.n. (1906) (as L. velutinum) (FH) ; Theissen s.n. (1905) (as L. velutinum) (FH), Theissen s.n. (as L. velutinum) (FH), Theissen s.n. (1907) (as L. velutinum), Rio Grande do Sul: Parecy Novo, Rick s.n. (1923) (as L. velutinum) (FH); Rick 51 (1928) (as L. velutinus) (FH), Rick s.n. (1928) (as L. velu- tinus) (FH); Santa Cruz, Rick s.n. (1927) (as L. velutinum) (FH); Santa Maria, Rick s.n. (1935) (as L. velutinum var. purpurea) (FH); Nova Petropolis, Rick s.n. (1923) (as L. velutinum) (FH); Rick 129

PONCE DE LEON: REVISION OF MORGANELLA

[ cm

FIG. 3. M. puiggarii (Spegazzini) Kreisel & Dring. Type of Batista puiggarii Spegazzini (LSP). X 3.

(Lloyd 24666) (BPI); Torrend 158 (Lloyd 30570) (BPI); Torrend 196 (Lloyd 30071) (BPI); Rick 5413 (Lloyd 30572) (BPI); Rick 4019 (Lloyd 22891) (as L. epixylon) (BPI) ; Rick s.n. (ex Mus. Paris) (Lloyd 22892) (BPI). VENEZUELA. Orinoque: Gaillard s.n. (ex Hb. Patouillar 1664) (FH). TRINIDAD. Cedros, Watafiels s.n. (NY). BOLIVIA. Vaca Diez, Guayaramerim, Singer B2096, B2029 (ex Fund. M. Lillo) (F); Nor-Vungus, Rio Yariza, Singer B1413 (ex Fund. M. Lillo) (F). PERU. Dpto. Loreto, Island in Amazon River opposite Iquitos. Simpson F77 (F). UNITED STATES. GEORGIA: Bartlett 1575 (FH) (BPI). FLORIDA: Dunne 3 (Lloyd 122893) (BPI). AFRICA. No location cited: Uscher s.n. (Lloyd 30569) (BPI). HUNGARY: Hollos s.n. (ex Hb. Hollos) (FH). CHINA. HAINAN: Deng 7204 (BPI); Deng 5752 (BPI- 4321) (BPI).

Morganella puiggarii (Spegazzini) Kreisel & Dring, Feddes Rep. 74: 116. 1967. Bovista puiggarii Spegazzini, Bol. Acad. Nac. Cienc. C6rdoba 11: 470. 1887. Radiigera puiggarii (Spegazzini) Singer, Wright & Horak, Darwiniana 12: 603, 1963.

Fructifications 10-15 mm. diameter, pear-shaped or globose to obovate; mycelium rhizoid-like, white; exoperidium pallid-whitish when fresh (according to Spegazzini), blackish-brown when dried, surface velutinous or somewhat tomentose; endoperidium white or pallid-whitish, not separating from the exoperi- dium or from the gleba; mouth not seen; gleba grayish-olive (according to Spegaz- zini) when fresh, milk-coffee-colored when dried, pulverulent, tramal plates radial, anastomosing; columella with a convex upper part; subgleba small, compact, of

34 FIELDIANA: BOTANY, VOLUME 34

filamentous, thick-walled elements; spores 3.4-4 /* diameter, globose, brown; paracapillitium irregular, septate, not or rarely branched, hyaline (cyanophilous), 3.5-4 n diameter.

Type.— Apiai (Apiahy), leg. Puiggari, Sao Paulo, Brazil (LPS).

Habitat. Gregarious on rotting fallen trunks in forest.

Distribution . Brazil, Paraguay.

Discussion. This species' principal character is the tricho- dermal palisade of elements in chains that integrate its exoperidium. The swollen terminal cells separate it from M . fuliginea.

This species has been considered as belonging in the genus Radiigera, subgenus Endoplectidium by Singer et al. (1963), but the lack of capillitium, and the exoperidium of two layers, place it in Morganella. It differs from Bovista, where Spegazzini put it, by the lack of capillitium.

Material observed. BRAZIL. SAO PAULO: Apiai, Puiggari s.n. Type (LSP). PARAGUAY. Anisits 369 (as Bovista puiggarii) (LSP).

Morganella purpurascens (Berkeley & Curtis) Kreisel & Bring, Feddes Rep. 74: 115. 1967. Lycoperdon purpurascens Berkeley & Curtis, Proc. Amer. Acad. Arts Sci. 4: 124. 1860. Bovista purpur- acea (Berkeley & Curtis) De Toni in Saccardo, Syll. Fung. 7: 97. 1888, sphalm. Lycoperdon tephrum Berkeley ex Masse Journ. Roy. Microsc. Soc. 1887: 723. 1887. L. lignigenum P. Hennings & E. Nyman in Warburg, Monsunia 1: 23. 1900. L. subincarnatum

1 cm

FIG. 4. M. purpurascens (Berkeley & Curtis) Kreisel & Dring. Wright (U.S.N. Pacific Exp. Expd. no. 118). Bonin Island. Type of Lycoperdon purpurascens Berkeley & Curtis (FH).

PONCE DE LEON: REVISION OF MORGANELLA 35

Peck sensu Cunningham, Gaster. Australia & New Zealand 147. 1944.

Fructifications 2-3 cm. diameter, subglobose, sessile; exoperidium brown, lighter below, covered with minute, conical tubercles consisting of chains of roughly globose cells which often collapse; endoperidium papyraceous, pitted like a thimble, light brown; mouth small, torn; gleba olivaceous; subgleba obsolete; pseudo- columella large but inconspicuous; paracapillitium flaccid, branched, hyaline; glebal membranes inconspicuous; spores 3.5-4 /* diameter, almost smooth to minutely spiny, with an oil drop inside, pedicel hyaline, collapsed.

Type.— Wright, U.S.N. Pacific Expl. Exped. 1853. Bonin Is. (K) (FH).

Habitat. On decaying wood.

Distribution. Australia, Pacific Islands, Philippines, India.

Discussion. This species is confined to Australia, India, and the Pacific Islands. The pitted endoperidium and the almost smooth spores are the principal characteristics of this species.

Material observed.— BONIN ISLAND. Wright (U.S.N. Pacific Expl. Exped. No. 118) Type ex Hb. Gray (FH). NEW CALE- DONIA. LeRat 117 (as L. mixtum) (Patouillar 1696) (FH). INDIA. Ceylon: Fetch 2983 (Lloyd 37967); Fetch s.n. (Lloyd 51760) (BPI). PACIFIC ISLANDS. Fetch s.n. (Lloyd 37957) (BPI). PHILIPPINES. LUZON: Reinking 9990 (Lloyd 30560) (BPI); Basilan Isl., Isabela, Yatess.n. (Lloyd 41527) (BPI). AUSTRALIA. Cheel s.n. (Lloyd 30561) (BPI). SAMOA. Lloyd 5039 (30563) (BPI); Lloyd 5039 (Lloyd 30562) (BPI); Lloyd 5 (5039) (Lloyd 30564) (BPI).

Morganella samoensis (Bresadola & Patouillard) P. Ponce, comb. nov. Globaria samoensis Bresadola & Patouillard, in Lloyd Myc. Writ. 1: 50. 1901. Lycoperdon samoense (Bresadola & Patouillard) Saccardo, Syll. Fungorum 17: 233. 1902.

Fructifications 3-5 cm. diameter, sessile or subsessile, obovoid or utriform; mycelium white; exoperidium furfuraceus, brown, with minute conical tubercles consisting of elongated irregular cells which later collapse; endoperidium papyra- ceous, reticulated, light brown; mouth lacerate, dehiscing irregularly; gleba olivaceous to dark brown; subgleba obsolete; paracapillitium flaccid, the threads often fused, subhyaline, 3-5 » diameter; glebal membranes present; spores globose, smooth, tawny with a yellowish center, 3-3.5 » diameter.

Type.— Lloyd 5056 (Lloyd Cat. 30584) Lycoperdon samoense n.s. (BPI).

Habitat. On wood.

36 FIELDIANA: BOTANY, VOLUME 34

1cm

FIG. 5. M. samoensis (Bresadola & Patouillard) P. Ponce. Type of Globaria samoensis Bresadola & Patouillard (BPI). X 1.

Distribution. Samoa.

Discussion. This species is very close to Morganella purpurascens, from which it differs by its large size, elongated cells of the exoperi- dium, and its smooth spores.

Material observed. SAMOA. UPOLU ISLAND: Lloyd 5056. Type of Globaria samoense (BPI). Lloyd 5016 as Lycoperdon samoense Bresadola & Patouillard ex Herb. Patouillard. Part of the type collection? (FH).

Morganella stercoraria P. Ponce, sp. nov.

Fructificationes 1-1.5 cm. diam., globosae vel pyriformes, sessiles; mycelium album, filiforme; exoperidium spadiceum vel pallido-fuscum, spinae breves, apices conniventes; endoperidium firmum, laeve; gleba lutea et leviter cinereascens; sub- gleba albido-lutea, parva, cellulosa (cellulae irregulares) ; pseudocolumella in- conspicua; paracapillitium 3-3.4 /u diam., pallido-luteum, laevis vel leviter rugosum, aliquot minute pedicellatum.

Fructifications 1-1.5 M diameter, globose or short-pyriforme, sessile; mycelium white, thread-like; exoperidium tan to light-brown, composed of small pluricellular spines connivent at the apices, sometimes persisting at maturity; endoperidium firm, smooth; mouth irregular; gleba yellowish, with a gray shadow; subgleba whitish-yellow, small, with large and irregular chambers; pseudocolumella incon- spicuous; paracapillitium rarely branched, irregular, hyaline, septate, 3-4 » diameter; spores globose, 3-3.4 M diameter, smooth to moderately rugose, with a very thick wall, some with a minute pedicel.

Habitat. On cow dung.

Type.— Fink 7-40, El Yunque, Puerto Rico (Lloyd 43945) (BPI).

Distribution. Puerto Rico.

PONCE DE LEON: REVISION OF MORGANELLA 37

Discussion. This species belongs in the section Subincarnnta. and is separated from M. subincarnata by its spores (those of the latter being spinose), the lighter color of the exoperidium, and the smooth endoperidium. It is distinguished from M. afra by the larger spines of the exoperidium, smoother spores, and larger chambers of the subgleba .

This collection, Fink 740, has been cited (Stevenson, 1936) as the probable basic material of Lycoperdon fimicola Lloyd. However, this is a nomen nudum, as there is no description of the material by Lloyd.

Morganella subincarnata (Peck) Kreisel & Dring, Feddes Rep. 74. 117. 1967. Lycoperdon subincarnatum Peck, Ann. Rep. New York State Mus. Nat. Hist., Bot. 24: 83. 1872.

Fructifications cespitose, 2-4.5 cm. diameter, depressed-globose, or short pyriform, sessile, some with a broad point of attachment on a white fibrous myce- lium; exoperidium brown to reddish-brown, composed of small, separate, sub- pyramidal tubercles or spines with their tips connivent, consisting of chains of cells, which fall at maturity; endoperidium firm, tough, pitted like a thimble; mouth 3-4 mm. diameter; gleba white at first, becoming yellowish and finally olive umber, with obviously radiating membranes, subgleba chambered, white to light brownish, occupying one-third of the total fructification, pseudocolumella distinct; paracapillitium rarely branched, hyaline, septate, 3.5-6.5 » in diameter; spores 4-5 ^ in diameter, light yellowish-brown, globose-echinulate, with a minute pedicel.

Type.— Peck 2959, Sandlake, New York (NY). Habitat. On decaying wood. Distribution. North America.

Discussion. This species is easily distinguished by the small pits outlined by acute ridges on the endoperidium, that make its surface

1cm

FIG. 6. M. stercoraria P. Ponce. Fink 7^0 El Yunque, Puerto Rico. Type. (Lloyd Cat. 43945) (BPI).

38 FIELDIANA: BOTANY, VOLUME 34

1 cm

FIG. 7. M. subincarnata (Peck) Kreisel & Dring. Peck 2959 Sandlake, New York. Type of Lycoperdon subincarnatum Peck. (NY). X 2.

appear like a thimble under the lens. It is separated from M. compacta by the large spines of the exoperidium of this species.

Material observed.— UNITED STATES. NEW YORK: Sandlake, Peck 2959 Type (NY). Peck 10 annotated by C.T. Rogerson as probably part of the type (NY); Cazenovia, Morgan s.n. (NY); Adirondacks, Lake Placid, Murrill 543 (NY); Adirondacks, Camp Kanasa, Murrill 74 (NY); Delaware Co., Arkville, Murrill s.n. segregated "A" by D.M. Dring (NY); Burnham s.n. (Lloyd 7039) (BPI); Lodenick s.n. (Lloyd 16465) (BPI); Clute s.n. (Lloyd 31353) (BPI); Schum s.n. (Lloyd 31557) (BPI); Long 1833 (Lloyd 31558) (BPI);H6. ofNYBGs.n. (Lloyd 31559) (BPI) ; Peck 8 (Lloyd 31560) (BPI); Darker s.n. (FH). MAINE: Bar Harbor, White s.n. (NY). VERMONT: Sudbury, Wright s.n. (ex Hb. Bart.) (FH); Hadley 17 (Lloyd 31544) (BPI). MASSACHUSETTS: Page 9 (Lloyd 31547) (BPI). NEW HAMPSHIRE: Jones s.n. (Lloyd 31555) (BPI); Theobald s.n. (Lloyd 31556) (BPI). NEW JERSEY: Paddock s.n. (NY). PENNSYL- VANIA: Buck Hill Falls, Delafield s.n. four collections (NY); Bilgram s.w. (Lloyd 31548-31552) (BPI); OHIO: Preston, Monroe Co., Stifler s.n. (1931, 1934) (F) (NY) ; Morgan s.n. (NY); Smith 96 (Lloyd Hb. 8671) (NY) (BPI); Smith s.n. (Lloyd Hb. 7329) (Lloyd 31534) (BPI). MICHIGAN: Bessey s.w. (Lloyd 16462), (Lloyd 16464) (BPI); Upper Tahquamenon Falls, Luce Co. Alexia s.n. (MICH.) INDIANA: Bichtel s.n. (Lloyd 14827) (BPI). WASHINGTON D.C.: Braendle s.n. (Lloyd 31545) (BPI). VIRGINIA: Schum s.n. (Lloyd 31537) (BPI); Boutlou 54 (Lloyd 31538) (BPI); Lloyd 5691 (Lloyd 31539) (BPI). WEST

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40 FIELDIANA: BOTANY, VOLUME 34

VIRGINIA: Lloyd 5691 (Lloyd 31540); Lloyd 2606 (Lloyd 31541); Lloyd 2619 (Lloyd 31542) (BPI). NORTH CAROLINA: Coker 8870 (ex Hb. U. of N.C.) (FH) ; Winston-Salem, Schallert 2304 (F). FLORIDA: Singer F 237 (Fungi Floridani) (FH) ; Thaxter 3442 (FH) . CANADA. ONTARIO :Cain 7438 (asL. timagani) (NY) ; Fowler s.n. (Lloyd 31543) (BPI); Harraly, Lake Rosseau, Harper s.n. (Field Mus. 1286998) (F). No location cited: Patouillar 5662 (FH).

Morganella velutina (Berkeley ex Massee) Kreisel & Dring, Feddes Rep. 74: 114. 1967. Lycoperdon velutinum Berkeley ex Massee, Journ. Roy. Microsc. Soc. 1887: 718. 1887.

Fructifications 3 cm. diameter, depressed-globose to pulvinate, usually with a marked umbo, sessile on a white cord-like mycelium; exoperidium mahogany color shading to tan below, darker when dry, persistent, densely velutinous, consisting of setose hyphae mostly 100 n long and clavate; endoperidium smooth, very thin; mouth small and torn; gleba umber, membranes not prominent, pseudocolumella small and flattened, not well marked; subgleba tan, compact; paracapillitium branched, smooth, hyaline, cyanophilus, septate, 3-4 p. diameter; spores globose, 3.5-4 ju diameter excluding ornament, spines up to 1 n long, often with flattened tips.

Type.—Fendler 235 and 236, Colonia Tovar, Venezuela (K).

Habitat. On decaying wood.

Distribution. Eastern North America: Florida, North Carolina, New Jersey. South America: Venezuela, Brazil.

Discussion. This species may be separated from related species in the genus by its setose exoperidium of clavate hyphae.

Material observed.— UNITED STATES. NEW JERSEY: Newfield, Ellis 5013 (as paratype of M. mexicana) (NY). FLORIDA: Alachua Co., E. West s.n. (as L. subincarnatum) (NY). VENEZUELA. Ellis (ex Hb. Fendler) (NY); Massee s.n. (NY); Hennings s.n. (Lloyd 22897) (BPI); Hennings s.n. ex type Kew (Lloyd 50776) (BPI). AMAZONAS: Cano Tucano, Rio Cauaburi, Maguire & Steyer- mark 60243 (NY) (F) ; Cerro de la Neblina, Maguire, Wurdack and Bunting 36907 (NY) (F). BRAZIL. No location cited, Rick s.n. (Lloyd 22894), (Lloyd 22895) (BPI); Pazschke s.n. (Lloyd 22896) (BPI); Rick 52 (Lloyd 25311) (BPI). RIO GRANDE DO SUL: Nova Petropolis, Rick s.n. (FH). TRINIDAD. Santa Cruz, Broadway 7037 (F).

SPECIES DUBIAE

There are two species of Lycoperdon, L. brasiliense Fries and L. pisiforme Hennings, which in the opinion of several authors can be placed in Morganella.

PONCE DE LEON: REVISION OF MORGANELLA 41

Lycoperdon brasiliense Fries, Syst. Myc. 3: 40, 1829. For this species we have only the description of Fries, as the type cannot be located. He said,

"L. brasiliense, peridio membranaceo persistente globoso punctato- scabro, ore obtuso, floccis laxis, sporidiis fuliginosis . . .

Peridio membranaceo floccido, extus tantus punctato-scabro, ut Lycogala, cui exoleto simile est . . .

Ad truncos Brasiliae. Beyrich!"

The type was collected by Beyrich in Brazil, but his specimens have not been found in any of the herbaria to which he sent his collections.

Lycoperdon pisiforme Hennings, Bot. Jahrb. 23: 556. 1897.

Bring (1964) and Kreisel (1967) have suggested that this species might be placed in Morganella, mostly because of its small size, its habit of growing from a subiculum on dead wood, and its hyaline capillitium. However, the spores from the type of this species in the Lloyd collection have a different episporium, which is thicker than that of Morganella, and its sculptures are not spines or warts, but elongated ridges as in the spores of Lycogalopsis.

Because of the characteristics observed in the episporium, the shining peridium, and the extensive white shining subiculum men- tioned in the description, it is assumed that this species may belong in the genus Lycogalopsis Fischer.

SPECIES EXCLUSA

Restudy of the type of Lycoperdon albidum Cooke (Morganella albida P. Ponce) and comparison with the type of Lycogalopsis solmsii Fischer shows these two to be identical in their clamp con- nections, in the disposition of the basidia in groups, the short sterigmata of the spores, and the constitution of the peridia. It is necessary, therefore, to place the first two names in synonomy under Lycogalopsis solmsii Fischer, as indicated by Dennis (1953) and as suggested by Bring in a personal letter to me in April, 1970.

LIST OF ACCEPTED NAMES AND SYNONYMS

Accepted Names in bold face Synonyms in italics

afra Kreisel & Bring Morganella

albina (Cooke) P. Ponce (Morganella) Lycogalopsis solmsii

42 FIELDIANA: BOTANY, VOLUME 34

asterospora Mass. (Bovista) Morganella fuliginea

astrocaryi Berk. & Cke. (Lycoperdon) Morganella fuliginea

compacta (Cumm.) Kreisel & Dring Morganella

compactum Cumm. (Lycoperdon) Morganella compacta

confluens Pat. (Lycoperdon) Morganella fuliginea

cubensis Berk. (Lycoperdon) Morganella fuliginea

epixylon Berk. & Curt. (Lycoperdon) Morganella fuliginea

fuliginea (Berk. & Curt.) Kreisel & Bring. . . .Morganella

fuligineum Berk. & Curt. (Lycoperdon) Morganella fuliginea

fuligineum Berk. & Curt, sensu Dring (Lycoperdon) Morganella afra

golungense Welw. & Curr. (Lycoperdon) Geastrum sp.

golungense Welw. & Curr. sensu Beeli. (Lycoperdon) Geastrum sp.

mexicana Zeller (Morganella) Morganella fuliginea

puiggarii (Speg.) Kreisel & Dring Morganella

puiggarii Speg. (Bovista) Morganella puiggarii

puiggarii (Speg.) Singer, Wright, & Horak (Radiigera) . .Morganella puiggarii purpurascens (Berk. & Curt.) Kreisel & Dring. .Morganella

purpurascens Berk. & Curt. (Lycoperdon) Morganella purpurascens

purpurascens (Berk. & Curt.) De Toni (Bovista) . . .Morganella purpurascens pyriforme Pers. var. tesselatum Pers.

sensu Dissing & Lange (Lycoperdon) Morganella afra

samoensis (Bres. & Pat.) P. Ponce Morganella

samoensis Bres. & Pat. (Globaria) Morganella samoensis

samoense (Bres. & Pat.) Saccardo (Lycoperdon) .... Morganella samoensis

stercoraria P. Ponce Morganella

subincarnata (Peck) Kreisel & Dring Morganella

subincarnatum Peck sensu Cunn. (Lycoperdon) . . . Morganella purpurascens

subincarnaium Peck (Lycoperdon) Morganella subincarnata

tephrum Berk, ex Mass. (Lycoperdon) Morganella purpurascens

velutina (Berk. & Curt.) Kreisel & Dring . . . .Morganella

veluntium Berk. & Curt. (Lycoperdon) Morganella velutina

LITERATURE CITED

BERKELEY, M. J. and M. A. CURTIS

1869. Fungi Cubenses. Journ. Linn. Soc. 10: 280-392.

BOTTOMLEY, A. M.

1948. Gasteromycetes of South Africa. Bothalia 4: 473-810.

COKER, W. C. and J. N. COUCH

1928. The Gasteromycetes of the Eastern United States and Canada. Univ. of North Carolina Press. Chapel Hill.

CUNNINGHAM, H. G.

1926. Lycoperdaceae of New Zealand. Trans. New Zealand Inst. 57: 195.

DENNIS, R. W. S.

1953. Some West Indian Gasteromycetes. Kew Bull. 8: 307-328.

PONCE DE LEON: REVISION OF MORGANELLA 43

DISSING, H. and M. LANGE

1962. Gasteromycetes of Congo. Bull. Jar. Bot. de L'Etat Bruxelles 22: 326-416.

BRING, D. M.

1964. Gasteromycetes of West Tropical Africa. Commonwealth Myc. Inst. Myc. Papers 98: 44-45.

FRIES, E.

1829. Systema Mycologicum 3: 40.

GARNER, J. H. B.

1956. Gasteromycetes from Panama and Costa Rica. Mycologia 48: 757-764.

HEMRICH, M. H.

1969. Etude de quelques Gasteromycetes du Rio Grande do Sul. Rev. Myc. 34 (1):3-16.

HENNINGS, P. 1897. Fungi Camerunenses II. Bot. Jahrb. 23: 556.

KOTLABA, F. and Z. POUZAR

1964. Preliminary results on the staining of spores and other structures of Homobasidomycetes in cotton blue and its importance for taxonomy. Brit. Myc. Soc. Trans. 47: 653-654.

KORF, P. R.

1952. Two techniques for showing spores markings in operculate Discomycetes. Trans. Brit. Myc. Soc. 35 (1): 224-225.

KREISEL, H. and D. M. DRING

1967. An emendation of the genus Morganella Zeller. Feddes Rep. 74 (2): 109-122.

LLOYD, C. G.

1902. The Genera of Gasteromycetes, 1-24.

1905. The Lycoperdaceae of the United States. Myc. Notes 20: 221-238.

1905. The Lycoperdaceae of Australia and New Zealand, 1-42.

1916. Lycoperdon albidum. Myc. Notes 24: 582.

1924. The Genus Lycogalopsis. Myc. Notes 71: 1244.

PATOUILLAR, N.

1902. Champignons de la Guadalupe. Bull. Soc. Myc. Fr. 18: 175-300.

SEAVER, F. J. and C. E. CHARDON

1926. Scientific Survey Porto Rico and Virgin Islands (Mycology) 8: 180.

STEVENSON, J. A. and E. K. CASH

1936. The new fungus names proposed by Lloyd. Bull. Lloyd Library 35, Myc. Ser. 8: 185.

SINGER, R., J. E. WRIGHT and E. HORAK

1963. Mesophilliaceae and Cribbeaceae of Argentina and Brazil. Darwiniana 12 (3): 598-611.

WELWITSCH, F. and F. CURREY

1870. Fungi Angolenses. Trans. Linn. Soc. London 26: 269.

44 FIELDIANA: BOTANY, VOLUME 34

ZELLER, S. M.

1948. Notes on certain Gasteromycetes, including two new orders. Mycologia 40: 639-668.

1949. Key to the orders, families, and genera of Gasteromycetes. Mycologia 41:36-58.

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