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THE GASTEROMYCETES

OF THE

EASTERN UNITED STATES
AND CANADA

THE UNIVERSITY OF NORTH CAROLINA PRESS
CHAPEL HILL, N. C.

THE BAKER AND TAYLOR CO.
NEW YORK

OXFORD UNIVERSITY PRESS
LONDON

MARtJZEN-KABUSHIKI-KAISHA
TOKYO

THE GASTEROMYCETES

OF THE

EASTERN UNITED STATES
AND CANADA

BY

WILLIAM CHAMBERS COKER

Kenan Professor of Botany, University of North Carolina

AND

JOHN NATHANIEL COUCH

Assistant Professor of Botany, University of North Carolina

WITH 123 PLATES

u C

>. LIBORTASi?

CHAPEL HILL
THE UNIVERSITY OF NORTH CAROLINA PRESS

1928

COPYRIGHT, 192 8, BY
THE UNIVERSITY OF NORTH CAROLINA PRESS

PRINTED IN THE UNITED STATES OF AMERICA
BY THE WAVERLY PRESS, BALTIMORE, M D.

CONTENTS

Preface ix

Artificial Key to the Families 1

Family Phallaceae 3

Family H ysterangiaceae 15

Family Hymenogastraceae 24

Family Secotiaceae 53

Family Lycoperdaceae 58

Family Arachniaceae 144

Family Sphaerobolaceae 146

Family Tylostomataceae 149

Family Sclerodermataceae 160

Family Xidulariaceae 173

Family Calostomataceae 185

Literature 194

Index 197

34

PREFACE

It seems to be generally felt that there is need for a handbook of Gasteromycetes.
There has been in this country no comprehensive treatment of the species as a whole
and in logical order with convenient keys, and as a result this highly interesting and
varied group has been neglected by many who would otherwise enjoy their study.
Lloyd, Morgan, and Peck have been the principal recent students of the Gasteromycetes
and their publications are indispensable. Morgan included all the families except
Hvsterangiaceae and Hymenogastraceae, which were nearly unknown in America at that
time, and established several new genera. Lloyd was occupied with these plants for
years and has published a large number of good photographs, several monographs, and
many notes. His collections, which are the largest in America, are available, and so are
Peck's, but so far as this group is concerned Morgan's, unfortunately, have been almost
destroyed.

In preparing this book we have studied all available species microscopically and
have compared types and authentic specimens in many cases. To this end the senior
author has visited the following herbaria: Persoon's at Leiden, Bresadola's at Stock-
holm, Kew Herbarium, New York Botanical Garden Herbarium, Curtis Herbarium,
Schweinitz Herbarium, Lloyd Herbarium, and the Pathological and Mycological
Herbarium at Washington. This last contains Michener's herbarium which includes
many important plants from the Schweinitz Herbarium. To a number of correspond-
ents we are indebted for generous gifts of material. Among these are Dr. W. A. Murrill,
Mr. C. G. Lloyd, Miss Edith Cash, Mr. John Dearness, Dr. Homer D. House, Mr. H. C.
Beardslee, Dr. Roland Thaxter, Mr. F. W. Pennell, Miss E. M. Wakefield (Kew),
Professor N. Patouillard (Paris), Dr. G. F. Weber, Dr. C. L. Shear, Mr. Elam Bar-
tholomew and Mr. G. H. Cunningham (New Zealand).

Other members of the botanical staff of this University have been of much assist-
ance. Most of the microscopic work has been done by Dr. Couch and the drawings
have been made by him and Miss Alma Holland, the latter having done all the ink work
and drawn many of the spores. Nearly all the written matter and most of the photo-
graphs are by the senior author and the microscopical work and drawings have been
carefully supervised.

The literature has been referred to throughout this work under families, genera
and species, but for convenience we list on page 194 the more important general works
dealing with the entire group or with several families.

w. c. c.

Chapel Hill, N. C.
June, 1927.

ARTIFICIAL KEY TO THE FAMILIES

Plants emerging at maturity from a soft volva or "egg;" the spores bornein a slimy, brown, bad-smelling

liquid at the top of a stalk or net or several columns Phallaceae (p. 3)

Not as above

Plants with a distinct firm or gelatinous stalk which carries the spore-bearing sac, the latter in our
representatives having a distinct apical mouth through which the spores escape as a dry
powder.

Stalk firm and fibrous; mouth not red Tylostomataceae (p. 149)

Stalk gelatinous; mouth red Calostomataceae (p. 185)

Plants without a distinct, terete or gelatinous stalk; if a stalk is present it is thick and expands

gradually into the swollen spore-bearing part above.

Plants small, shaped like cups or tumblers or subspherical, at maturity opening in most of the

species over the entire top by the collapse of a veil (or rarely by the crumbling of the

peridium) to expose a number of small flattened peridioles with hard coats which

contain the spores, the whole looking like a little nest containing eggs.

Nidulariaceae (p. 173)
Plants growing on wood; very small, spherical, exploding at maturity and throwing out to some

distance a single minute peridiole Sphaerobolaceae (p. 146)

Not as above

Peridium without a distinct outer layer that falls away or splits; at maturity opening at

the top by irregular lobes, or by an irregular tear, or by crumbling or rotting

away. Elongated threads (capillitium) not present among the spores.

Gleba not formed of hollow chambers, but of sterile plates cutting out irregular blocks

which are stuffed with the fertile tissue; at maturity crumbling into a dusty

powder Sclerodcrmataceae (p. 160)

Gleba formed (at least when young) of hollow chambers which are lined with the
hymenium.
Peridium rotting away after maturity, the gleba (at least in species we are treat-
ing) turning into a slimy or gelatinous mass.
Tramal plates radiating from the base and not intimately onnected with

the peridium Hysterangiaceae (p. 15)

Tramal plates not radiating from the base and intimately connected with

the peridium Hymenogaslraceae (p. 24)

Peridium crumbling away after maturity, the glebal chambers remaining intact

and falling apart as fine sand-like particles Arachniaceae (p. 144)

Peridium without a distinct outer layer; at maturity opening ventrally by separating
around the stem and expanding more or less; capillitium not present.

Secoliaceae (p. 53)
Peridium with a distinct outer coat (Calvatia rubro-flava is an exception) which falls away
in flakes or wears away by degrees at maturity, or dehisces equatorially, or (in
Ceaster) splits into star-like rays to expose the thin, pliable or (in Calvatia)
fragile and brittle inner peridium; gleba composed of small, hollow chambers
lined with the hymenium (obscure in Disciseda); spores mixed with a true
capillitium of long, slender, branched or unbranched threads (broken up into
short sections at maturity in Disciseda) and escaping as dust through a definite
(except in Calvatia) pore or slit. (In several small species of Lycoperdon the
outer peridium is very thin, obscure and persistent, and in Myriostoma there
are several pores) Lycoperdaceac (p. 58)

PHALLACEAE*

Plants consisting at first of a white, elastic, oval or subspherical "egg," which
consists of three coats, the central one soft and gelatinous, which break at maturity to
allow the elongation and exposure of the curious, spongy, and in some species brightly
colored receptaculum of various shapes which bears above either on itself or on a
specialized appendage the slimy, deliquescing gleba which contains the spores, and which
in nearly all cases has a very strong and offensive odor by which insects are attracted to
scatter the spores. Basidia, so far as known, club-shaped (very peculiar in Anlhurus),
with several (4-8) sessile (no mucro), smooth, minute, apical spores.

The family is divided into two subfamilies or by some authors (as Corda and
Fischer) into two distinct families which are separated by the position of the gleba
and by other important microscopical characters. These subfamilies may be simply
defined as follows :

Gleba (and spore slime) borne on the inner side of the receptaculum Clathreae

Gleba borne on the outer surface of the receptaculum Phalha?

The unique genus Phallogaster , placed by Morgan, its author (1. c, 15: 171, pi.
11. 1892), and by Thaxter (Bot. Gaz. 18: 117, pi. 19. 1893) in the Phallaceae,
but differing from them in the absence of a volva and in other ways, is treated by Fischer
in the Hysterangiaceae (1. c, Pflanzenfamihen, p. 307), and we are following him in this.
PkaUogasler, Protubera, and Protophallus mark a distinct advance toward the Phalloids
and indicate their descent from the Hysterangiaceae. The last mentioned genus is,
in fact, so Phalloid-like that it is placed by its author among them. It is distinctly
intermediate between the two families and if put in the Phalloids, their family character-
istics would have to be modified.

As the number of our species of Phalloids is small, we key all of them under the
family, and for the convenience of students we include all species described from the
United States.

Literature

Atkinson. Origin and Taxonomic Value of the Veil in Diclyoplwra and Ilhyphallus. Bot. Gaz. 51:

1, pis. 1-7 and one text figure. 1911.
Bambeke. La Relation du Mycelium avec le Carpophore chez Ithy phallus impudicus et Mutinus

caninus. M6m. Acad. Roy. Belgique, 2nd ser., 2: 1910.
Bambeke. Recherches sur certain elements du mycelium d'lthypliallns impudicus. Bull. Acad. Roy.

Belgique, No. 6, p. 280. 1914.
Burt. A Xorth American. ■Inthiirus. Mem. Boston Soc. Nat. Hist. 3, No. 14: 1894.
Burt. The Phalloideae of the United States— I. Bot. Gaz. 22: 273, pis. 11 and 12. 1896.
Burt. The Phalloideae of the United States— II. Bot. Gaz. 22: 379. 1896.
Burt. The Phalloideae of the United States— LU. Bot. Gaz. 24: 73. 1897.

• A peculiar new plant, Claustuta Fischeri, recently described from New Zealand (Curtis, 1. c.) is obviously related to this
family, but if included would necessitate modification of the family characters. The receptaculum is a complete hollow sac
bearing the hymenium on the entire inner surface. The spores are larger than in typical Phalloids and have a very distinct
mucro. There is also no obvious slime.

4 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Conard. The Structure of Simblum sphaerocephalum. Mycologia 5: 264, pis. 96 and 97. 1913.

Corda. I.e., 5: 70, pi. 6, fig. 49, pi. 7 and pi. 8, fig. 51.

Cunningham. Clathrus cibarius, the "Bird-cage Fungus." New Zealand Journ. Sci. and Tech. 5:

247, figs. 1-4. 1922.
Cunningham. Aseroe rubra, an Interesting New Zealand Phalloid. New Zealand Joum. Sci. and

Tech. 6: 154, figs. 1-4. 1923.
Curtis, K. M. The Morphology of Clauslula Fischeri, gen. et sp. nov., a New Genus of Phalloid Affinity.

Ann.Bot.40: 471, pi. 15 and 6 text figs. 1926.
Fischer. Beitrage zur Morphologie und Systematik der Phalloideen. Ann. Myc. 8: 314, pi. 5. 1910.
Fischer. Unters. z. vergleichenden Entwicklungsgeschichte u. Systematik d. Phalloideen — I. Denk-

schriften d. Schweiz. Naturforschenden Gesellschaft 32: 1. 1890; II. Ibid 33: 1. 1893;

III. Ibid 36: 2. 1900.
Gerard. Additions to the U. S. Phalloidei. Bull. Torr. Bot. Club 7: 29. 1880.
Gerard. Correlation between the Odor of Phalloids and their Relative Frequency. Bull. Torr. Bot.

Club 7: 30. 1880.
Holttum. Observations on the expansion of Dictyophora indusiata Desv. Gard. Bull. Straits Settle-
ments 3: 281. 1924.
Lloyd. The Phalloids of Australasia, figs. 1-25. Cincinnati, 1907.

Lloyd. Concerning the Phalloids. Myc. Notes No. 24: 293, figs. 131-135 and pis. 91-93. 1906;
No. 26: 325, figs. 160-163 and pis. 112-121. 1907; Nos. 28-30: 349, figs. 167-223. 1907-08.
Lloyd. The Phalloids of Japan. Myc. Notes No. 3 1 : 400, figs. 236-242. 1908.
Lloyd. Synopsis of the Known Phalloids, figs. 1-107. Cincinnati, 1909.

Lohwag. Der tjbergang von Clathrus zu Phallus. Arch. f. Protistenkunde 49: 237, with 7 figs. 1924.
Long. The Phalloids of Texas. Joum. Myc. 13: 102, pis. 102-106. 1907.
Moller. Brasilische Pilzblumen. Jena, 1895.
Penzig. Ueber Javanische Phalloideen. Ann. Jardin Bot. de Buitenzorg, ser. 2, 16: 133, pis. 16-25.

1899.
Schroeter. In Cohn's Krypt.-Fl. Schlesien 3, pt. 1: 687. 1889.

For other literature see p. 194 and Fischer as cited there.

Key to the Phalloids of the United States*

Clathreae

Receptaculum composed of a stout, netted globe Clathrus cancellatus (p. 6)

Receptaculum composed of two to five stout columns fused only above Clathrus columnatus (p. 5)

Receptaculum composed of a relatively short stalk which divides above into three or more arms which
are simple, widely separated and united at their tips, the gleba enclosed in the apical part of

the arms Coins Schellenbergiae (p. 7)

Receptaculum composed of a single distinct, relatively long stalk bearing a subglobose, netted, spore-
bearing part on the end; red above Simblum sphaerocephalum (p. 7)

Receptaculum as above, but color yello w all over Simblum texense

Receptaculum stalked as in Simblum, but the apical spore-bearing part composed of several (usually
six) short, hollow arms, incurved and meeting above at first and enclosing the gleba; stalk
white Antkurus borealis (p. 8)

PhaLleae

Gleba (and spore slime) borne on the upper part of the stalk itself

Stalk 10-17 cm. long, tapering gradually from the center upward to a rounded point; spores 4.7/i

long Mulinus Curlisii (p. 10)

Stalk 6-8 cm. long, nearly cylindrical except for the tapering base (at times) , the tip rather abruptly

rounded and the spore-bearing part more or less sharply delimited from the sterile part;

spores 3.7—1.8ju long Mutinus Ravenelii (and M. caninus, if that is different.) (p. 9)

• All except Simblum texense are found east of the Mississippi.

PH.YLLACEAE 5

Gleba borne on a pendent cap hanging from the stem tip
An obvious netted veil absent

Outer surface of cap strongly pitted by reticulated plates Itliyphallus impudicus (p. 12)

Outer surface of cap not pitted but minutely granular; stalk white /. Ravenelii (p. 11)

Outer surface not pitted but nearly smooth to longitudinally rugose; stalk red

/. rubicundus (p. 13)

An obvious netted veil-like indusium pendent from the stem apex and exposed for some distance

below the cap Dictyophora duplicata (p. 13)

CLATHRUS Mich.

Receptaculum without a single distinct stem, but composed of an inflated hollow
network or of several curved columns meeting and fused above; in all cases having the
spore-bearing gleba attached to the inner side above as a slimy mass. Odor offensive.
Only a few species are known and but two have been found in the eastern United States.

For other species see Lloyd, Syn. Known Phalloids, p. 54. For literature see under
the family.

There is evidence that both of the following species are poisonous. Farlow reports
the killing of hogs within twelve to fifteen hours by C. columnatus in North Carolina
(Bot. Gaz. 15:45. 1890).

Clathrus columnatus Bosc
Laternea columnata Nees

Plates 1 and 105

Eggs subspherical, about 2.5 cm. thick, bursting above into several flaps to allow
the expansion of the receptacle which consists of 2-5 (in our plants 4-5) stout, spongy,
curved and very delicate columns with separate, pointed, basal ends which remain in
the volva, the distal ends incurved and completely fused into a flat roof from which
hangs within the dark slimy mass of the spore-bearing gleba. Color of the receptacle
rosy red, the base pale to colorless ; volva watery white, attached at base by a cord-like
root. Odor about that of a stink horn, strong and fetid.

Spores (of No. 4949) smooth, elliptic, minute, 1.8-2.4 x 3.7-4.8ju.

This plant seems to be entirely southern in its range (but see Saccardo's Sylloge
7: 19) and confined to the sandy coastal plain of North Carolina southward. It was
originally described from South Carolina and is also known from the Gulf states (see
Long, Mycologia 9: 274. 1917). An interesting anomaly is shown by one of our
plants (pi. 1). There is a natural perforation on one side at the top just about the
center of one of the columns, exactly like the more numerous perforations in C. cancel-
latus and in Simblum. For development and microscopic detail, see Burt, Bot. Gaz.
22: 273, pis. 21 and 22. 1896.

Illustrations: Bosc. Mag. der Gesell. naturforschender Freunde Berlin 5: pi. 5, fig. 5. 1811.
Burt. As above.

Fischer. Pfianzenfamilien 1, pt. 1: fig. 120 B.
Lloyd. Myc. Works, pi. 92.
Lloyd. Phalloids of Australia, fig. 20.

Moller. Brasilische Pilzblumen, pi. 2, figs. 3 and 4; pi. 7, fig. 17 (as Laternea).
Nees von Esenbeck. Syst. Pilze Schw., pi. 36b, fig. 262. 1817.

6 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

North Carolina. Smith's Island. In a sandy road, December 29, 1921. Couch and Grant, colls.
(U. N. C. Herb., No. 4949).
Fayetteville. Lawrence, coll. Reported by Farlow (Bot. Gaz. 15: 45. 1890).
South Carolina. Georgetown. In sandy soil near Silver Hill Farm, Dec. 29, 1922. Coker, coll.
(U. N. C. Herb., No. 6013).
Santee Canal. Ravenel, coll. (Curtis Herb.).

Also from South Carolina by Ravenel in Fungi Car. Exs. No. 70 (N. Y. B. G. Herb, and Phil.
Acad. Herb.).
Florida. Eustis and Orange Bend. Underwood, coll. (N. Y. B. G. Herb.).
Mississippi. Trice, coll. (Curtis Herb.).

Louisiana. St. Martinsville. Langlois, coll. (N. Y. B. G. Herb.).
Porto Rico. Cook, coll. (U. S. Nat'l. Herb.).

Clathrus cancellatus Linn.
C. ruber Mich.
C. albits Mich.
C . flavescens Mich.
C. volvaceus Bull.

This beautiful species has been reported in the United States only from Georgia
(Schweinitz; Syn. Fung. Car. Sup., No. 537, as C. ruber) and Florida (Lloyd; Myc.
Notes, p. 296; Syn. Known Phalloids, p. 54) and New York (Peck; Bull. Torr. Bot.
Club 7 : 29. 1880). The last reference is in some doubt and may refer to C. columnatus,
as suggested by Lloyd (Myc. Notes, p. 150). It differs from C. columnatus in the more
globular receptaculum in the form of an open net-work. The color is red above, paler
below; base surrounded by the conspicuous white volva with a central cord-like root.
The dark, slimy, ill-smelling gleba containing the smooth, elongated spores is said to
cover the entire inner surface of the thick, spongy elements of the net-work.

Illustrations: Barla. Champ. Nice, pi. 45, figs. 5-12.
Bulliard. Champ. Fr., pi. 441.

Fischer. Pflanzenfamilien l1: fig. 129 A. Copied by Lloyd on pi. 92, fig. 7.
Lloyd. Myc. Notes, p. 326, fig. 160; Syn. Known Phalloids, fig. 70; also pi. 92, fig. 7, and pi. 112.
Micheli. Nova Plant. Gen., pi. 93 (as C. ruber).
Nees von Esenbeck. Syst. Pilze Schw., pi. 36b, fig. 261 (as C. ruber). 1817.

Jamaica. Howe, coll. (N. Y. B. G. Herb.).

Bahamas. Acklin's Island. Brace, coll. (N. Y. B. G. Herb.).

Porto Rico. Britton and Wheeler, colls. (N. Y. B. G. Herb.).

COLUS Cav. & Sech.

Fruiting bodies with a distinct hollow stalk which is seated in a smooth volva, the
wall with or without chambers, dividing above into several elongated arms which are
united at their tips and simple, or capped by a group of meshes like those of Simblum.
Gleba borne within the arms or meshes, deliquescing into a dark slime.

The original species (C. hirudinosus) and one other (C. Mulleri) are the only ones
with a netted apex, the others have simple arms which taper up and are united at their
tips only. Lloyd has separated the latter group as the genus Pseudocolus (Synopsis
of the Known Phalloids, p. 51). Through C. Mulleri the genus approaches Clathrella.
The species given below is the only one ever found in the northern hemisphere and is
known only from the type locality.

For literature see Fischer, Lloyd and Penzig as cited under the family.

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THALLACEAE '

Colus Schellenbergiae Sumstine

This species has been collected but once, and this is the only record of a Colus in
the northern hemisphere. We copy below the original description (Mycologia 8:
183. 1916):

"Volva dark-brown, smooth, globose, 2-3 cm. in diameter, breaking at maturity
into several irregular segments; receptaculum stipitate, the stipitate portion cylindric,
hollow, reticulate-pitted, white below, orange-colored above, slightly enlarged upward,
dividing into three arms; arms arched outward, united at the apex, transversely
wrinkled, cylindric or sometimes compressed so as to appear triquetrous, orange-
colored, attenuate upward, 3-6 cm. long; gleba dark green, borne on the inner side of
the arms, foetid; spores hyaline, ellipsoid-ovoid, 4.5-5.5 x 2-2.5^."

The species is near C.javanicus (See Penzig, 1. c, p. 160) and may be the same, as
no important differences appear from the descriptions. Colus Garciae Miiller from
Brazil has much the same appearance, but differs principally in the absence of chambers
in the wall of the stem, a remarkable peculiarity for a Phalloid.

Pennsylvania. Pittsburg. Sumstine, coll. (Carnegie Mus. and N. Y. B. G. Herb, and U. N. C. Herb.).

SIMBLUM Klotzsch

Receptaculum formed of a distinct, hollow, delicate stalk which is transformed
above into a somewhat larger inflated network which bears the gleba slime on the inside.
Yolva watery white; receptaculum bright colored, red (in our species) or yellow, rarely
whitish; odor offensive.

Only a few species are known and only two are North American. For literature
see under the family.

Simblum sphserocephalum Schlecht
Simblum rubescens Gerard
5. rubescens var. Kansensis Cragin

Plates 2, 3 and 105

Stalk long, club-shaped, hollow, spongy, tapering downward to a narrow attach-
ment in the bottom of the large, white, toughish, inflated volva; terminated above by
the subglobose structure of thick, anastomosing strands that holds the dark slime con-
taining the spores. Color bright red above, fading to pale below; odor strong and re-
pulsive. The entire plant is about 7-9 cm. high and the stalk about 1-1.6 cm. thick
above, its honeycombed wall 3-4 mm. thick near the top and only 1 mm. thick below.

Spores (of No. 1427) elliptic, smooth, 1.4-2 x 3.7-4.4/*.

This is a rare plant in the United States and has not been reported before from
North Carolina (for distribution see Lloyd's Myc. Notes, p. 220. 1905). It is said
to be common in South America. The only other North American species is S. texense
which differs from the present one in its yellow color and longer spores, which are
3 x 7M (see Long, 1. c, p. 112, pi. 106, fig. 11).

8 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Our plate 3 shows an abnormal variation of two plants springing from one volva
and fused at their tips. This is apparently of not rare occurrence as a similar example
is shown in Lloyd's Myc. Notes No. 24, fig. 133, from a photograph of a Brazilian plant
by Rick, and another such is illustrated by Gerard in his plate 2 (as cited below).

Illustrations: Gerard. Bull. Torr. Bot. Club 7: 8, pis. 1 and 2. 1880.
Long. 1. c, pi. 106, fig. 10.

1426. In apple orchard, October 26, 1914.

1427. In grass on campus, fall of 1902. A large number of plants.
5910. On ground in pasture, November 20, 1922.

5917. On ground in pasture by branch, November 22, 1922. Several plants.
7142. On ground in pasture, September 25, 1923.

ANTHURUS Kalch.

Phalloids with a hollow, spongy, subcylindrical stem springing from a volva and
bearing at the top several spongy arms which in youth embrace the dark gleba which
covers their inner faces and sides; at maturity either remaining connivent at the tips or
bending more or less strongly outward.

There is a difference of opinion as to the correct definition of the genera Lysurus
and Anthnrus. As originally described, in Lysurus the gleba is borne on the outside
faces of the arms, while in Anthurus the gleba is described as enclosed within the arms
in youth and covering their inner and side faces at full maturity, a dorsal zone remaining
bare. We are following Burt and Fischer (Phalloideen, part 3, p. 41) in including our
species in Anthurus. Lloyd (Syn. Known Phalloids, p. 35) and Rea (Trans. Brit.
Myc. Soc. 2: 57. 1904) would put the species in Lysurus, modifying that genus to
include certain species with the gleba on the inner surfaces of the arms.

The genus Anthurus is described by Kalchbrenner (Grevillea 9: 2. 1880) as
having the stem conspicuously flaring at the top and passing into several broadly spread-
ing arms which bear the spore slime on the inner surface. The description does not
state that the arms pass directly into the stem without a groove or abrupt change of
texture, but the original figure of the type species (A. Mullerianus) has this appearance,
and Fischer's figures of the same species are in agreement. Anthurus aseroeformis, a
better known plant, is said to have this structure. Lloyd thinks these two species may
be the same (Myc. Notes, p. 408, fig. 244).

In the eastern United States there is but one representative, and that is found in the
northern and central sections. There may be another American species, as a name
(Lysurus Texensis Ellis) has been referred to without description (Bull. Torr. Bot.
Club 7: 30. 1880), and Lloyd mentions a red Lysurus in Texas seen by Prof. Long
(Syn. Known Phalloids, p. 40; also Myc. Notes, p. 407). In Mycological Notes, pp.
586 and 647, Lloyd reports L. Mokusin from hothouses at Chico, California, where it
was probably introduced from the Orient.

Anthurus borealis Burt

We take the following description from Burt (Mem. Boston Soc. Nat. Hist. 3:
504. 1894) :

"Solitary or subcaespitose. Stipe white, clavate, divided above into 6 erect,
narrowly lanceolate, hollow arms incurved above, and with pale, flesh-colored backs

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PLATE 3

SlSIBUM SPHAEROCEPHALUM. No. 1426

PHALLACEAE 9

which are traversed their entire length by a shallow furrow having its surface con-
tinuous with the surface of the stipe; cavity of the stipe nearly closed at the base of the
arms by a thin diaphragm opening above into a closed chamber with dome-shaped
wall even on its inner surface and adherent to the arms for about one-sixth their length;
gleba brownish olive-green, supported upon the dome and closely embraced by the
arms; spores simple, olive-green, ellipsoidal, 3-4 x 1.5/x, 5-8 on septate and constricted
basidia.

"Total height of plant 10-12 cm. ; arms one-sixth of this; greatest diameter of stipe
15 cm."

As Lloyd supposes, this species is almost certainly introduced from the tropics or
south temperate regions.

The basidia as described by Burt are most remarkable. They are constricted at
intervals so as to appear like a row of beads, a form apparently unknown in other
Basidiomycetes. Individual plants vary greatly in size: in those we have seen, the
arms in the dry state varying from 1 to 3.6 cm. in length. In addition to plants seen
which are cited below, the species has been reported from Ohio, Connecticut, Mas-
sachusetts, England and Germany. (See Lloyd, Myc. Notes, pp. 183, 219, 386, 515;
Beardslee, Ann. Rept. Ohio St. Acad. Sci. 9: 19. 1901; Sumstine, Ohio Naturalist
6: 474. 1906; Stockberger, Ohio Naturalist 6: 517.)

Lloyd thinks it very probable that L. anstraliensis (see figure by Rea in Trans.
Brit. Myc. Soc. 2: pi. 3, fig. A. 1904), L. Clarazianus and L. Gardneri are all the
present species (Myc. Notes, p. 594).

A variety of this species ( Klitzingii Hennings) has been described from Germany
(Hedwigia 41: 167. 1902), differing only in paler color and absence of green in the
gleba (see Rea, as cited under the genus).

Illustrations: Burt. I.e., pis. 49 and 50.

Lloyd. Myc. Xotes, p. 386, fig. 219; p. 513, fig. 510.
Volkert. Mycologia 4: pi. 68, fig. 8. 1912.

New Jersey. Ramsay. In a flower garden. Lutz, coll. (N. Y. B. G. Herb.).
New York. Columbia University campus. Griffiths, coll. (N. Y. B. G. Herb.).

BlackweU's Island. In a mushroom bed. Dr. Baker, coll. (N. Y. B. G. Herb, and U. N. C. Herb.).

New York Botanical Garden. In a flower bed. Boynton, coll. (N. Y. B. G. Herb.).
Pennsylvania. Sumstine, coll. (Herb. Carnegie Mus., Pittsburgh, Pa., and U. N. C. Herb.).

MUTINUS Fr.

Receptaculum formed of a distinct, delicate, hollow stalk as in Simblum, but
differing in the spore slime being borne on the outside of the upper part of the stalk
itself, which is smooth and more or less pointed, the tip often perforated. Color rosy
red above (under and below the deep olive slime), fading downward. Volva soon col-
lapsing against the base of the stalk.

For literature see under the family and species.
Mutinus Ravenelii (B. & C.) E. Fischer

Plates 1, 4 and 105

Plants about 6-8 cm. high, the ample volva 2.5-3.5 cm. long; stalk up to 1.3 cm.
thick in center and typically tapering a little downward, again nearly cylindrical, the

10 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

spore-bearing part about 2-2.5 cm. of the apex, more or less abruptly marked off from
the sterile part by the narrower, radially elongated, definitely one-layered compartments
of the wall. Color bright rosy red above (under and below the slime), fading down-
wards; apex with or without a small opening; odor strong and offensive. Wall of the
stalk about 3 mm. thick in center and 1.5 mm. thick near base; that of the spore-bearing
part about 2 mm. thick below and 1 mm. thick above; walls of the stalk compartments
55-75/1 thick, with 3-4 layers of cells, those of the spore-bearing part 90-95/1 thick,
with about 5-7 layers of cells.

Spores (of No. 741) smooth, elliptic, 1.6-2.2 x 3.7-4.8/1.

This is more common with us than M. Ciirtisii. For treatment of its development
see Burt in Ann. Bot. 10: 343, pis. 17, 18. 1896; see also Bambeke, Mem. Acad. Roy.
Belgique, 2nd. ser., 2: 1. 1910. Lloyd considers M. Rawnelii as different from M.
caninus, both occurring in the eastern United States (Myc. Notes No. 24: 300; No. 26:
325; Syn. Known Phal., p. 28). Burt considers the two species the same (1. c, p. 344).
Lloyd's interpretation would probably refer the left hand plant on our pi. 4 to M.
caninus, the two right hand plants to M. Ravenelii. We cannot find any important
difference in structure among the forms we have collected. Mutinus caninus is common
in Europe.

Illustrations: Long. Journ. Myc. 13: pi. 104 (as M. caninus). 1907.

Lloyd. Myc. Notes No. 24, fig. 135; No. 28, fig. 183.

Marshall. Mushroom Book, pi. opposite p. 136 (as M . caninus).
For illustrations of the European M. caninus see :

Fischer. 1. c, fig. 142 A-E.

Hollos. 1. c, pi. 1, figs. 3-11.

Long. 1. c, pi. 104, fig. 9.

Nees von Esenbeck. Syst. Pilze Schw., pi. 36b, fig. 260. 1817.

Sowerby. Engl. Fungi, pi. 330.

22a. By stone wall near Battle's Park, October 24, 1902. Spores 1.4-2 x 3.7-4A)i.

24a. Old Chapel Hill collection without label. Spores smooth, 1.6-2.2 x 3.7-4.8/1.

741. On ground near stream, September 12, 1913.

1312. Near brook just above Meeting of the Waters, September 17, 1913. Spores 1.2-2.2 x 3.7-4.6/1.

1729. In low place by branch, September 10, 1915.

1925. In grass in front of Davie Hall, October 25, 1915. Spores 1.6-2.2 x 3.7-4.8/1.

Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb., as M. elegans).
Also collection from Pennsylvania by Sumstine. (Herb. Carnegie Mus., Pittsburgh, Pa., as M. caninus,
and U. N. C. Herb.)

Mutinus Curtisii (Berk.) E. Fischer
M. bovinus Morgan
M . elegans Mont.

Plates 5, 6 and 105

Plant about 10-17 cm. high, the white volva at first spherical then elongating and
on rupturing at the apex collapsing against the stalk which is almost cylindrical for the
first third and up to 2.5 cm. thick, then tapers gradually upward to the blunt point;
spore-bearing (slimy) part composing about 3-5 cm. of the apical end, similar in super-
ficial appearance to the rest of the stem except for the slime; color bright rosy red under
and below the slime, gradually fading to watery white or flesh color below; apex per-
forated by a small opening; volva rooted by a strong cord; odor of the brownish slime
very strong and offensive.

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PLATE 5

MrTiMs Curtisii. No. 2421

PHALLACEAE 11

Spores (of No. 5113) smooth, elliptic, 2-3 x 4-7/i. Wall of the very hollow stem
about 2-3 mm. thick and composed of one or two layers of small thin-walled chambers
which are irregularly isodiametrical below and change gradually to radially elongated
and definitely to a single layer in the slime-bearing part. Walls of the chambers of
the stalk 90-95/1 thick, with only 3-4 layers of cells; of the spore-bearing part 55-75/1
thick with 3-4 layers of cells.

The plants usually appear singly in woods or groves.

Illustrations: Hard. Mushrooms, pi. 56 and fig. 453.
James. Bull. Torr. Bot. Club 15: pi. 86. 1888.
Lloyd. Syn. Known Phalloids, fig. 24 (as M. elegans).
Lloyd. Myc. Notes No. 28: fig. 182 (as M. elegans).
Morgan. Journ. Cin. Soc. Nat. Hist. 11: pi. 3 (as M. bovinus). 1889.

2421. In wood mold, Battle's Park, July 24, 1916. Spores smooth, elliptic, 2-2.8 x 4-7/i.
51 13. In rotting leaves, May 16, 1922.

Asheville. Beardslee.

Blowing Rock. Seen by the authors.

ITHYPHALLUS Fr.

Volva and stalk as in Mutinns, but the spore slime is borne on the outside of a thin,
pendent, campanulate membrane or cap which is free from the stem except at the tip.
Between this cap and the stalk above and between the volva and the stalk below another
delicate, white membrane or veil is obvious (at least in /. Ravenelii).

We are following Fischer and Atkinson in placing in this genus Phallus Ravenelii,
P. impudicus, and P. rubicundus. Atkinson has shown that the membranous veil
of this genus is not homologous with the veil-like indusium of Dictyophora.

For literature see under the family and species.

Ithyphallus Ravenelii (B. & C.) E. Fischer
Dictyophora Ravenelii (B. & C.) Burt.

Plates 7-9 and 105

Stalks usually 10-16 cm. high, tapering gradually upward or nearly equal, about
1.7-2.5 cm. thick, springing from an ovate egg or volva, which is pinkish, tough, thick,
wrinkled below and connected with the earth and with other eggs by purplish pink
strands or roots which grow out from the base. The eggs are large and just before
rupturing may reach a height of 5 cm. and a thickness of 3.5 cm. The cap or pileus is
conical and is attached around the raised white ring which terminates the stem. The
upper part of the membranous veil is concealed beneath the cap while the lower half
remains in the volva around the base of the stem. At times parts or rings of the veil
may be torn loose in expanding and cling to the stalk so as to be visible beneath the cap.
Surface of the cap minutely granular, not veined or honeycombed, covered at first by
the dark, bad-smelling slime of the deliquescent gleba.

Spores (of No. 41a) smooth, elliptic, 1.2-1.8 x 3.7-4.4/1.

This interesting species is generally found growing in soil containing rotten wood
such as old wood piles and rotting trash piles and often appears in large numbers from
one colony. For discussion of sclerotia in this species, see Overholts, Mycologia 17:
109, pi. 11, figs. 5, 6. 1925.

12 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustrations: Atkinson. 1. c, pi. 2; pi. 3, fig. 7; pi. 4, fig. 10; pi. 6, fig. 14.
Hard. Mushrooms, figs. 447^149.
Lloyd. Syn. Known Phalloids, figs. 7 and 8.
Lloyd. Myc. Notes No. 28: fig. 168.
Overholts. As cited above.

Patterson and Charles. U. S. Dept. Agric. Bull. 175: pi. 36, figs. 2 and 3 (as Diclyophord). 1915.
Peck. Bull. Torr. Bot. Club. 9: pi. 25. 1882.
Scofield. Minn. Bot. Survey 2: pis. 29-31. 1900.

41a. Chapel Hill. No other data.

618. In pile of chips and trash in road, October 24, 1912.

649. Same spot as No. 618, October 31, 1912.

Asheville. Beardslee.

New York. NewRochelle. Seaver, coll. (N. Y. B. G. Herb.).

Ithyphallus impudicus (L.) Fr.

Plate 10

This is a rare plant east of the Mississippi and, except for doubtful early records,
as Schweinitz and Curtis, seems to have been recorded only twice in that territory.
The American form seems to have consistently a pink volva which is elongated or egg-
shaped and may be best considered as I. imperialis or as a variety of /. impudicus.
The latter is said by some to have in Europe a round, white volva (see Lloyd, Myc.
Notes, p. 327 and 328, also p. 508), but Rea (British Basidiomycetae, p. 23) says the
volva is round or oval and Hollos shows one with an oval, very pink volva. We have
not seen the plant alive and take the following measurements from Dr. W. B.
McDougall.

Buttons oblong or egg-shaped, pinkish, much like those of /. Ravenelii, 3-4.5 x
3.5-6 cm.; expanded plant 7.5-15 cm. tall; cap up to 4.5 cm. long, deeply reticulated
with large chambers, which are at first filled with olivaceous slime, apically attached
(in the plant we have) to a broad (1.3 cm.) white expansion of the stem apex which is
not perforated; a thin, membranous, non-perforated veil is present between the cap
and the stem and a thicker basal portion surrounds the stem attachment in the volva
base. Odor very putrid.

Spores (of plant from Urbana, 111.) color of the slime, 1.4-1.7 x 3-3.7/a, short-
elliptic, smooth. Spores of a typical plant from England in the New York Botanical
Garden (Kew Gardens; Murrill, coll.) are 1.3-2 x 3.2-4.2/1.

Illustrations: Bambeke. Bull. Acad. Roy. Belgique, No. 6, pis. 1-2. 1914.
Bambeke. Mem. Acad. Roy. Belgique, 2nd. ser., 2: pis. 1-3. 1910.
Dumee. Nouvel Atl. Champ., 2nd ser., pi. 59.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 2. Arkiv fur Botanik 17: 1921.
Hollos. PI. 1, figs. 1-2, 12-15.
Lloyd. Myc. Works, pi. 114.
McDougall. Trans. 111. Acad. Sci. 15: fig. 4. 1922.
Nees von Esenbeck. Syst. Pilze Schw., pi. 36. 1817.
Patterson and Charles. U. S. Dept. Agric. Bull. 175: pi. 37, fig. 2. 1915; also Farmer's Bull.

796, fig. 19. 1917.
Schaeffer. Der Gichtschwamm mit griinschleimigem Hute, pis. 1-5. 1760.
Schaeffer. Fung. Bavar., pis. 196-198.
Tulasne. Fungi Hypogaei, pi. 21, fig. 10.

PLATE 6

Mutinus Curtisii. Xo. 5113

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PHALLACEAE 13

Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).

Colorado. Seavcr, coll. (N. Y. B. G. Herb., as P. impcrialis). Seaver says this rosy plant is common

in the West and is a rosy form of /. impudicus.
Minnesota. Elk River. Baitey, coll. (U. S. Nat'l. Herb., as P. Ravendii).
Kansas. Lawrence. Det. by Underwood. (U. N. C. Herb.)

Ithyphallus rubicundus (Bosc) E. Fischer

We have not seen this and adapt the following from Long (I.e., p. 109):

Eggs white, solitary or in groups of 2-6 individuals from a common mycelium,
when in groups usually one large plant surrounded by small ones, ovate to globose,
2-3 cm. tall by 1-3 cm. thick. Stipe cylindric-fusiform to fusiform, hollow, scarlet,
9-15 cm. tall by 1.5 to 2.5 cm. in diameter, walls of several chambers thick, which open
onto outer and inner surfaces of stipe as pits; chambers isodiametric, pseudo-paren-
chyma tous; apex perforate or imperforate, but usually perforate as the plant ages, by
scarlet top of apex falling entirely off of plant; joined to pileus by a narrow irregular
scarlet collar or ring. Pileus conic, smooth or rugose, scarlet, sometimes extending
below gleba into a narrow sterile border, whose edges are finely crinkled to dentate,
pseudo-parenchymatous, 1-2 cm. wide to 2-3 cm. tall. Gleba at first isabella color,
becoming a dirty yellowish brown when deliquescing, odor very foetid. Veil wanting
or when present, membranous floccose, white beneath pileus or in bands or patches on
the stipe or clinging to stipe within volva as in /. impudicus. Spores oblong, 2 x 4/j.
In lawns and open grassy places, Austin, Tex., April, May, and Nov., 1900, or in old
sandv fields near rotting oak stumps and along fences in sandy soil. Denton, Tex.,
Nov. to Jan., 1902 and 1903.

Lloyd thinks that Phallus aurantiacus Mont, is the same as this (Myc. Notes, p.
330 and Syn. Known Phalloids, p. 14).

Illustrations: Bosc. Mag. Gesell. Naturf. Freunde, Berlin, 5: pi. 6, fig. 8. 1811.
Long. Journ. Myc. 13: pi. 104, fig. 5; pi. 105. 1907.
Lloyd. (Copied from Long.) Syn. Known Phalloids, fig. 5.
Lloyd. (Copied from Long.) Myc. Works, pi. 116, figs. 1-3.

South Carolina. Bosc, coll. Type locality.
North Carolina. Reported by Curtis.

DICTYOPHORA Desv.

With the characters of Ithyphallus except that there is a large campanulate, veil-
like, netted indusium that is attached to the stem tip under the cap and extends far
below it. Atkinson has shown (1. c.) that this indusium is a distinct organ and not
homologous with the short, membranous veil of Ithyphallus. The species here included
is the only one known in the United States.

For literature see under the family.

Dictyophora duplicata (Bosc) E. Fischer

Plates 11-14 and 105

Our largest and most massive Phalloid, arising from a large egg which is subspheri-
cal, ovate or sometimes flattened, about 4-4.5 x 4.5-5 cm., when flattened up to 7 cm.
broad and 5 cm. high, white and plicate below as in a peeled orange, the upper half
smooth, pale flesh color to deep fleshy brown; in the center below is given off a large,

14 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

fleshy root, and sometimes one or two smaller, more lateral ones. The expanded plant
may reach a height of 17 cm. with the fertile, pendent, apical cap about 5 cm. long and
broad, its outer surface strongly chambered by anastomosing plates, over which the
brownish olive slime is spread. Between the cap and the stem, and hanging from the
top, is a beautiful net-like veil (technically the indusium) of a light rosy pink color that
extends below the cap for about 3-5 cm., the perforations being rather regular and
about 1-2 mm. broad except towards the margin where they become much smaller.
Stalk about 4.5 cm. thick, nearly cylindrical, very hollow and with chambered walls;
between its base and the volva is a thick, brownish yellow slime which is separated from
the stem by a thin membrane. The odor of the dark spore slime above is offensive but
not nearly so much so as in the species of Mutinus, being weaker and not so distressingly
fetid.

Spores (of No. 5195) smooth, elliptic, 1.2-1.8 x 3.7-4.4/x.

Not rare in Chapel Hill, occurring usually in a scattered colony of several in woods
mold in deciduous woods. Dictyophora phalloidea {Phallus indusiatus) is a closely
related tropical species. Fischer considers it the same as the above (1. c, 1890). See
this paper "for a long list of synonyms of D. phalloidea.

Illustrations: Atkinson. 1. c, figs. 6, 7, 11, 13, 16.
Hard. Mushrooms, pi. 55.
Lloyd. Syn. Known Phalloids, fig. 16.

Murrill. Chart of Edible and Poisonous Mushrooms, fig. 34.
Nees von Esenbeck. Syst. Pilze Schw., pi. 35, fig. 258. 1817.
Patterson and Charles. U. S. Dept. Agric. Bull. 175: pi. 35, fig. 1. 1915.
Rau. Bot. Gaz. 8: pi. 4 (a.s Phallus togatus). 1883. See Farlow, ibid., 8: 258.

836a. On lawn of president's house, October 22, 1911.

2286. On a ditch in woods, June 28, 1916.

5195. In rich woods, June 20, 1922.

5343. In deciduous woods near Forest Theater, July 8, 1922. Plant 12 cm. long.

Asheville. Beardslee.

Alabama. Winston Co. Underwood, coll. (N.Y. B. G Herb.).

New York. New York Botanical Garden. Miss Eaton, coll. (N. Y. B. G. Herb.).

Syracuse. Underwood, coll. (N. Y. B. G. Herb.).
Massachusetts. Middlesex Falls. Underwood, coll. (N. Y. B. G. HerD.).
Indiana. Greencastle. Underwood, coll. (N. Y. B. G. Herb.).

PLATE 8

Ithvphallus Rayexelii. No. 649

PLATE 9

Ithyphai.i.us R.wexelii. No. 618
Reduced. Largest plant 7J in. long

HYSTERANGIACEAE

Fruit bodies usually subterranean (in Phallogastcr and Protophallus exposed),
tuberous to pyriform or spherical, the mycelium basal from a central point or with
several scattered fibrils; peridium simple, thin, not intimately connected with the
tramal plates, in some cases disappearing by maturity ; gleba very irregularly chambered,
the walls more or less obviously radiating from a sterile base which extends upward
and outward into plates and strands that do not extend into the peridium; texture
toughish, subgelatinous, after maturity deliquescing into a glutinous mass. Basidia
elongated; spores 2-S, apical, rough or smooth.

Of the few genera of this family, Phallogaster and a species each of Gautieria and
Bysterangium are the only ones heretofore reported east of California. We are adding
Bysterangium clalhroides, a species not before found in the East, and a new species of
Gymnomyces. There is no doubt that many other species of this and related families
await discovery in the eastern United States. On account of its peculiar interest we
are including Protophallus from Jamaica.

The following genera have been described since Fischer's treatment in Pflanzen-
f amilien :

Mortellia Mattirolo. Malpighia 14: 78. 1900. See Saccardo, Syll. 16: 252.

Clathrogaster Petri. Malpighia 14: 125. 1900. See Saccardo, Syll. 16: 250.

Torrendia Bres. Atti dell' R. Acad, di Sc. Lett, ed Arti degli Agiati in Roveredo, 3rd. ser., 8: 132.

1902. See Saccardo, Syll. 17: 241.
Dendrogaslcr Bucholtz. Beitr. Morph. Syst. Hypog., p. 148. 1902. See Saccardo, Syll. 17: 240.
Jaczeuskia Mattirolo. Reale Accad. d. Sci. d. Torino, 2nd. ser. 63: 213. 1912.
Protophallus Murrill. Mycologia 2: 25. 1910. See our treatment.
Callacea Lloyd. Lycoperdaceae of Australia, p. 37: Myc. Notes, p. 1201. See also Cunningham,

Trans. Brit. Myc. Soc. 9: 193, pis. 9 and 10. 1924. The hollow structure of the mature plant

is shown by Cunningham not to exist in the younger condition but to be due to gelatinization and

collapse toward maturity.
Phallobala Cunningham. Trans. Xew Zealand Institute 56: 73, pis. 12, 13. 1926.

Key to the Genera

Peridium ephemeral, entirely or almost absent from the mature fruit body.

Spores melon-shaped, with longitudinal ridges Gautieria (p. 21)

Spores spherical, rough to spiny Gymnomyces (p. 23)

Peridium present at maturity

Fruit body not stalked, buried or only partly exposed Bysterangium (p. 17)

Fruit body entirely exposed

Fruit body stalked Phallogaster (p. 16)

Fruit body not stalked and with the exact appearance of a Phalloid "egg"

Protophallus (Jamaica) (p. 15)

PROTOPHALLUS Murrill

Characters of a true Phalloid except for the entire absence of a stalk, the gleba
remaining permanently enclosed in the volva until its liberation as a slime by irregular
rupture. A monotypic genus known at present only from Jamaica.

15

16 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Protophallus jamaicensis Murrill

Plates 15 and 105

Sessile on very rotten wood; nearly spherical, 2-3.5 cm. thick, whitish, quite smooth
and glabrous, attached by a single, strong basal strand as in Phalloids, and the whole
with exactly the appearance of a Phalloid "egg." Peridium (volva) single, tough and
pliable, membranous, about 0.5 mm. thick, internally lined by a hyaline gelatinous
sheath, 1-2.5 mm. thick, which extends inwards as plates between the deep brown,
fertile tissue which forms rather narrow, elongated, irregular masses, connecting in
center to a distinct, slender, subgelatinous columella, which extends up to about the
center. At maturity the sheath, plates, and columella break down into a homogeneous
slime like white of an egg, in which the unchanged fertile tissue floats. Dehiscence by
rupture of the peridium, probably by bursting from internal pressure.

Spores 1.5-1.8 x 3.5-4.8/x, smooth, short-elliptic. Basidia subcylindrical, 3-3. 7/j
thick at the enlarged end.

This remarkable plant is obviously nearest Prokibera from Brazil, but is distinctly
more Phalloid in appearance, the smooth, even, spherical egg being indistinguishable
from that of a typical Phalloid. The gleba also has almost exactly the appearance of
that in the egg of Dictyophora. We have here in fact a Phalloid in every respect except
the absence of a stalk. It is a beautiful connecting link between Phallogaster,
Jaczewskia, Protubera and the true Phalloids.

Doctor Murrill found the type collection in 1909 (Mycologia 2: 25. 1910) in the
same general locality from which our plants were taken nine years earlier. Although
extra-limital, we include this plant because of its great interest and because no illus-
trations of it have heretofore been published.

Jamaica. Cinchona. 1900. Coker, coll. (U.N. C. Herb.).

PHALLOGASTER Morgan
We take the following from Thaxter (Bot. Gaz. 18: 120. 1893):

"Mycelium fibrous, branching. Peridium spherical to pyriform, stipitate or sub-
stipitate, consisting of a single layer covered by an evanescent cortex and coarsely
reticulated through the presence of numerous irregular thin areas which become per-
forate at maturity, the perforation commonly associated with a general terminal
dehiscence of the peridium into several divergent lobes. Gleba irregularly lobed, the
lobes continuous with slight prominences from the surface of the peridium from which
they are elsewhere separated by a gelatinous layer continuous with a central gelatinous
axis which penetrates the gleba and separates its lobes. The entire contents deliques-
cent at maturity, adhering in distinct masses to the inner surface of the ruptured
peridium."

Phallogaster saccatus Morgan

We can add nothing of importance to Thaxter's description and take the following
from him (Bot. Gaz. 18: 120, pi. 9. 1893):

"Solitary or rarely subcespitose. Peridium spherical to pyriform, 20-50 x 10-25
mm., stipitate or nearly sessile, the surface smooth, slightly uneven, whitish stained with
dull flesh-color at maturity, becoming coarsely clathrate from the formation of irregular
perforations, the perforation usually associated with a terminal dehiscence of the
peridium into from three to five divergent lobes; the dark sage-green gleba adhering in
definite masses of irregular size and shape to the inner face of the peridial wall. Spores

PLATE 10

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Ithyphallus impudicus. Urbana, Illinois

IIYSTERANGIACEAIC 17

greenish, subcylindrical, 4-5.5 x 1.5— 2a/, 6-8 on each basidium." The spores are
smooth; basidia elongated, club-shaped.

r/iallogiister Whitci, described by Peck (Bull. N. Y. St. Mus. 116: 31. 1907)
from Connecticut, was based on small plants which in the opinion of Lloyd are a de-
pauperate form of this species.

Illustrations: FiUpatrick. Ann. Myc. 11: pi. 4, figs. 8 and 9; and text figs. 1, 3, and 7. 1913.
Lloyd. Syn. Known Phalloids, figs. 93 and 94.

Mattirolo. RealcAccad. d. Sci. di Torino, 2nd scr., 63: figs. 9-12. 1912.
Morgan. Journ. Cin. Soc. Nat. Hist. 15: pi. 2, 1892.
Thaxter. As cited above.

Pennsylvania. Ohiopyle. Murrill, coll. (N. Y. B. G. Herb.).

Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.).
New York. West Park. Earle, coll. (N. Y. B. G. Herb.).

Onondago Co. Underwood, coll. (N. Y. B. G. Herb.).
Connecticut. West Goshen. Underwood, coll. (N. Y. B. G. Herb.).
Wisconsin. Blue Mounds. (Univ. Wis. Herb, and U. N. C. Herb.)

Reported also from Massachusetts, Ohio, Kentucky, West Va., Iowa. See Lloyd, Myc. Notes, p. 1327,
and Martin, Iowa Acad. Sci. 32: 222. 1925.

HYSTERANGIUM Vitt.

Fruit body subterranean or partly exposed, subglobose, not stalked; peridium thin
and easily removable at maturity; gleba tough and elastic, finally deliquescing into a
slime; cavities irregular, elongated radially, extending to and abutting on the peridium,
with which the tramal plates are not continuous. Spores elongated, slightly or dis-
tinctly rough, and in all of our species furnished with a basal cup.

Literature
Setchell and Watson. Some Ecological Relations of the Hypogaeous Fungi. Science 63: 313. 1926.

Key to the Species

Plants 3-15 mm. thick, surface fibrous and sandy or trashy; rooting by a bunch of fine fibers; basidia
mostly 2-spored

Gleba distinctly olive; spores pointed at the distal end 77. dathroides (p. 17)

Gleba not greenish; spores rounded at the distal end H. pompholyx (p. 19)

Plants 1-2.5 cm. thick, surface smooth and even; rooting by a single basal cord; spores pointed at the
distal end; basidia mostly 3-spored B. sloloniferum var. americanum (p. 20)

Hysterangium clathroides Vitt.

Plates 16 and 105

Fruiting bodies spherical to subspherical, 3-14 mm. wide by 4-11 mm. high, sessile
or partly buried, rooted by a more or less branched bunch of fibrils, some of which may
pass up on the side of the plant as in Rhizopogon; mycelium fibrous and obvious in the
soil. Peridium surface myceloid with numerous minute particles of earth and trash
adherent, rarely nearly smooth above, at times slightly areolate, whitish when young
with tints of flesh color, becoming buff to clay colored when maturing, consisting of a
single layer 145-190/i thick, composed of much entangled, closely packed threads which
in section appear like closely packed parenchvma cells; easily separable from the gleba.
Gleba tough-cartilaginous when young, becoming somewhat mucilaginous upon del-
iquescing, usually with a more or less central, branched columella which arises from the

UJ LIBRARY 5

18 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

base and from which the tramal plates radiate. Color at first pale greenish, becoming
darker and finally about dark Saccardo's olive (Ridgway). Tramal plates with a
gelatinous hyaline appearance which persists even in mature plants; quite variable in
thickness from one hundred to several hundred microns ; cavities of the gleba labyrinthi-
form, long, and narrow, nearly closed with spores when fully mature.

Spores (of No. 7416) 5-6.2 x 12-15. 5yu, subelliptic, often narrowed toward the
distal end, smooth under low power, but appearing minutely punctate under oil im-
mersion, with a distinct cup at the mucro end (appearing under ordinary power as two
prongs), subsessile on the basidia. Basidia long cylindrical, 5.5-6.6 x 30-42/j, usually
with somewhat sinuous walls, bearing two spores, rarely four.

Odor quite variable: fresh unripe plants with a very faint not unpleasant odor;
deliquescing plants when first collected with a very strong odor suggesting that of
Clathrus; while deliquescing plants kept covered in the laboratory give off an odor like
that of fermenting wine. Taste of fresh plants slightly sweet, not unpleasant.

It seems that this species, from the literature we have at hand, has been reported
only from Europe, North Africa and California. Our plants agree well with the
descriptions of both Vittadini and Tulasne. The former author describes his plant
as varying from about the size of a pea to a hazel nut, rarely larger, with a closely
fitting but separable peridium, from the base of which numerous fibrils extend,
some of which run over the surface of the ground. The odor is described as
being strongly nauseous, much like that of Clathrus. He reports his plants as
being found deeply buried in humus, and occurring in the months of March and
April. With the latter author's description and elaborate notes and figures, our plants
agree in all essential characters. Tulasne described two varieties which differ from each
other in size and in the fact that in one the peridium was much more easily separable
from the gleba than in the other. Our plants vary enough in size to include both of his
varieties, while in our specimens the separability of the peridium seems to be dependent
on the age of the plant, the peridium being separable, but with some difficulty in young
specimens, while it often breaks and slips off in collecting mature plants.

We have examined plants identified by C. Torrend (No. 90) as H. clathroides, and
find that his plants agree with ours in practically all characters, the only difference
being that the spores of his plants are slightly smaller than in ours.

Of the six other species of Hysterangium described or recorded by Tulasne, four:
E. nephriticum, H. Thwaitesii, H. fragile, and H. stoloniferum, while showing certain
points of superficial resemblance, have spores too large for our present plant and show
differences in other characters. Of the other two species noted by Tulasne, H. mem-
branaceum is admittedly very close to H. clathroides, but differs in having smaller spores
according to Hesse's measurements; while B. pompholyx differs in odor and in having
spores which are argillaceous in color. The peridium of H. clathroides is described by
Hesse as being composed of pseudoparenchymatous cells, a structural character with
which our Chapel Hill plant is in agreement, while the peridium of H. pompholyx
according to Hesse is without pseudoparenchymatous cells in the peridium. Of the
three new species described by Hesse, H. rubricatum differs in being larger, in having a
much thicker peridium, and in the clay colored gleba; H. coriaceum differs in having a
much thicker reddish peridium and much smaller spores; H. calcareum differs in having
the peridium covered with a grayish white, silky flocculence and in slightly smaller
spores.

PLATE 11

I

\

DlCTYOPHORA DUPLICATA. Xo. 2286

Much reduced (actual length 18. 5 cm.i

PLATE 12

DlCTVOPHOEA DUPLICVTA. Xo. 5343

HYSTERANGIACEAE 19

The only other record of Hysterangium in the eastern United States is H. sloloni-
fcrum var. amcricanum Fitzpatrick. We have compared material sent us by Dr.
Fitzpatrick with our Chapel Hill //. clathroides and find that the two disagree in size,
peridial structure, and in the much more myceloid basal attachment of the latter plant.

For a detailed comparison see notes under Dr. Fitzpatrick's plant.

Illustrations: Bucholtz. Beitrage zur Morphologie und Systematik der Hypogaeen (Tuberaceen und
Gastromyceten), pi. 1, fig. 16. 1902.
Fries, Th. C. E. Sveriges Gasteromycetcr, fig. 9.
Hesse. Die Hypogaeen Deutschlands 1: pi. 1, figs. 1-14; pi. 7, fig. 19.
Rehsteiner. Bot. Zeit. 50: figs. 7-11. 1892.
Tulasne. Fungi Hypogaei, pi. 2, fig. II-II4.
Vittadini. Monog. Tub., pi. 4, fig. 2.

7416. On very rich, nearly bare soil by Bowlin's Creek, July IS, 1924.
7435. On soil near Meeting of Waters, July 19, 1924.
7490. Under soil and humus by branch, Aug. 9, 1924.
7496. On soil by Fern Walk, Aug. 10, 1924.

7501. Under beech trees, Aug. 14, 1924. Spores elliptic with distal end often pointed, 5.5-6.8 x 14-20m,
with prongs at the proximal end.

Hysterangium pompholyx Tul.

Plate 105

Largest plants up to 1.5 cm. thick, subspherical, rooted by branched fibrils. Per-
idium 180-260/4 thick, not readily separable from the gleba, covered externally with a
white myceloid layer to which particles of sand and trash adhere in abundance, giving
the surface a mottled appearance; in section composed of an inner layer made up of
parenchyma-like cells and the outer myceloid part of loosely woven, clamp-connected
threads, 3-7/i thick, which are heavily encrusted with triangular or rosette-shaped
crystals. Gleba toughish-pliable, pale buff when young, becoming darker and finally
brown upon maturity; quite often with a fairly distinct, hyaline, sterile base which
passes up irregularly toward the center, branching out into the hyaline tramal plates.
Cavities rounded to elongated, often sinuous.

Spores 6.6-7.4 x 13-14.8/u, blunt-elliptic, rounded at the distal end and with a
distinct cup at the proximal end. Wall very remarkable in structure: the outer and
inner layers becoming separated from each other as the spores mature, leaving the
adjacent surfaces of the walls irregularly rough. The outer wall is usually attached to
the distal rim of the cup while the base of the cup rests on the inner wall. Spores buff
to brownish in mass without any tint of green. Basidia 6-7.6 x 16-23/t; 2-spored, club-
shaped.

Our only collection of this plant occurred along with the plant with greenish gleba
which we are calling B. clathroides. The present species seems quite distinct from the
latter in glebal color and structure, and in peridial, basidial and spore structure: the
color of the gleba in the species here described is buff to brown without any tint of green
and the chambers are pouch-like and only slightly elongated, while in our collections
of H. clathroides the gleba always has a greenish tint and the chambers are usually
elongated and narrow; in the present plant the peridium is covered with flocculent
threads which are covered with crystals, while in our specimens of H. clathroides the
flocculence, though usually present, is nothing like so evident as in the present plant.
The basidia in the present plant are two-spored while in H. clathroides they are 2-4-

20 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

spored; the shape and color of the spores furnish the sharpest distinction of all. In the
present plant they are blunt-elliptic, rounded at the distal end, and buff to brown colored,
while in H. clathroides they are subelliptic, distinctly pointed at the distal end, and
always with a greenish tint.

Of the species described by Tulasne, the present plant seems closest to H. potn-
pholyx, especially in the appearance of the gleba in structure and color. Tulasne states
(p. 84) "when one cuts this fungus vertically, a crowd of chambers is found in section,
and their transparence makes it easy to see the argillaceous color of the spores ac-
cumulated on the walls. One could believe, in this case, in the existence of short tubes,
of little fertile sacks, joined together by an agglutinative substance." The spores
agree well enough in size and some of Tulasne's figures agree in shape, being rounded at
the distal end; others however show a distinctly pointed distal end. Tulasne does not
mention or figure any peculiarity in the spore wall, if it did exist, and he figures all of
the spores as smooth. We have found in all specimens examined that the spores under
high magnification present a more or less rough appearance. It is quite possible that
Tulasne in working under rather low magnification overlooked any peculiarity in struc-
ture which might have been present.

Though this slight difference in the spore shape renders our determination somewhat
unsatisfactory, it seems better, with the rather limited first hand information on the
group, not to multiply names.

Illustrations: Tulasne. Fungi Hypogaei, pi. 2, fig. 3; pi. 11, fig. 6.

7490a. Under soil and humus by branch in woods, August 9, 1924.

Hysterangium stoloniferum var. americanum Fitzp.

Plate 105

We adapt the following in condensed form from Fitzpatrick's description (Ann.
Myc. 11 : 130. 1913). In addition the article contains an account of the development
of the fruit body in this plant and in Phallogasler and Ganlieria as well as a discussion
of their relationships and the derivation of the Phalloids.

Fruit bodies subterranean, gregarious, 1-2.5 cm. in diameter, usually globose and
with a basal depression from which a thick, cord-like rhizomorph attaches the plant;
peridium at first snow white, later mottled with dingy brown, not changing color on
exposure, easily separable even in youth, fleshy to cartilaginous, in age tough and
papery and scaling off the gleba; only 0.3-0.4 mm. thick, but composed of two layers, a
thicker, pseudoparenchymatous, large-celled one and a thin fibrous inner one. Gleba
elastic and cartilaginous, olive green in section, the surface bluish just under the pe-
ridium; sterile base and columella distinct, with radiating, anastomosing branches ex-
tending to the peridium; chambers minute, irregular, elongated. At maturity the
gleba softens and the columella and radiating plates become gelatinous, and a not
unpleasant odor resembling gasoline is given off. In alcohol the peridium turns red-
brown.

Spores smooth, narrowly ellipsoidal, tapering at both ends, the base with a short
stalk up to lju long, olive green in mass, 5-6 x 14-17//, at maturity with gelatinous walls.
Basidia long, irregularly cylindrical, 3-spored.

t-l

<

PLATE 14

DrCTVOPHORA DtTLICATA, Egg." No. 2286

HYSTERANGIACEAE 21

The variety differs from the type in the olive green and less firm gleba, thinner
peridium and smaller spores; differs from var. nuitabile Bucholtz in smaller spores,
and white peridium which does not become reddish in age or on handling.*

It seems after a careful comparison of the present plant with Vittadini's, Tulasne's
and Hesse's descriptions of //. clatkroides and with European material of H. clathroides
determined by Torrend and with Tulasne's description of H. stoloniferum that this
plant should stand as a variety of the latter. Fitzpatrick's plant varies from 1-2.5
cm. in diameter, while //. clathroides according to Vittadini, Tulasne, and Hesse varies
from the size of a pea to a hazel-nut, seldom larger. The great majority of the examples
we have seen are about 5-8 mm. thick. The presence of a single, thick, white, cord-
like rhizomorph at the base of each fruit body which is very persistent and in most cases
is unbranched for an inch or more from the fruit body, together with the smoothness
of the surface of the peridium are structural characters of the most fundamental im-
portance with which Vittadini's, Tulasne's, and Hesse's figures and descriptions of H.
clatkroides are in total disagreement. In the microscopic structure of the peridium the
two plants disagree: The peridium of Fitzpatrick's plant is composed of large pseudo-
parenchymatous cells which, cut at any angle or even teased apart, present practically
the same appearance, and there is no outer flocculent layer; while in our study of H.
clathroides we have found that the peridium presents a pseudoparenchymatous cell
structure only when cut at certain angles and when cut at other angles or teased apart
it presents a filamentous structure and there is present in our specimens of H. clathroides
an outer flocculent layer (Tulasne and Hesse report the same condition). In the color
of the peridium and gleba the two plants agree fairly well according to Fitzpatrick's
description, but in a specimen which he sent us the gleba is deep brown with an ex-
tremely faint tint of olive. In basidial and spore characters the two plants are practi-
cally inseparable (Fitzpatrick reports three spores to a basidium but we find a few
with two and some with four) .

It is a noteworthy fact that the peridium of Dr. Fitzpatrick's plant has a striking
resemblance in structure to that of Proto phallus jamaicensis Murrill. It is also obvious
that in the more globular form and single cord-like root, this species approaches nearer
the Phalloids than any other species of Hysterangium.

Illustrations: Fitzpatrick, 1. c, pi. 4, fig. 10; pi. 6; also text figs. 2 and 6.

New York. Ithaca. In humus under leaves in dense woods. Fitzpatrick, coll. (U. N. C. Herb.).

GAUTIERIA Vitt.

Plants subterranean or partly so, subglobose to tuberous; peridium thin and dis-
appearing entirely or in large part by maturity so as to expose the gleba and some of the
cavities; base attached by one or more strands, columella and radiating plates present
or absent; glebal cavities not filled at any stage. Spores shaped like an oblong canta-
loupe with longitudinal ridges and grooves.

Seven species have heretofore been described, two of which are doubtful. Of the
five good species three occur in America and two of these in the eastern states. Gym-
nomyces, in which the peridium is thin and often entirely or partly lacking, is dis-
tinguished by the very different spores.

• Note below that in the plants sent us, preserved in formaldehyde, there is practically no green color, while in all conditions
of 11 . clathroides the gleba is a clear olive.

22 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Literature

Zeller and Dodge. Gautieria in North America. Ann. Mo. Bot. Garden 5: 133, pi. 9. 1918.
See other citations' under the species.

Key to the Species

Odor strong and offensive; cavities small G. graveolens

Odor not offensive; cavities large G. morchelliformis

Gautieria graveolens Vitt.

We copy the following description from Zeller and Dodge:

"Fructifications globose, 1-2 cm. in diameter, light ochraceous-buff to Prout's
brown; stipe slender and fragile, up to 1 cm. long, 1 mm. thick; columella frequently
reaching the center of the fructification, forking; odor very strong, suggestive of decay-
ing onions; peridium thin, composed of delicate, thin- walled, loosely woven hyphae,
soon rupturing and disappearing; gleba ochraceous-tawny to cinnamon-brown; cavities
globose or elongated, minute, empty; septa 40-80/z thick, composed of small hyphae,
compact; cystidia clavate to subfusiform, hyaline, often obscured by the spores; paraph-
yses linear, septate; basidia broadly clavate, 2-spored, 12-16 x 8-9/i, with long filiform
sterigmata; spores ochraceous-tawny, usually with 10 prominent striations, the latter
smooth or nearly so, apex rounded, base pedicellate, 18-19 x 11-12/z, often with a large
oil globule."

"Deeply buried under leaf mold. Europe and North America. Summer."

Illustrations: Bucholtz. Ann. Myc. 1: pi. 5. fig. 14. 1903.
Chatin. La Truffe, pi. 15, fig. 4.
Corda. Icon. Fung. 6: pi. 7, fig. 63.

Fischer in Engler & Prantl. Die Nat. Pflanzenfam. 1. 1**: 304.
Fitzpatrick. Ann. Myc. 11: pi. 4, fig. 11; pi. 7, figs. 29-39, also text figs. 4 and 5.
Hesse. Hypog. Deutschl. 1: pi. 2, figs. 5-9; pi. 7, figs. 4-6.
Vittadini. Monog. Tuberac, pi. 4, fig. 3.

New York. Ithaca. Fitzpatrick, coll.

Gautieria morchelliformis Vitt.

We copy the following description from Zeller and Dodge:

"Fructifications globose to oblong, 1-3 cm. in diameter, with a basal stalk-like
rhizomorph, usually much branched; columella rudimentary, merely a subglobose
summit of the rhizomorph; peridium thin in early stages, quickly evanescent; gleba
ochraceous-tawny to hazel; cavities 1-6 mm. in diameter, subglobose to irregular;
septa white when broken, hyaline to cream-colored under the microscope, composed of a
stupose mat of hyphae, about 75/z broad; basidia about as large as the spores, hyaline,
granular, 2-3-spored; sterigmata filiform, as long as the spores; cystidia in the upper
cavities of the fructification, not prominent; paraphyses clavate, septate, hyaline;
spores fusiform to citriform, ochraceous, longitudinally striate, with 8-10 usually
smooth striations, 1-2-guttulate, pedicellate, 12-24 x 8-12.5//."

"In clay soil. Europe and United States. Spring and summer."

Illustrations: Bucholtz. Ann. Myc. 1: pi. 5, figs. 12 and 13. 1903.
Corda. Icon. Fung. 6: pi., 7, fig. 62.
Bail in Nees v. Esenbeck. Syst. d. Pilze 2: pi. 27, figs. 1-4.

PLATE 15

Protophallus jamaicensis. From Jamaica

PLATE 16

W1

9

vj

Gvmnomyces vesiculosus. No. 7470 [two small plants in upper left corner]
Hvsterancium clathroides. No. 7490 [upper right] X 3; No. 7416 [below]

HYSTERANGIACEAE 23

\'ittadini. Monog. Tuberac, pi. 3, fig. 6.

Klotzch in Dictr. Fl. Boruss. (Fl. Konigr. Prcuss.) 7: pi. 764.

New York. Washington Co., Hudson Falls. Burnham, coll.

GYMNOMYCES Massee & Rodw.

Fruit body subglobose to irregular, peridium delicate or entirely wanting. Gleba
fleshy, rather pale; cavities box-like to labyrinthiform; septa not scissile, composed of
delicate hyphae or bladdery cells. Basidia forming a distinct hymenium, lining the
empty cavities, sterigmata 2—4, Spores hyaline to pallid brown, globose, echinulate
to rugose-reticulate.

Three species have so far been described, one of which is from the United States
(California), and we are adding a fourth from Chapel Hill. The genus is distinguished
from Gaulieria by the rough, spiny, spherical spores; from Arcangeliella, by the absence
of milk; and from Macowanites by the absence of a stalk.

Our Chapel Hill plant described below is clearly marked and is very peculiar in the
friable texture of the tramal plates.

Literature

Zeller and Dodge. Arcangeliella, Gymnomyces, and Macowanites in North America. Ann. Mo.
Bot. Gard. 6: 49, text figs. 1-3. 1919.

Gymnomyces vesiculosus n. sp.

Plates 16, 17 and 105

Fruiting body subspherical, about one cm. thick when fresh, drying to about half
size, the base attached by a few basal fibrils which spring from a narrow depression;
light buff yellow (about Naples yellow) ; odorless. Peridium nearly absent at maturity,
some of the chambers being exposed, others separated from the air only by the tramal
plates and visible as small depressions, the whole giving the surface a spongy look;
gleba pallid gray brown (pale earthy buff), firm, but pliable, not tough or elastic, about
like a potato; chambers rather large and box-like, empty, about 0.4-0.8 mm., walls
thick, about 0.17-0.2 mm. including the hymenial layers which are about 30,u thick;
structure of the walls remarkable, composed of large bladdery cells about 20-40/i thick
except for a very thin layer under the hymenium where the cells are small. Peridium,
where present, composed of a very thin layer of delicate, loosely woven, yellowish
threads. A sterile base and radiating plates are entirely lacking.

Spores color of the gleba under a lens, darker under the microscope, spherical,
7.5-10/u thick including the strong, blunt spines, often stuck together in groups of 4.
Basidia short and thick, the 4 sterigmata nearly half as long as the spore diameter.

We take the name from the peculiar cells of the tramal plates, which are different
from those of any other Gasteromycete we have seen. They remind one of the flesh
of a Russida, and the friable texture of the plant is a reflection of this structure. The
texture is not at all tough or elastic or subgelatinous.

Gaulieria Trabuti from California (N. Y. Bot. Gard. Herb.; Parks, coll., and ap-
parently correctly determined) has vesiculose cells just beneath the hymenium, but the
central region of the trama is composed of filamentous threads as in most species and the
cavities are much more tortuous. The spores are entirely different, having the surface
characteristic of Gaulieria, longitudinally ridged, 9-11 x 11.5-20ju.

7470. Found exposed on soil in frondose woods after a hard rain had washed away the humus, August
4, 1924.

HYMENOGASTRACEAE

Plants produced underground or on the surface, more or less globular or irregular.
Peridium connected intimately with the tramal plates which form the irregular, elon-
gated to more or less box-like chambers which in most cases are empty in youth, with or
without a distinct hymenium; at maturity partly or completely ruled with spores.
Basidia clavate with 1-8, more or less stipitate or sessile spores of a great variety of
size, shape and structure. Capillitium none; sterile base or column in some species
obvious, again practically absent. At maturity not dehiscing but rotting slowly, the
gleba not becoming powdery but (in our species at least) deliquescing inside to a dark
slimy mass or slowly undergoing a viscid disintegration from above downward.

Four of the genera have been reported from the eastern United States and we are
adding Sclerogaster and Melanogaster.

Literature

For special literature on this family see Fischer, Harkness, Hesse, Massee, Tulasne, and Vittadini
as cited on p. 194.

Cavara. Sur la Morphologie et la Biologie d'une Espece Nouvelle d'Hymetwgasler. Rev. Myc. 16:

152, one plate. 1894. Also published in Italian.
Cavara. Nuov. Giorn. bot. ital. 7: 126. 1900.
Fischer. Mykologische Beitrage 25. Jugendstadien des Fruchtkorpers von Leucogaster. Naturf.

Ges. Bern, Mitt. 1921: 301. 1922.
Lloyd. The Hymenogastraceae. Myc. Notes, p. 1138, figs. 2152-2176. 1922.
Zeller and Dodge. Arcangeliella, Gymnomyces, and Macowanites in North America. Ann. Mo. Bot.

Gard. 6: 49, text figs. 1-3. 1919.
Zeller and Dodge. Leucogaster and Leucophlebs in North America. Ann. Mo. Bot. Gard. 11: 389,

pi. 11. 1924.

Key to the Genera

Fruit bodies very small, white; gleba chambers very minute Sclerogaster (p. 25)

Not as above

Spores strongly angled and lobed Nigropogon (p. 37)

Not as above

Fruit bodies veined with superficial fibers; spores elliptic, smooth Rlrizopogon (p. 26)

Fruit bodies attached by scattered fibrils; superficial veins short or absent; cavities box-like,
nearly or completely filled at maturity; spores smooth; basidia irregularly distrib-
uted, not forming a distinct hymenium Melanogaster (p. 38)

Fruit bodies attached by one or a few fibrils, superficial veins present or absent; cavities
mostly polyhedral, when young filled with tissue which later gelatinizes and embeds
the spores; spores pale, usually spherical to oval, rough or pitted but surrounded by a
smooth, hyaline, gelatinous sheath; basidia irregularly distributed, not forming a dis-
tinct hymenium Leucogaster (p. 42)

Fruit bodies without obvious vein-like fibers running over the surface and attached only at
one place

Spores not spherical, in our species surrounded by a bladder Hymenogasler (p. 45)

Spores spherical, reticulated or warted Octaviania (p. 47)

24

PLATE 17

Gymnomyces vesiculosus. No. 7470. X 130
[above]

Hymenocaster foetidus. No. 7473. X SO
[below]

SCLEROGASTER MINOR. No. 7474. X 25

[above]

Hymenocaster foetidus. No. 7473. X 15
[below]

HYMENOGASTRACEAE 25

SCLEROGASTER Hesse

Fruiting bodies white, subglobose, small, rooted by several fibrils; peridium soft,
flocculent, but persistent, often connecting with byssoid mycelium which runs about
conspicuously through the substratum; cavities very minute, round to oblong, becoming
nearly completely filled with spores; hymenium disappearing, being replaced by spores;
septa very inconspicuous at maturity; spores abundant, spherical, warted, yellow.

The genus was established by Hesse to contain the plant called Octaviania compacta
by Tulasne and up to the present no other species has been added. Our plant is the
first record of the genus in America outside of California.

Tulasne, recognizing the striking distinctness of the species 0. compacta, reluctantly
placed it in the genus Octaviania saying, "This species differs much from all the hypog-
enous fungi which we have seen, and it is with doubt that we place it in the genus
Octaviania. In fact it has none of the sterile base, or large, lacunate, sessile chambers
of 0. astcrosperma, differing from it in its compactness and in that its chambers become
nearly entirely filled with spores. It should perhaps rather be placed with Hydnan-
gium, after H. liospermitm or H. hysterangioides; but it differs from these fungi by its
mycelium which is much like that of Hysterangium."

Sclerogaster minor n. sp.

Plates 17, 18 and 106

Fruiting bodes small, about 0.3-1 cm. wide by 0.3-0.7 cm. high, subglobose, often
irregularly lobed ; buried 2—4 cm. under cedar humus, often more or less covered with a
pure white byssoid mycelium which extends through the humus, covering twigs and
cedar berries so that the latter may often be mistaken for the fruiting bodies themselves;
usually with several considerably branched, mycelioid fibrils extending out from the
base or sides of the plants, often connecting two or more plants. Odor faintly
earthy. Peridium pure white on young fresh plants, changing only slightly on drying,
0.3-0.45 mm. thick, composed of two layers: an outer, very thin, byssoid layer made up
of delicate threads which are in places continuous with the mycelium, and the thick
inner layer which in young fresh plants is composed of chains of large bladder-like cells,
but which in mature plants appears to be composed of large parenchymatous cells;
peridium readily separable from the gleba in areas. Gleba pure white when young
and fresh, becoming pale buff and finally deep ochraceous yellow when the spores ripen.
Tramal plates, 60-185^ thick composed of septate, branched threads 3-5^ thick,
continuous with the peridium, continuous also (when it is present) with a sterile plug-
like area which may be basal or lateral, and also with a more or less central columella.
The plates become very inconspicuous as the cavities and space occupied by the hy-
menial layer become filled with spores. Cavities 37-100 x 74-300/u, varying from nearly
spherical to long and narrow, becoming almost or completely filled with spores. Scat-
tered throughout the plant are numerous crystals. Hymenium forming a distinct layer
lining the cavities, obvious only in young plants while the spores are forming, dis-
appearing entirely upon their formation, the space occupied by the hymenium being
filled with spores; composed of basidia and long, slightly acuminate cells; basidia long,
irregularly cylindrical, 4—7.4 x 28-40/j, bearing 1-5 spores, three being the commonest
number, sterigmata quite conspicuous, thick and of equal diameter throughout their
length, part or all of the sterigmata remaining on the spore as a distinct stalk which
varies in length from a very small fraction of the diameter of the spore to about as long
as half the spore's thickness.

Spores 7.4-9.5;u thick, spherical, at first smooth, becoming covered with conspicu-
ous warts upon maturing, often with a pedicel.

26 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

The present plant presents some points in structure which show the difficulty in
separating the families Hysterangiaceae and Hymenogastraceae. In the former the
tramal plates arise from a sterile, more or less conspicuous base and hence the peridium
is readily separable from the gleba, an excellent type of this order being Hysterangium
clathroides. In the latter family the tramal plates arise from the peridium, and
hence the latter is more or less inseparable from the gleba. In the present plant
tramal plates usually arise entirely from the peridium but there is quite often a
fairly distinct sterile base or central columella from which the tramal plates also origi-
nate and in sections, of a few young specimens, small spaces have been observed where
the gleba was not even in contact with the peridium. But since the sterile base and
columella, when present, are almost always continuous in all directions with the perid-
ium, and since it is separable only with difficulty from the gleba, we are considering
our plant as belonging in the Hymenogastraceae. It seems clearly a Sclerogaster , as
shown by the presence of numerous mycelial strands which often extend out into byssoid
mycelial plates, the minute glebal chambers, the hymenium with long, irregularly
cylindrical basidia and the accompanying slightly acuminate, sterile cells (which may
be young basidia), and the spherical, warted spores with short stalks. Our plant
differs from S. compactus (Tul.) Sacc. ( = Octaviania compacta Tul. and S. lanatus
Hesse) in smaller size, larger spores, and fewer spores to the basidium. We have
examined the spores of an authentic specimen of Oclaviania compacta, through the kind-
ness of the Director of Kew Gardens, and find them to be subspherical, warted but not
strongly so, pale olivaceous under the microscope, 4.5-6/* thick. They are much less
rough than is shown in Tulasne's drawings, but possibly immature. Aside from micro-
scopic characters the present plant can readily be distinguished from other hypogenous
fungi, which we have observed, by its very small size, white color, minute chambers,
and by the very obvious byssoid mycelium.

7474. Buried in humus mixed with cedar leaves and twigs, August 5, 1924. An ample collection of
about 100 plants.

RHIZOPOGON Fr.

Fruit body subglobose, growing on the surface of the ground or buried up to several
inches in humus or soil. Peridium tough, its surface more or less netted or lined with
adherent vein-like fibrils which fuse below into one or several threadlike rhizomorphs;
gleba attached to peridium, composed of small crumpled or box-like chambers which
are lined by a distinct hymenium and contain an empty cavity in youth, but which at
maturity may or may not be stuffed with spores. After maturity the gleba deliquesces
slowly into a deep brown slime which escapes by the decay of the peridium or its rupture
by insects. Basidia subclavate to nearly cylindrical, thin-walled, collapsing after the
spores are formed; spores smooth, elliptic, 2-8 on a basidium, the sterigmata short or
none, sometimes represented by a little cup which remains attached to the spore.

Tulasne says that the spores are sessile in Rhizopogon, and in all species we have
studied except R. parasiticus the sterigmata are very short or absent. Zeller and Dodge
speak of and illustrate long sterigmata in several cases, but where we have been able to
check these we find the usual short sterigmata. Their fig. 2 on pi. 3, of R. roseolus,
we are confident is an erroneous interpretation of their material, both in the length
of the sterigmata and the gelatinized basidial walls. The gelatinized cells there shown

m

oo
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^

^^

J3
bo

f

g

5 g

a «

s

« h

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8 z

S S

^S

HYMEXOGASTRACEAE 27

are other cells of the hymenium, and the supposed sterigmata are probably the col-
lapsed basidia lying among them. In all species we have seen the basidia are very
delicate and soon collapse.

A very peculiar species is R. parasiticus, which grows as a parasite on the rootlets
of pine. It is the only one of our subterranean Basidiomycetes which is certainly known
to be a parasite. Rliizopogon graveolens has been reported by Zeller and Dodge from
Alabama (Earle, coll.). Through the kindness of Dr. C. L. Shear we have seen a good
plant of this collection and find that it agrees well with our R. rubescens, No. 1932.

Literature

Lloyd. The Genus Rliizopogon. Myc. Notes, p. 1170, figs. 2312-2322. 1923.
Zeller and Dodge. Ann. Mo. Bot. Garden 5: 1. 1918.

Key to the Species

Parasitic in clusters on the roots of pine; plants small R. parasiticus (p. 27)

Not noticeably parasitic; plants larger

Spores without a cup and not appearing truncate
Peridium composed of a single layer

Cavities only partially filled with spores at maturity

Fibrils few; peridium in section reddish brown to black when treated with 7% KOH
Peridium when cut turning pink or rosy throughout; spores 2-2.7 x 5.5-7.4//

R. rubescens (p. 28)
Peridium when cut turning rosy in inner part only, outer part remaining citron

yellow; spores 3-3.7 x 6.4-8.5/* R. roseolus (p. 32)

Fibrils many; peridium turning distinctly purplish in section when treated with 7%

KOH; plants deep sordid brown to black when dry R. nigrescens (p. 30)

Cavities filled with spores at maturity; fibrils many

Surface woolly; septa 60-90^ thick; odor unpleasant R. luteolus (p. 33)

Surface not woolly; septa 74-111m thick; odor pleasant R. piceus (p. 34)

Peridium double; many of the cavities filled with spores at maturity R. maculalus (p. 35)

Spores with a vague cup which is often extended into a collapsed filament (a part of the basidium),

but spores scarcely truncate, 3.7^1.6 x 8-ll/i R. luteolus (p. 33)

Spores with an open cup on the proximal end and thus appearing truncate; glebal chambers very
small and barely visible without a lens

Spores 3-3.7 x 7.4-8.5/» R- atlanticus (p. 35)

Spores 3-4.4 x 6.8-7.2/i R- truncatus (p. 36)

Rhizopogon parasiticus Coker and Totten

Plates 19 and 20

Fruit body up to 1.5 cm. broad and high, though usually much smaller, the great
majority about 2-5 mm., sometimes almost evenly globose but more often lobed and
convoluted; attached at any point to one or several branching, flocculent, rhizomorphous
threads which run in the humus and connect the fruit bodies on different roots; rarely
some of these branching threads may run along and cohere with the surface of the
fruit body; color of both mycelium and the fruit body varying from a light ochraceous
salmon to a warm buff at all ages until decay sets in. Peridium of mature plants du-
plex, 50-1 30/i thick, the outer layer a spongy mass of loosely woven threads that col-
lapse when the plant is cut or bruised or when decay sets in; the inner layer more closely
woven, lighter in color and intimately connected with the internal hyphae; threads of
the peridium soft and delicate, 2.6-lOA/i thick, in young plants more closely woven.

Gleba when fully formed containing many cavities that are minute, irregular,

28 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

20-200/u broad, hollow and lined with the hymenium. Septa 40-1 15/z thick, delicate
and intimately connected with the peridium, the threads that compose them much
branched, segmented, thin-walled, without clamp connections, 2.6-10.4/^ thick and
having much the appearance of those of the peridium. The hymenium contains no
obvious specialized cystidia, but certain cells among the basidia are of more fusiform
shape and these have not been seen to bear spores.

Spores brown, fusiform, smooth, 3-3.5 x 7.8-10.4ju. Basidia short-clavate, 5 x 17/*,
2-4-spored, with slender sterigmata, which are 2.5-3.5/j. long.

Gregarious and often crowded in large numbers in connected colonies in humus
just under or at the surface, and parasitic on rootlets of pine in damp places.

Illustrations: Totten. Joum. E. M. Sci. Soc. 39: pis. 1-7. 1923.

3990. On roots of Pinus cchinata and Pinus lacda, by branch south of Pritchard's, January 10, 1920.

5383. Same place as No. 3990, July 22, 1922.

6051. Roots of Pinus taeda, edge of swampy place north of cemetery, January 19, 1923.

6057. Same place as No. 3990, January 28, 1923.

Rhizopogon rubescens Tul.

Plates 21, 22 and 106

Plants above ground or partly or completely buried; single plants up to 5.7 cm.
broad by 4 cm. high (one cluster almost fused into a single plant 7 x 6.5 cms. by 4.5 cm.
high), irregularly subglobose, not rarely lobed, surface nearly smooth except for ridges,
channels and depressions, usually somewhat flattened above; not viscid; young under-
ground plants pure white when fresh, turning a distinct rosy pink when bruised; ex-
posed plants pure white at first, then yellowish with a tint of olive above (about citron
yellow, amber yellow or buffy citrine of Ridgway with russet areas where bruised),
remaining whitish below, becoming darker in age and an earthy tan with brown or black
areas on drying. Fibrils very few, usually absent above, innate-appressed on the sides,
becoming free below, and passing into one or several more or less conspicuous rhizo-
morphs; underground plants usually without any fibrils and with more or less con-
spicuous myceloid rhizomorphs which may be attached to the top, sides or base of the
plant. Peridium single, 0.4-0.6 mm. thick, pure white when fresh, turning fleshy pink
or rosy red throughout when cut, not viscid, composed of closely packed threads 4-9/i
thick, and numerous brownish amorphous crystals which do not change color upon the
application of 7% KOH. Gleba pure white when fresh, becoming pinkish when bruised
(often changing when simply cut with a sharp knife), after some hours losing the pink
color and becoming dull brown where cut, drying with a pale olive tinge; texture rather
tough and firm. Cavities irregularly flattened, 2-3 to a mm., not rarely up to 1 mm.
long, labyrinthiform; septa 85-1 00/* thick, composed of threads which become gela-
tinized; becoming very scissile upon maturing.

Spores (of No. 7204) light brownish yellow, oblong-elliptic, smooth, 2-2.7 x
5.5-7.4/i. Basidia more or less obscure, clavate, 4.8-6.8 x 16-35/*; thin-walled, often
extending out beyond the hymenium, collapsing more or less completely after the spores
are formed; bearing 4-8 sessile spores; associated with the basidia in the hymenium are
numerous cells which become slightly gelatinized.

Taste and odor at first practically none; plants drying without deliquescing are
nearly odorless, deliquescing plants becoming fragrant with an odor much like that of
strawberry preserves or scuppernong grapes, or decaying apples, in which condition the
plants are infested with numerous fruit flies.

This species can be distinguished from R. luteolus as represented by a European

PLATE 19

Rhizopogox parasiticus. X 40

HYMEXOGASTRACEAE 29

specimen (Hohnel, Krypt. exs. No. 1607) by the more numerous fibrils, tawny colored
peridium and distinctly larger spores of the latter, which in mass and under the micro-
scope are decidedly more colored.

The present plant can also be distinguished from Rliizopogon nigrescens (see below)
by the peridium of the latter, which is viscid when fresh and which dries a much darker
color, becoming inky black in most specimens, by the slender but numerous fibrils,
and by the pleasant but weak odor of the latter plant. A section of the peridium of the
present plant when put in a 7 per cent solution of KOH changes its color only slightly
if at all, while the peridium of nigrescens becomes a distinct orange lavender when so
treated. For differences between this species and R. roseolus, see that species.

This is not at all rare with us in fall and winter, growing on bare soil or in thin
grass and always near or under pines or cedars; rarely found completely buried to a
depth of several inches. It agrees best with R. rubescens among the species described
by Zeller and Dodge, but has a thinner peridium in the dry state, distinctly smaller spores,
and a strong and pleasant fragrance. In R. rubescens the odor is said by Tulasne to
be weak or almost none. The variety Vittadini has spores as small (3 x 6-8yu) as in our
plants, but is said to have a thicker peridium when dry and not to redden so distinctly
when bruised. Tulasne gives the spores as 3 x 7-9^ and Hesse as 3-4 x 6-9/a. The
simple peridium is in section dull yellow or rosy when cut in the fresh state and sordid
brown or reddish brown, or blackish in areas when dry. We have examined Ravenel's
specimen in his Fung. Car. 1: No. 75, labelled R. rubescens Tul. with R. luteolus Vitt.
and R. albus Schw. given as synonyms (Herb. Dept. Agric, Washington). Our plants
are certainly the same, agreeing in all structural characters. The spores of the Ravenel
plant are elliptic, smooth, 1.8-2 x 4.4-6.2/u. Dr. Dodge has seen some of our plants
and thinks it best to retain them in R. rubescens for the present. The strong odor
and small spores is a discrepancy, and it is obvious that this group is not yet in a satis-
factory state. If the sterigmata are as long as stated by Zeller and Dodge, that would
be a further discrepancy. The plant from Hartsville (R. atlanticus) differs conspicu-
ously in the separable peridium which is white in section when dry, in the much smaller
chambers, larger spores with cupped base, and in not turning rose when rubbed.

We have made over 20 collections of this species around Chapel Hill from 1911 to
date. Excellent lots of material have been collected for three successive years during
the fall and early winter from the same location in a pine grove near Chapel Hill. The
spores are very consistent in size in different collections, e.g., 2.2-2.6 x 5.5-7.4;it in No.
6003, 1.6-2.3 x 5.5-7.5m in No. 58.

Collections number 7450, 7457, 7499, and 7563 differ in certain particulars from the
typical pleasant smelling plants such as 6003 and 7565, though two of the first men-
tioned collections, 7450 and 7563, came from the same location as 6003. The peridium
in No. 7450 has a basic color of about tawny with a tint of vinaceous, in places Rood's
brown or burnt umber, and not rarely with black areas, whereas in No. 6003, etc.,
the peridium has a basic color of sordid buff to clay with brown or blackened areas.
In the odor of the deliquescing plants the two sets of collections differ in that the former
plants (No. 7450, etc.) usually have an odor of decaying Irish potatoes when kept covered
in the laboratory for several hours while the odor of No. 6003 is very pleasant. In the
texture of the dried plants the two sets of collections show some differences. Numbers
7450, etc., are all hard and brittle when dry while many of the plants in No. 6003, etc.,

30 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

have a texture only slightly harder and firmer than a raisin; many plants of No. 6003,
however, are bony hard when dry. The above mentioned plants agree in all characters
except the ones mentioned above, and we are therefore retaining numbers 7450, etc.,
under R. rubescens as a color form.

North Carolina. Chapel Hill. Nos. 58, 1444, 1932, 6003, 6043, 6049, 7199, 7203, 7204, 7212, 7213,
7218, 7223, 7450, 7457, 7499, 7563, 7564, 7565, 7567, 7570, 7573, 7579, 7583.

Saxapahaw. Coker, coll. August 1922. No. 5503. On ground under cedars. Strongly parasi-
tized internally by a mold with large, spherical, spiny, golden yellow spores, 18-22jk
thick. These spores were so abundant as to give their color to the entire gleba, but the
spores of the host had also matured and were like those of our collections. There were
no external signs of a parasite, the surface being to all appearances normal.

Asheville. Beardslee, coll.

South Carolina. Sandy soil in pine and oak woods, just under loose surface leaves, about 7 miles above

Georgetown, Dec. 29, 1926. Coker, coll. (U. N. C. Herb.) . Spores 2.-2.8 x 6-8.5,*.
Alabama. Auburn. Earle, coll. (Path, and Myc. Herb., as R. grazcolens).

Rhizopogon nigrescens n. sp.

Plates 23 and 107

Fruiting body above ground, about 2-6 cm. wide by 1.5-3 cm. high, subglobose,
pure white when young and fresh, becoming mottled with several colors when maturing,
about light cadmium (Ridgway) beneath, shading to russet on top, turning orange
below or reddish brown above when rubbed and later almost black; becoming chestnut
to bay or inky black upon drying; fibrils numerous but slender, quite conspicuous in
fresh plants, innate-appressed above, becoming more conspicuous below, passing into
several rhizomorphous strands, reddish when fresh, becoming concolorous with the
peridium and so inherently appressed above and on the sides on drying as to be quite
inconspicuous, usually remaining free and conspicuous below. Fresh peridium slightly
sticky, about 290-400yu thick, simple, filled with a bright red or orange amorphous
material which gives a strong color to a section and which seems to cause the surface
color when rubbed; color changing slowly to a deep purple upon the application of
7% KOH ; in old specimens in which deliquescence has proceeded far the peridium may
be very thin or even absent in places; threads of peridium septate, only slightly branched
1.%-Aij. thick. Gleba pure white when fresh, turning light clay color, sometimes with a
tint of olive on drying; cavities labyrinthiform, up to 0.5 mm. long, not filled with
spores; septa 80-llOju thick (fresh) but varying much in the same collections and some-
time even in the same plant, only slightly if at all scissile.

Spores 2.4-3 x 6-9/i, smooth, long-elliptic, 2-guttulate, 4—8 on each basidium, and
usually remaining attached even after the basidium has collapsed. Hymenium 22-30^
thick; basidia 5-7.4 x 28— 38p, with thin, non-gelatinized walls; basicna extending out
considerably beyond the hymenial surface, collapsing more or less completely after the
spores are formed. In the hymenium are numerous more or less cylindrical cells which
along with the tramal threads become very much gelatinized as the plant deliquesces.
Sterigmata very short and inconspicuous.

Plant nearly odorless when fresh, deliquescing plants fragrant, the odor faintly
suggesting wine or candy; the quality of the odor the same but diminishing in strength
on drying.

In fresh condition the distinctive characteristics of the plants are the peridial colors
and the abundant slender fibrils. In the dried condition the peridium becomes very

PLATE 20

PLATE 20

Fig. 1. Fungal threads attacking cortical cells of the pine. The threads force their way between and
over the pine cells which are the unshaded areas in the figure. X 690.

Figs. 2 and 3. Basidia with nearly mature spores.

Fig. 4. Young basidia. X 1290.

Fig. 5. Mature spores. X 1290.

Fig. 6. Tramal thread, showing nuclei. X 1290.

Fig. 7. Trama and hymenium of mature plant, some of the basidia with two, others with four sterigmata.
X 1290.

Figs. 4-6 stained to show nuclei.

PLATE 20

Rhizopogon parasiticus

1IYMI \OGASTRACEAE 31

dark, sordid brown to a dull or shiny black and the fibrils, so conspicuous in the fresh
condition, usually become so innately appressed above as to appear as though there
were none, while below and occasionally on the sides they remain free and conspicuous.
The pleasant wine-like odor is also distinctive.

The present species seems close to R. roseolas and we were at first inclined to refer
our plant to that species, but upon comparing a specimen of the latter plant kindly sent
us by Dr. Fitzpatrick (from Dept. PL Path. Herb., N. Y. State Coll. Agric. at Cornell
Univ.; Whetzel, No. 598) and determined as R. roseolus by Zeller and Dodge with
our collections of the present plant, they were found to differ in the following ways:
In color the peridium of R. roseolus is citron yellow to olive brown with blackish areas
when dry and the fibrils are very few, innate-appressed and black when dry; the per-
idium of the present plant is chestnut to bay or inky black upon drying, and the fibrils
are very numerous and conspicuous. The peridium of the present plant is viscid when
fresh while in Chapel Hill collections of R. roseolus the peridium is not at all viscid.
In section the peridium of R. roseolus is more or less reddish brown and does not change
color upon the application of 7% KOH, whereas the peridium of the present plant con-
tains a red or orange amorphous material which changes to a purplish orange upon the
application of 7^ KOH. In the macro- and microscopic characters of the gleba the
two plants agree except for the somewhat larger spores of R. roseolus.

In some respects the present plant resembles R. provincialis but the hairlike threads
making up the peridium of the latter plant and the much thinner scissile septa seem to
separate the former from the latter plant.

The species differs from R. rubesccns in having a very slight though pleasant odor,
while the latter is very fragrant while deliquescing and retains the odor for several
months after drying. The almost total absence of fibrils in R. rubescens also serves to dis-
tinguish that plant from the present species, in which the fibrils are quite numerous and
conspicuous in the fresh state and are usually quite obvious in the dried condition.
The present plant turns black with rather inconspicuous reddish black areas on drying,
while R. rubescens becomes light to dark earthy tan (or dull black on crushed places)
on drying. In section the two plants are quite distinct: (1) the peridium in R. rubescens
having brownish, amorphous, crystalline material which does not change color on the
application of 7% KOH, the present plant having embedded in its peridium a bright
red or orange amorphous material which changes slowly to a purplish orange on the
application of 7% KOH; (2) in R. rubescens the septa are slightly scissile in the fresh
and very much scissile in dried plants, while in the present plant the septa are usually
not at all scissile in either the fresh or dried plants.

From R. piccus which is also blackish when dry R. nigrescens easily differs in the
smaller, more densely packed chambers, the absence of purple color in the peridium and
in the more regular spores.

Attention has already been called to the peculiarities of the basidia in the present
plant, a condition found in young fresh specimens of Nos. 7372 and 7569. In older
specimens such as Xo. 1910 no plump basidia could be found at all, whereas conditions
such as those represented in fig. 7 (showing collapsed basidium) are fairly common, but
could be made out only after crushing a section of the gleba under the cover glass. It was
only after studying young fresh material which showed the different stages in the
development of the basidia that we were able to account for the extreme difficulty in

32 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

finding basidia in the old dried specimens. It seems probable that, since Zeller and
Dodge were studying dried specimens on the hymenia of which the spores are usually
so abundant as to render such delicate objects as sterigmata, even when present,
extremely hard to see, the long prongs figured and described by them (especially in
R. roseolus) as sterigmata are nothing more than the remains of the collapsed
basidia (fig. f).

591. On bank of a brook, Oct. 18, 1912. Spores 2.2-2.8 x 6-7.5/*.
1011. Under cedars, Oct. 21, 1913.

7177. On rocky ground in pine woods, Sept. 30, 1923. Spores 2.5-3.2 x 6.5-9**.
Also Nos. 976, 1031, 1910, 7198, 7210, 7227, 7372, 7478, 7566, 7569, 7582, 8139, 8168.

South Carolina. About 7 miles above Georgetown, Dec. 29, 1926. Coker, coll. (U. N. C. Herb.).

Spores 1.8-2.6 x 5-7.6/t.
Alabama. Spring Hill. Nov. 9, 1902. (Univ. Wis. Herb., as R.rubescens, and U. N. C. Herb.)

Rhizopogon roseolus (Corda) Hollos

Plates 24 and 107

Plant irregularly subglobose, usually flattened above, not rarely flattened laterally
to about kidney-shaped, 0.5-2.5 cm. thick, nearly white when young and fresh, turning
reddish or purplish red (nearly Hessian brown of Ridgway) where bruised, becoming
citron yellow to olive brown with blackish areas when dry; fibrils inconspicuous, absent
above and appearing as innate ridges below, blackening upon drying, passing into one
or more conspicuous and considerably branched rhizomorphs. Peridium when fresh
0.4-0.6 mm., not rarely up to 1 mm. thick; outer part citron yellow, glabrous above,
thinly felted below, composed of loosely packed, only slightly entangled hyphae, 4.4-1 1/*
thick when fresh, septate, without clamp connections, encrusted with pale olive-colored
granules; inner part rosy salmon in section when fresh, composed of more closely packed
and much entangled hyphae with reddish granules intermingled; when dry more or less
reddish brown throughout and not changing color in 7% KOH; 200-300/* thick. Gleba
pallid white at first, drying warm buff, not changing when cut, rather tender when fresh,
becoming quite brittle upon maturing; tasteless, odor faintly farinaceous; cavities
averaging 2-A to the millimeter, labyrinthiform, not rarely up to 2 mm. long in section,
empty; septa 90-110/*, averaging about 100/* thick, occasionally scissile, hyphae loosely
packed, much entangled and branched. When dry the peridium is only about 200-300/*
thick, the outer part 30-80/*, the inner 110-230/* thick.

Spores (of No. 7207) elliptic, smooth, with two conspicuous droplets, 3-3.7 x
6.4-8.5/*. Basidia 6-9 x 18-30/*, with thin, nongelatinized walls; extending out con-
siderably beyond the hymenial surface, collapsing more or less completely after the
spores are formed. On the hymenium are numerous more or less cylindrical cells
which along with the tramal threads become very much gelatinized as the plant
deliquesces.

We have compared our plants with a specimen of R. roseolus studied by Zeller
and Dodge (Whetzel, No. 598; N. Y. State Coll. of Agric. at Cornell Univ.) , kindly sent
us by Dr. Fitzpatrick and find that they agree in all respects except for the size of the
spores which in the plant from Ithaca are somewhat larger (measuring 3.6-3.9 x
8.1-11.1/*) than those of the Chapel Hill plants.

This plant is closest to R. rubescens and epigeal forms of the latter might easily be
confused with the present plant since the exposed surface of R. rubescens is quite often
more or less citron yellow. The citron yellow color when present, however, in the

PLATE 21

Rhizopocon rubescens. No. 7199 [two top plants and one on
left just below]; No. 7213 [all others].

PLATE 22

%

f.'H

Rhizopogon rubescens. No. 7565

HYMENOGASTRACEAE 33

latter, usually extends throughout the thickness of the peridium; while in the present
plant the yellow color is limited to the outer part, the inner part being a distinct rosy
salmon. The two plants are quite distinct in the color changes in the gleba, the present
plant showing no change in the gleba upon cutting, while in R. rubescens the cut surface
of the gleba changes to a distinct, rosy pink. Furthermore, the cavities of the gleba
in R. roseolus are larger than in R. rubescens, and the septa are considerably thicker,
more gelatinized and much less scissile.

7207. On very damp, moss-covered earth, mixed wood, November 11, 1923.

7212a. On mossy ground in old road, pine woods, near No. 7212, November 13, 1923.

7214. On damp grass and moss covered earth, open place in thin mixed woods, November 17, 1923.

7229. On ground, partly under pine needles, near New Hope Creek, December 1 1 , 1923.

New York. Ithaca. On ground in moss. Whetzel, coll. (N. Y. St. Coll. Agric. Herb., No. 598, and
U. N. C. Herb.).

Rhizopogon luteolus Fries & Nordholm emend Tulasne

Plate 106

We have not found this species in North Carolina but have studied a piece of a
plant preserved in formalin kindly sent us by Dr. Fitzpatrick from Ithaca, N. Y.
This same collection was examined by Zeller and Dodge and included under this species.
These authors give the size of the spores as 3-5 x 7-16yu, while our measurements are
3.7-4.6 x 8-13/li. They give the basidia as 9—10 x 12-13/*, with sterigmata as long
as the spores, while we find the basidia about 5.5/* thick in the uncollapsed half
by 24-28/1 long, and the spores nearly sessile, as is the case in most species we have
seen. In this material the thin-walled basidia had collapsed in the distal half or
throughout, but often were easily recognizable from the 2, 3 or 4 spores attached to
the end. We find the great majority of the cavities to be completely filled with spores.

We have examined also a European specimen distributed by von Hohnel as R.
luteolus (Krypt. exs. No. 1607; U. S. Nat'l. Herb.), and find the surface to be distinctly
netted below, the peridium thin; chambers stuffed with spores; no olivaceous color
obvious in the gleba. Basidia 5.5-9 x 20-30/j, with 4-8 subsessile spores which are
3.2-4.4 x 5.5-S/i, rarely up to 11/u, mostly elliptic but some bent or angular. As usual
in RJiizopogon, the basidia are thin-walled and collapse after maturing the spores, and
the hymenium is also furnished with numerous elongated cells with highly gelatinized
walls. This plant differs from R. luteolus from Dr. Fitzpatrick in the size and numbers
of spores on the basidia and in the presence of gelatinized cells in the hymenium of the
European plant, but because of other resemblances and the scanty material we are
including both under this species.

Krombholz's figure (Abbild. Schwamme, pi. 60, fig. 15) shows a green gleba, and
Hesse notes the gleba as becoming olivaceous. Tulasne rejects Krombholz's figure
as not being his interpretation of R. luteolus.

The following description except for the italicized parts, which are ours, is from
Zeller and Dodge (Ann. Mo. Bot. Gard. 5:11. 1918) :

Fructifications subglobose to oblong and often pear-shaped, diameter up to 3 cm.
when dry, color warm buff to mummy brown when dry: odor weak at first and later

34 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

stercoreus (Tulasne) ; fibrils numerous but not prominent, fine, elastic, about the same
color as the peridium or darker, composed of septate hyphae, innate-appressed above
and not very prominent below; peridium thick, 240-400^, simplex, context meshy and
quite loose, stupose, ochraceous-buff to ochraceous- tawny under the microscope; gleba
white at first, then yellowish when dry; cavities narrowly labyrinthiform, empty or
filled with spores where the cavities are small; septa 60-90p. thick, made up of hyaline,
branched hyphae mostly extending parallel with the surface of the hymenium, becoming
scissile early; basidia almost entirely collapsed, a few were seen which were completely
collapsed only in the distal half, the uncollapsed part about 5.5/u thick; the length of the basidia
24-28fi. Spores acrogenous, distinctly colored, ochraceous tawny, usually ellipsoidal,
but often irregularly bent or angular, 3.7-4.6 x 8-lly. rarely up to 132y.s smooth, with a
vague cup which is often extended into a collapsed filament (a part of the basidium) but
spores scarcely truncate.

In sandy coniferous woods. Cosmopolitan.

Illustrations: Hesse. Die Hypogaeen Deutsch., pi. 2, figs. 1-4; pi. 5, figs. 5-7; pi. 7, fig. 26; pi. 9, fig. 26.
Tulasne. Fungi Hypogaei, pi. 1, fig. 5; pi. 11, fig. 5.
Zeller and Dodge. Ann. Mo. Bot. Gard. 5: pi. 1, fig. 4.

New York. Ithaca. Reddick, coll. (Dept. PI. Path. Herb., 7359, in N. Y. State Coll. Agric. at Cornell

Univ. and U. N. C. Herb.).
Reported by Zeller and Dodge also from Massachusetts and North Carolina.

Rhizopogon piceus B. & C.

Plate 107

Plants irregularly globose, up to 1.5 cm. thick, glabrous, greenish yellow when
first dug, soon sordid yellowish and then brown to blackish on exposure and handling;
fibrils abundant and netting the surfaces all over, inherent and only sparingly free
below, dark brown, several entering the ground and there branching and holding a ball
of earth. Peridium 300-500/u thick, single, mottled with dark and light areas, in
microscopic section composed of densely packed, small threads about 3.7/j thick, and
of dark colored, subspherical to oblong or irregular bodies up to about 10/i long which
turn reddish brown in large part when fresh upon the application of 7% KOH. When
dry a section in KOH becomes largely black, mottled with pale areas. Gleba brownish
yellow, toughish and elastic, the chambers small, 70-300/i broad or long (most about
100-200/u), irregular, but less so than in other species, remarkable in being blocked out
into small groups by obvious plates; empty at first, becoming densely filled with spores
at maturity, lined with a hymenium; septa not scissile or very slightly so, 74— 111/z
thick, the context composed of densely packed, much-entangled hyphae about 2-3. 7/i
thick. Odor distinct but not nearly as strong as in R. rubescens (No. 6049, etc.),
pleasant and suggesting walnuts or wine.

Spores brown in mass, pale olivaceous when single, smooth, subelhptic with two
oil drops, very peculiar in that a good many are slightly to considerably bent and less
often deviating to irregular and even subtriangular, occasionally slightly constricted,
2.9-3.7 x 7.2-8/x, not rarely up to 11/*. Basidia totally collapsed in dried specimens;
in the hymenium are numerous highly gelatinized thick-walled cells.

According to Dr. Dodge our plants are just like the co-type in the Curtis Herbarium
(Hong Kong), including the abundant and conspicuous bodies in the peridium. A
note with the Curtis plants says, "Germinating beneath the surface, and by its expan-
sion causing small areas of a hand-breadth size to cleave off. Black or brown after
exposure."

IM.ATE 23

Rhizopocon nigrescf.ns. No. 7210 [above] ; No. 591 [below]

HYMEXOGASTRACEAE 35

From R. lutcolus which is somewhat similar in color of fresh plant, abundant
fibrils and irregular spores, this species differs in the abundant dark bodies in the
peridium. giving to the dried plants a blackish color, by the smaller and more densely
packed spore chambers and the plates which divide the gleba into larger areas, each
containing several spore chambers. This last peculiarity is the rule in Melanogasler
and indicates an affinity to that genus. For a comparison with R. nigresccns see that
species.

6059. Under pines in upland wood? near a branch, rooted in sandy soil under several inches of rotting
1 umus, Jan. 30, 1923.

Rhizopogon maculatus Z. & D.

Plate 107

Plants subglobose, ovoid, oblong, or irregularly lobed, flattened, 0.6-1 cm. high by
1-2 cm. broad, becoming much smaller upon drying; sordid yellow to deep brown and
mottled with lighter colored areas; attached to the earth by one to several conspicuous,
dark brown fibers which are free from the peridium below, anastomosing to form a loose
web which is innate-appressed laterally and which branches into little fibrils that are
sparse and inconspicuous above. Peridium 70-14% thick, composed of two layers:
the outer one 30-80/j thick, made up of hyaline, densely entangled hyphae; the inner
layer 35-60/z thick, consisting of brown, much entangled hyphae and brown bodies.
The outer peridial layer may become worn off or mashed down so as to expose the inner
dark layer. Gleba dark grayish to brown, tough, elastic when fresh, becoming hard
upon drying; cavities of gleba labyrinthiform, appearing long (up to 1 mm.) and narrow
in section, many of them filled with spores at maturity; tramal plates 45-80/x thick,
composed of the two hymenial layers which are about 20^ thick and a non-scissile layer
of hyphae which are about 3.7/* thick.

Spores smooth, elliptic, regular, rounded at both ends, 3-3.7 x 5.5-7.4/*. Basidia
clavate without gelatinized walls, 4-5 x 10-14/*, with 4-8, usually 8, nearly sessile
spores.

Of the species of Rhizopogon studied by us this plant is nearest R.piceus, but differs
from that in the more loosely attached and more conspicuous fibers, in the shorter,
elliptic spores which are not angular, in the much smaller basidia without gelatinized
walls, and in the double-layered and much thinner peridium. It may also be separated
from R. atlanticus, with which it agrees in having a double peridium, by the much more
numerous and conspicuous fibers, by the absence of the peculiar cups on the end of the
spores, and by the thinner peridium. Our description has been drawn from material
preserved in formalin. For a comparison of R. maculatus with others nearly related,
see Zeller and Dodge, 1. c, page 5.

South Carolina. Hartsville. Coker, No. 955. On the surface of damp soil near a branch in a
pasture, November 27, 1913.

Rhizopogon atlanticus Coker and Dodge n. sp.

Plates 25 and 106

Plants subglobose, oblong or irregularly lobed, 1-4 cm. thick; young plants pure
white then dull yellow toward maturity, not turning rose when rubbed, becoming
pinkish cinnamon or sordid yellow-brown upon drying; odor of fresh plants faintly of
woods mold, odor of deliquescing plants fragrant like Calycanthus blossoms; superficial

Lui LIBRARY )%

36 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

veins few, not conspicuous, innate appressed, darker colored than the peridium usually
attached at base by a single small strand which is not conspicuous, separating very
easily from the substratum. Peridium when fresh 300-450/* thick, when dried and
resoaked 250/* thick, simple, composed of loosely interwoven, much entangled, en-
crusted hyphae, pale under the microscope, if dried in the fresh state, except for an outer,
thin, brownish crusty area; easily separable from the gleba which is covered with a con-
tinuous white layer of the same tissue which separates the chambers. Gleba white
when young, becoming Isabella color to buffy brown, shrinking considerably upon
drying and not rarely separating from the outer peri dial layer; cavities very small,
almost invisible to the naked eye, about 5-8 to a mm., slightly larger toward the center,
narrowly labyrinthiform, hollow at first, then only partly filled with the brown spores;
tramal plates 45-95/* thick, hyaline, composed of a narrow central region made up of a
few gelatinized hyphae (3-3.9/* thick) and the two hymenial layers.

Spores 3-3.7 x 7.4-8. 5**, blunt-elliptic, brown in mass under the low power of the
microscope, furnished at the base with a large empty cup, formed by a continuation of
the spore wall. Basidia inconspicuous, 5.5-7.8/* thick, collapsing early, bearing 4-8
sessile spores on the ends, the lower part of the spore becoming the empty chamber.

This species is near R. truncatus described by Linder in Rhodora 26: 196, 1924,
from a single specimen collected by him under hemlocks in New Hampshire. Mr.
Linder has kindly furnished us with a section of his plant. The peridium of R. truncatus
is described as being 120/* thick and composed of a single layer, while the peridium of the
present plant is 300-450/* thick when fresh and up to 250/* thick when dried and re-
soaked, and is composed of two layers. The tramal plates of the present plant are much
thicker than those of R. truncatus, but have the same arrangement and general appear-
ance. The cavities between the plates are exceptionally small in both plants. The
basidia are alike except for a slight difference in thickness. The spores of the two
plants are remarkable for the genus in being the only spores with apparently truncated
ends. If the spores of R. truncatus are examined under an oil immersion objective they
can easily be seen to be provided with the same empty chamber formed by a continua-
tion of the spore wall as in the spores of the present plant, a point not mentioned by
Linder. The cup-like extensions are not so conspicuous as in the southern species
where they are easily visible under a high dry objective. The spores of our plant are
also longer, narrower and paler than those of R. truncatus. This cup on the spore
giving it a broadly truncate base distinguishes these two species from all other Rhizo-
pogons. A similar cup is found in Hysterangium and Melanogaster.

South Carolina. Hartsville. Coker, No. 5999 (type). In damp pine woods just below the surface of

humus, or a few with upper part exposed, December 25, 1922. (U. N. C. Herb.)
Florida. Eustis. At foot of a long leaf pine. Thaxter, coll. 1897.

Rhizopogon truncatus Linder

Plates 26 and 106

This species has recently been described from New Hampshire (Camp Algonquin
in Holderness; moist birch woods with scattered hemlocks) and a slide of the type has
been kindly sent us by Mr. Linder. The species is closely related to our R. atlanticus,
which see for a comparison.

We translate the following from the original Latin description (Rhodora 26: 196,
pi. 148, figs. 3, 5-7. 1924).

NOIIOI 'INAVM303

=

o

r
-

o

r

>

H
M
to

HYMEXOGASTRACEAE 37

"Fructifications subglobose and irregular, measuring 3 cm. in diameter, a beautiful
citrine when fresh, odor not strong, when dry like Phallus dupHcatus, chamois or honey-
yellow (Ridgway); funiculi attached, mummy brown (Ridgway) when dry; peridium
thin. 120m thick, simple, byssoid, context of slender hyphae, maize yellow (Ridgway);
gleba when dry cinnamon buff (Ridgway); cavities globose or irregular, empty; septa
In 27 ft thick, context of hyaline hyphae; basidia broadly clavate, 7 x 13— 15ju, hyaline,
with sterigmata which are 3-4.5/u long; spores truncate, broadly ellipsoid, creamy,
3-4.4 x 6.8-7. 2m, smooth."

In a letter Mr. Linder calls attention to the fact that the cavities grow larger toward
the center.

Illustrations: Linder. As cited above.

Xew Hampshire. Holderness. Linder, coll. (Farlow Herb.).

NIGROPOGON Coker and Couch, n. gen.

As in PJiizopogon except adherent superficial fibers entirely lacking, peridium and
tramal plates composed almost entirely of pseudoparenchymatous cells, and the spores
strongly angular.

Nigropogon asterosporus n. sp.

Plate 108

Fruit body hypogeal, subglobose, 11 mm. thick when fresh, shrivelling and drying
to a little more than half its original size ; peridium when fresh nearly pure white, drying
tawny or amber brown; rooting by an entangled network of very delicate black, hairlike
fibrils; adherent, vein-like fibrils entirely lacking on the surface. Peridium 100-220ju
thick, of one layer composed of large parenchymatous cells and delicate threads.
Gleba pure white at first but soon becoming pale vinaceous russet or pecan brown
(Ridgw.) as the spores are formed, drying sayal brown. Cavities up to 1.5 mm. wide
by 120/i deep, sinuous, remaining empty. Tramal plates quite variable in thickness,
80-200m thick, usually about 1 10m thick, composed of thin-walled pseudoparenchymatous
cells and threads, continuous with the peridium. Hymenium composed of short, more
or less rectangular cells which elongate to form basidia. Basidia collapsing in the
distal half immediately after the spores are formed, the proximal half persisting;
7.5-°m thick in the proximal half, 5.5-6.5m in the distal half by 26-38m long; usually
with 4 (rarely 2) nearly sessile spores. Odor when fresh slightly like an old Irish potato
but not unpleasant ; on drying becoming like that of ham, faintly nitrous or phosphorous.

Spores color of gleba, very peculiar in shape, strongly angular, usually with four
angles showing in optical section, the most acute angle being at the mucro end, 8.3-
9.8x11.5-12/*.

This remarkable species belongs in the Hymenogastraceae, close to PJiizopogon.
It is clearly distinct from that genus, however, in peridial and tramal plate structure and
in the shape of the spores. In the present species the peridium and tramal plates are
composed of pseudoparenchymatous cells and a few threads, while in PJiizopogon the
peridium and tramal plates are composed of threads which usually are rather delicate;
also the spores of this plant are angular while the spores of PJiizopogon are invariably
elliptic and smooth. The rhizomorphs, the empty glebal chambers, and the structure
and development of the basidia suggest a close relationship to PJiizopogon.

The development of the basidia in this species is very peculiar. The hyrreniumis

38 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

composed of cells which are almost as thick as they are long. These cells elongate at
the distal end to form a narrower tube and on the end of this tube the spores are borne.
Since there is no cross wall between the cell and the tube and since both empty their
entire contents into the spores, both together must be considered the basidium. After
the spores are formed the tube collapses. A somewhat similar condition has been found
and is described here for certain species of Rhizopogon and Odaviana purpurea.

8271. Under deep layer of decaying oak and dogwood leaves, slightly buried in the soil, Oct. 22, 1927.

MELANOGASTER Corda

Fruiting bodies rounded or irregularly lobed with branched fibrils springing from
any part of their surface. Peridium tough, fleshy, not readily separable from the
gleba. Gleba with rounded to polygonal chambers which are larger towards the
center and become nearly or completely filled with spores. Tramal plates directly
continuous with the peridium, bearing the numerous basidia which are not arranged
in a definite hymenial layer but are irregularly distributed throughout a broad periph-
eral zone which may nearly fill the cavity. Basidia club-shaped, 2-8-spored; spores
usually apical on the basidia but not rarely somewhat laterally placed, elliptic, smooth,
furnished with a cup at the proximal end. At maturity the gleba breaks down into a
slimy mass and escapes by the final rupture of the peridium.

The irregularly distributed basidia and the nearly solid interior at maturity gives
this genus a superficial approach towards Scleroderma. The essential structure is,
however, quite different as the chambers are not really stuffed solidly at all stages and
the basidia are not surrounded by a ground tissue which nourishes and matures the
spores. The distribution of the spores in a slime and not a powder is also a fundamental
difference. Rhizopogon seems the nearest relative.

We are including four species, two from New York State and two from North

Carolina.

Key to the Species

Spores less than 11m long and the distal end not pointed

Peridium 300-600^ thick, composed of irregularly swollen hyphae 7.4-33m thick

Odor strong and nauseous: basidia long, club-shaped M. nauseosus (p. 39)

Odor not strong; basidia short and thick M. rubescens (?) (p. 40)

Peridium 50-320m thick, composed of threads 3-7m thick M. variegatus (p. 38)

Spores over 11m long and the distal end pointed M. ambiguus (p. 41)

Melanogaster variegatus Vitt.
M. mollis Lloyd

Plate 109

Part of a collection of this plant preserved in formalin was sent us by Dr. Fitz-
patrick from Cornell University. From this collection the following descriptive notes
have been made:

Fruiting bodies up to 3.7 cm. in diameter, irregular in form; attached at base by
elaborately branched rhizomorphs. Peridium simple, composed of closely packed,
much entangled, clamp-connected threads, 3-7/* thick; surface of peridium covered with
numerous, thick-walled, swollen, club-shaped cells; peridium deep brown in section,
quite variable in thickness, 50-300/i thick. Tramal plates directly continuous with

ts

Pi

HYMENOGASTRACEAE 39

the peridium, concolorous with it in places but usually appearing white (?) to the naked
eye or hyaline under the microscope; 74-148/* thick. Threads of the tramal plates
2-6m thick, separate, considerably branched and much entangled, highly gelatinizable.
Glebal chambers quite variable in size, 0.3-2 mm. long, elongated or polygonal basidia
irregularly disposed.

Spores black in mass, dark brown under microscope, 3.5^1 x 7-10. 4/u, elliptic,
smooth, with a distinct hyaline cup (the remains of the short stalk which attached the
spore to the basidium) at the proximal end. Basidia irregularly arranged, 4.2-5 x
12-18p, club-shaped, usually with four spores, rarely with three or only two.

The spores of the present plant have been compared with type material kindly sent
us by Dr. Dodge from the Sprague Herbarium at Cambridge. The spore sizes of the
type vary from 3.7-4.5 x 7-9/j, rarely 10,u long, and have rather indistinct prongs (rings)
at the proximal end. The spores of the present plant, while agreeing with the type in
size, differ from the latter in having the proximal end extended into a rather long and
fairly conspicuous cup-like stalk. These cup-like extensions while present on the proxi-
mal end of the type spores are much less conspicuous than in the present plant. The
threads composing the tramal plates and peridium of the type material are so collapsed
that it is very difficult to make out their structure. The present plant agrees with
material kindly given us by the New York Botanical Garden which was collected and
identified by Dr. Hollos in Hungary (Krypt. Exs. No. 1812) except that the spores of
the latter plant are slightly thicker and without such a distinct stalk. The slight
differences in spore size and structure do not seem sufficient to exclude the present plant
from M. variegatus.

The present species is distinct from the Blowing Rock plant No. 5804, described
below, in size, in peridial structure and in spore color and size.

Illustrations: Bucholtz. Beitr. Morph. und Syst. der Hypogaeen (Tuberaceen und Gastromyceten),
pi. 1, fig. 19.
Fries, Th. C. E. Sveriges Gasteromyceter, fig. 36.

N'ew York. McLean, near Ithaca. J. H. Miller, coll. August 17, 1924. (Cornell Univ. Herb., No.

12657.) Two specimens collected together and attached at base by rhizomorphs.
Canada. Deamess, coll. Reported by Lloyd, Myc. Notes, p. 1064.

Melanogaster nauseosus n. sp.

Plates 27 and 108

Fruiting bodies gregarious, subglobose; up to 3 cm. wide by 2 cm. high, most
about 2 cm. wide by 1 .5 cm. high. Peridium light cinnamon brown when fresh, becoming
darker, about russet; outer part turning reddish upon being cut or bruised or eaten by
insects. Fibrils rather large below, few and very inconspicuous above, darker than the
peridium. Peridium, 300-600/j thick, single; composed of large, subglobose to elon-
gated vesicular cells usually about 20/i thick, often up to 33 x46/i; on the outer side are
occasional patches of septate irregular threads with clamp connections. Gleba at
first white, then becoming light tan and auburn as the spores mature and finally black
in the dried plants, except for the yellowish tramal plates; more or less firm and solid
when fresh and young, but changing upon maturing into a slimy gelatinized mass.
Glebal chambers 1-3 mm. wide, polygonally shaped, loosely filled from the first with
hyaline hyphae. In these are developed one to several "pockets" of spores. Chambers
without a distinct hymenial layer, basidia irregularly disposed. Basidia elongated,
club-shaped, 4.6-5.8/t thick, with 4-8, usually 6 or 7, nearly sessile spores. Basidia

40 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

gelatinizing and early disappearing. Odor very strong, penetrating and peculiar,
quite unpleasant, nauseous, but with a somewhat fruity element like decaying apples
and very attractive to fruit flies, even in the dried condition.

Spores smooth, elliptic, 3.9-4.2 x 7.4-8.4ju, with a hyaline cup at the proximal end.

This species seems close to M . variegatus in certain characters, but it can be easily
distinguished in the fresh condition by the very striking and unpleasant odor. In
M . variegatus the spores are 2-5 on a basidium and are larger than in the present species.
The shape of the basidia is also quite distinct as shown by Tulasne's figures. The name
of M. odoratissimus suggests this plant but the odor of that is described as pleasant
while the odor of this is decidedly otherwise; the basidia of that plant are short and
distinctly swollen at the distal end, and the spores have a distinct long stalk (Hesse).

The most important characters in distinguishing the present plant are the brownish
colored peridium which becomes red on being wounded, the large vesicular cells of the
peridium, and the exceedingly strong unpleasant odor.

This species is nearest the plant treated tentatively as M. rubescens, but careful
comparison shows them to be distinct. In section the dried gleba appears much alike
in the two plants except that in the Chapel Hill plant the tramal plates are much more
distinct. Sections, however, when soaked in water so that they regain approximately
the size when fresh show striking differences: in the Blowing Rock plant the "cavities"
are invariably stuffed with spores and the tramal plates are thin and rather inconspicu-
ous, while in the Chapel Hill plant the cavities are only partially filled, leaving the plates
broad and conspicuous. The structure of the tramal plates at maturity is quite
different: in the plant from Blowing Rock they are composed of entangled hyphae
which are partly gelatinized and which do not break apart easily when crushed; in
M. nauseosus they are composed of highly gelatinized hyphae which break apart when
crushed into separate units much like the cells in the subhymenial layer in Crucibulum.
Furthermore the spores in M. nauseosus are slightly larger and the basidia of the two
plants are quite different in shape.

8281. On and under beech leaves and humus, Nov. 4, 1927.
? Melanogaster rubescens (Vitt.) Tul.

Plate 109

During the summer of 1922 we collected in the mountains a single specimen of this
interesting plant. Unfortunately no notes were made on the fresh state. Our plant
in the dried condition is decidedly shrunken, subglobose, 11 mm. thick; surface dark
bay-brown, finely rivulose as if wrinkled in drying and marked on the lower half by a few
inconspicuous veins. Peridium simple, 320-450^ thick, composed of irregularly swollen
hyphae about 7.4-25/j thick and nearly hyaline under the microscope. Gleba very
hard, black and solid, with no cavities; when cut showing a shining surface with faint
whitish lines of the tramal plates outlining very large areas up to 1.5 mm. wide,
which are not sinuous, but box-shaped like parenchyma cells. These chambers are
solidly filled with the hard, black spore-mass, a condition different from species we are
referring to Rhizopogon. Tramal plates 35-1 10^ thick.

Spores smooth, black in mass, light brown under the microscope, subelliptic,
3.4-4.2 x 5.5-8.8^, the more mature ones showing the pale cup-like base as in M.
variegatus. Basidia short and thick, 7-9 x 12— 15yu, 4-8-spored, the sterigmata short.

PLATE 26

Rhizopogon truncatus. Type [above]
Calvatia elata. No. 2S43H [below]

HYMENOGASTRACEAE 41

Zellcr and Dodge have referred to Rhizopogon several species in which the glebal
chambers are solidly tilled with black contents at maturity. Of these R. pachyphloeus
seems nearest our plant, and Dodge seems inclined to put it provisionally in thatspecies.
To us, however, it differs strikingly in the far greater development of superficial fibers,
its much thicker peridium, thinner tramal plates and very much smaller glebal cham-
bers. From R. diplophlocus, also with solid contents, our plant differs in the simple
peridium, larger glebal cells, smaller basidia and larger spores. We think the plant is
best treated as a Melanogaster, but with only one plant and no notes on the fresh con-
dition, we think the specific name somewhat doubtful.

After referring to M. variegatus, Dr. Dodge in a letter says, "There is much closer
resemblance between your material and Melanogaster rubescens (Vitt.) as interpreted
by M. A. Curtis. He so determined (Ms. note in his copy of Schweinitz, specimen from
Schweinitz Herb, in Curtis Herb.) material collected by Schweinitz and referred by
Schweinitz to Rhizopogon acstivus Wulfen ex Fries.* The peridium of the Schweinitzian
specimen is about \ the thickness of your No. 5804 but it has a somewhat collapsed
appearance as do many of the older specimens which were repeatedly poisoned by the
old methods. The spores are much the same but darker than in your specimen. I have
never seen any authentic material of this species, so do not know whether Curtis's
interpretation is correct or not. Apparently Peters got the same thing in Alabama.
Your material has much lighter spores than anything I have ever seen in Melanogaster,
but I am not quite sure whether that is significant as in some other groups spores do not
assume their full color until late. For the present I am inclined to leave the specimen
in RJiizopogon pachyphloeus Z. & D. until I have had further opportunity to study
Melanogaster."

The relationship to M. rubescens is indicated by the basidia, which are similar to
those shown in Tulasne's figure, and by the appearance of the peridium.

Blowing Rock. Coker and party, No. 5804. Exposed on humus under chestnut and rhododendron,
August 26, 1922.

Melanogaster ambiguus Tul.

Plate 109

Dr. L. H. Pennington has kindly sent us a section of a plant found by him in New
York and reported by Lloyd as M. ambiguus (Myc. Notes, p. 1065, fig. 2005).

The dried section of the plant is slightly over a centimeter thick and has a con-
siderably wrinkled blackish bay peridium. In section the peridium is simple; about
300/i thick in dried condition and about 600ju thick when fresh (from prepared slide
of Pennington); composed of very irregularly interwoven threads with numerous
bladder-like swellings. Gleba jet black, the chambers rounded and filled with spores,
0.5-2 mm. wide; tramal plates very narrow, cream colored, contrasting strikingly with
the black spore mass.

Spores 7.7-10.4 x 13-18^, most about 8.5 x 15.9/j, broad-elliptic, more or less
pointed at both ends, with a distinct cup-like prolongation at the proximal end, deep
brown in color under the microscope. Basidia 8.5-11.8 x 22-26/z, club-shaped, with
2-4 sessile spores; cavities filled with interwoven threads, 3-3. 7/j thick, which give rise
here and there to basidia.

• Probably not in the sense of modern authors who use this name as an older name of Rhizopogon rubcuens Tul.

42 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

"Odor strong and penetrating, somewhat like that of onions" (Pennington).
This plant in the dried condition shows considerable resemblance to M. variegalus
but can readily be separated from that by the much larger spores.

Illustrations: Bucholtz. Beitr. Morph. und Syst. Hypogaeen (Tuberaceen und Gastromyceten),
pi. 1, figs. 22-24.
Hesse. Hypog. Deutsch., pi. 4, figs. 6-9; pi. 5, fig. 4; pi. 6, fig. 2.
Lloyd. Myc. Notes, fig. 2005.
Massee. Brit. Gastromycetes, pi. 1, fig. 5.
Smith. Brit. Basidiomycetes, p. 488, fig. 141.
Tulasne. Fungi Hypogaei, pi. 2, fig. 5; pi. 12, fig. 5.

New York. Syracuse. Pennington, coll. (Herb. N. Y. St. Coll. Forestry).

LEUCOGASTER Hesse

We have been able to study only one species of this genus, and take the generic
as well as other specific descriptions from Zeller and Dodge, who have recently mono-
graphed the American species, five of which they record from the eastern United
States (Ann. Mo. Bot. Gard. 11 : 389-408. 1924).

"Fructifications globose to irregular, hypogaeous to emergent, fleshy or waxy;
fibrils sometimes present, leading to rhizomorphs; columella, stipe, and sterile base
absent; peridium usually thin and fragile, sometimes rupturing at maturity; cavities
frequently polyhedral, usually filled with spores embedded in a gelatinous mass; septa
homogenous, with or without a distinct trama, often gelatinizing at maturity; basidia
from subglobose to ovoid and subcylindric, mostly 4-spored, sometimes 3- or 5-spored;
spores hyaline or slightly colored, with various surface markings embedded in a gelati-
nous spherical mass."

Key to the Species

Peridium double, thin, 60-80/* thick, white spotted with yellow, becoming reddish on drying; basidia
linear-oblong L- luteomaculatus (p. 42)

Peridium single, although sometimes with flocculent patches, then much thicker
Peridium thin (under 180/* when dry)

Tramal plates 40-50m thick; spores globose, 8-1 1m L. araneosus (p. 43)

Tramal plates 70-1 50M thick; spores subglobose, 8.5-11 x 11-14M L. caroliniamis (p. 43)

Peridium thick (over 200/*)

Septa thin, 60-120/x thick; basidia clavate, 22 x 8m; spores 11-13/* in diameter, sterigmata

short L. anomalus (p. 44)

Septa thick, 150-200/* thick; basidia pyriform

Basidia 20-24 x 12/*; spores 12-16/* in diameter; sterigmata short L. badius (p. 44)

Basidia 7-8 x 5-6/*, pedicellate; spores 8-10/* in diameter, sterigmata long

L. fnhimaculosus (p. 44)

Leucogaster luteomaculatus Z. and D.

"Fructifications globose, 2.5 x 1 cm., drying 1.5 x 0.8 cm., chalk-white with yellow
flecks (Thaxter's field notes), becoming ox-blood red to garnet-brown on drying, surface
uneven, shining; fibrils anastomosing, free in places, black, shining; peridium 60-80/*
thick, duplex, outer layer readily separating from the inner, leaving patches 20-30/*
thick' of large, septate, olive-brown hyphae, inner layer 40-55/* thick, of slender, reddish
brown, granular, closely woven hyphae; gleba milk-white when fresh, drying cinnamon
or clay-color; cavities globose to polyhedral, filled; septa thin, 50-60/*, hyaline, of
large, parallel, thin-walled, hyaline hyphae, scissile; basidia hyaline, 7 x 12/*, narrowly

PLATE 27

• •

# «

Leucogaster caroldciaxtjs. Xo. 8262 [above]. Dried plants
Melanogaster nauseosus. Xo. 8281 [below]. Fresh plants

HYMENOGASTRACEAE 43

oblong, 2-spored; sterigmata 2-iju long; spores light olivaceous under the microscope,
globose, minutely verrucose with a gelatinous sheath, 7-9/i in diameter.

"Under leaf mould in beech forests. Europe and North America. Summer.

"The duplex character of the peridium, the color both when fresh and upon dry-
ing, as well as the very thin peridium, should serve to distinguish this species from other
members of the genus."

Reported from Cranberry, North Carolina.

Leucogaster araneosus Z. &D.

"Fructifications globose, 0.6 cm. in diameter in preserved material, snuff brown to
bister; fibrils large but not prominent, few, half-immersed, somewhat branched; pe-
ridium 130-180/x thick, simplex, of closely woven, very slender, light brown hyphae;
gleba white with tawny spots; cavities subglobose, filled; septa thin, 40-50/* thick be-
tween hymenia, compact, of 3 layers, the middle layer being light brown, the other 2
layers hyaline; hymenial layer arachnoid; paraphyses none; basidia hyaline, granular-
guttulate, 8-10 x 6-8/i, pyriform, on pedicels up to 300/i long, 4-spored; spores hyaline,
globose, alveolate-reticulate, angles of alveoli projecting as blunt spines, in a gelatinous
sheath, 8-1 lju in diameter."

North Carolina. Summer.
Leucogaster carolinianus n.sp.

Plates 27, 28 and 108

Fruiting bodies hypogeal; subspherical, lobed, 5-17 mm. in diameter. Rooted
by several delicate, branched fibrils which may arise from either the top, sides or bottom
of the fruiting body. Mycelium fibrous and abundant in the soil, white. Peridium
50-220/t thick when fresh, about 60/* thick when dry, single, composed of threads 2.1/*
thick which soon collapse; nearly white in the young condition when first dug up, be-
coming antimony yellow or clay color or ochraceous buff or tawny in spots upon ma-
turing or drying ; peridium disappearing in small areas, either being worn away or slough-
ing off, thus exposing the whitish gleba; peridium in the young condition continuous
and homogeneous with the septa. Gleba pure white when fresh, becoming cream color
upon drying, composed of the numerous polygonal or subglobose sacs which are sur-
rounded by the tramal plates and filled when young with a gelatinous fluid in which the
spores are later embedded; upon ripening and drying some of the sacs come to contain
a large cavity, the walls of which are lined with the white spores, most of the sacs,
however, are filled with spores. Tramal plates, not measuring hymenium, 70-150/t
thick, usually about 100/1 thick, composed of the central region of densely packed, ir-
regular threads which, cut at the right angle, give the appearance of pseudoparenchy-
matous cells, quite variable in thickness, bordered on both sides by layers of irregular
threads, the walls of which become thickened and highly gelatinized as the plants
mature. Basidia arising from the layers which become gelatinized, long clavate, much
swollen at the distal end, 8.2-9.4/* thick at the end; apparently without a basal septum,
entire stalk 100-150/i long or perhaps longer; basidia not forming a definite hymenial
layer; sterigmata about 2/i long.

Spores 4 on a basidium, ovoid, rarely globose, heavily reticulated, covered with a
smooth layer of gelatinous material, 8.5-11 x 11-14/x (measuring sheath), very rarely
abnormally large ones 14.7 x 16.8/u, these being 2 on a basidium.

Taste faintly sweet, pleasant; odor slight, not unpleasant, faintly nutty, becoming
like machine oil as the fungus matures.

44 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

This species appears to be nearest L. araneosus Zeller and Dodge, which plant was
found by Dr. Thaxter in the mountains of North Carolina. Our plants are larger on
the general average than the size given by Zeller and Dodge, 6 mm. being their only
measurement, while ours are from 5-17 mm. thick. Their single measurement, how-
ever, comes within our range. In the color and the gross and minute structure of the
peridium the plants also agree. In the thickness and color of the tramal plates and the
size of the basidia and spores the plants are distinct. Zeller and Dodge give the tramal
plates as 40-50/* thick between hymenia while ours are 70-150/* thick, averaging about
100/* thick. In both plants the tramal plates are composed of three layers not including
the hymenium. In L. araneosus the middle of these three layers is light brown while
in our species it is white. The tramal plates in our plants are often scissile. The basidia
of L. araneosus are 6-8/* x 8-10/*; ours are 8.2-9.4/* thick and apparently without a
basal septum. The spores of L. araneosus are globose, 8-11/* in diameter; while those
of the present species are subglobose, rarely globose, 8.5-11 x 11-14/*. Dr. Dodge
has compared a specimen of our plant with his and thinks that there is no question but
that our species is new.
8262. Under deciduous leaves and earth, upper Laurel Hill, Oct. 19, 1927.

Leucogaster anomalus (Peck) Z. & D.

Hymenogaster anomalus Peck

"Fructifications globose to irregular, 1-2.5 cm. in diameter, cinnamon-buff, clay-
color, and tawny olive to Mikado brown above, Hay's russet and liver-brown below,
'glabrous, slightly lacunose, often with a root-like strand of mycelium at the base,
sterile base obsolete or nearly so, odor slight, not disagreeable,' (Peck) ; per-
idium 240-520/* thick, grenadine to English red near the surface to hyaline within,
composed of very slender interwoven hyphae; gleba amber-brown to Sudan brown,
sometimes lighter; cavities large, mostly more than 1 mm. in diameter, subglobose to
irregular, empty; septa 60-120/*, composed of very closely woven, hyaline hyphae, not
scissile; cystidia none, paraphyses clavate, granularly guttulate; basidia clavate,
22-24 x 8/*, hyaline, 4-spored; spores almost sessile, dilute cream-colored to hyaline,
globose, 11-13/* in diameter, uninucleate, surface pitted, giving the appearance of
hexagonal reticulations, surrounded by a hyaline, gelatinous sheath.

"Hypogaeous, in woods. District of Columbia. August to September."

Leucogaster badius Mattirolo

"Fructifications subglobose, 1 cm. in diameter, raw sienna to raw umber, surface
pitted, glabrous; stipe very slender, from a very slight, inconspicuous, sterile base;
peridium 200-340/* thick, ochraceous-bufi, stupose, composed of very slender hyphae;
gleba raw umber; cavities polygonal, filled with spores in a gelatinous mass due to
deliquescence, 0.7-1.0 mm. in diameter; septa 160-200/* thick, composed of compactly
woven, gelatinous, cream-colored hyphae; cystidia none; basidia hyaline, pyriform,
20-24 x 12/*, 4-spored, guttulate; spores almost sessile, subglobose, 12-16/* in diameter,
cream-colored, surface pitted, giving the appearance of hexagonal reticulations, sur-
rounded by a hyaline, gelatinous sheath.

"On the ground, under leaves. Italy and New York, July and August."

Leucogaster fulvimaculosus Z. & D.

"Fructifications globose, 2.0-2.5 cm. in diameter, drying cinnamon-buff, spotted
with tawny; fibrils not prominent, few, con-colorous; peridium 375-425/* thick, simplex,

PLATE 28

Leucogaster caroliniaxus. No. 8262. X 44

HYMENOGASTEACEAE 45

compact, of closely woven, slender hyphae; gleba warm buff or lighter; cavities globose
or polyhedral, half tilled; septa 150-180*1 thick, hyaline, of closely woven slender
hyphae, not scissile; basidia hyaline, 7-8 x 5-6^, pyriform, on pedicels about 90/x long,
3-spored; sterigmata conical, 2p long; spores hyaline in preserved material, brown in
dry material, spherical to ovoid, verrucose to reticulate, leaving a pore where detached
rom the sterigma, inclosed in a gelatinous sheath, 8-10ju in diameter.

"Damp woods. New York. Summer.

"The peridium of this species appears to be variable in thickness, new cavities
seeming to form in the inner layer as the fructification increases in size, with the hyphae
next the basidia-bearing hyphae gelatinizing and finally disappearing. A study of
young material is very desirable for further interpreting this phenomenon."

HYMENOGASTER Vitt.

Subterranean or partly exposed; peridium without superficial veins, not easily
removable, in our species white or nearly so; glebal chambers empty at first, sometimes
stuffed at maturity; spores oval, lemon-shaped or elliptic, in most cases pointed at the
distal end and rough, in some species rounded at the distal end and smooth, but inclosed
in a hyaline sac. The spores seem to be the only generic character of any value and
they are of two distinct types (if descriptions can be trusted).

Except for H. anomalus Pk., which Zeller and Dodge have transferred to Lenco-
gaster, only one species has been reported from the eastern United States. This was
determined by Lloyd as H. decorus, but we are transferring it to H. Thwailesii. We are
herewith adding one other.

Key to the Species

Odor very strong, not pleasant; spores 6.5-7.4 x 10-13/i, surrounded by a sac H. foctidus

Odor slight; spores 13-15 x 24-28/1 E, Thwailesii

Hymenogaster foetidus n. sp.

Plates 17, 18 and 110

Plants partly or entirely buried in soil under humus, single or cespitose or often
fused, 0.3-1.3 cm. wide by 0.3-0.7 cm. high, when fresh, shrinking to about two-thirds
this size on drying, rooting by several basal rhizomorphic strands; subspherical or
lobed, fleshy, odor when fresh very strong and persistent, not pleasant, to some sug-
gesting aniline oil, to others witch hazel. Peridium 0.5-0.8 mm. thick, simple but
often partly covered with white byssoid mycelium, whitish when fresh, changing to
creamy buff on drying, not changing color when cut or wounded; tender and fragile
when fresh, becoming only slightly toughish on drying; smooth and without ridges or
fibrils, but often encrusted with sandy particles, becoming wrinkled on drying ; composed
of much entangled, densely packed, thin- and thick-walled threads, the latter often
terminating in large spherical swellings 37-55^ thick, which strikingly resemble the
oogonia of certain species of Cystopus, except that the swellings are usually empty.
Gleba whitish at first, changing'to pale green and then to light brown with a greenish
tint, and finally blackish brown on drying; without any signs of a sterile base. Tranial
plates arising from the peridial wall and passing inwards in a more or less radial fashion,
in some uniting in the center to form a fairly distinct columella. Cavities of the gleba
subrotund in the central region, up to 0.6 mm. wide, becoming narrow and radially
elongated toward the peripheral region; empty at first, becoming nearly or completely
filled with spores. Hymenial layer concolorous with the tramal plates; composed of

46 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

young and old basidia 3-7 x 16-25/u long, which, when young, are cylindrical but which
collapse after the spores are formed.

Spores 2-4 to a basidium, smooth, elliptic, deep brown under the microscope,
6.5-7.4 x 10-13,u (not counting the utricle), with an open cup on the proximal end (at
first appearing like two prongs), the spore body surrounded at some distance by a very
thin, hyaline, bladdery utricle, which is in contact with the spore only at the cup end.
The surface of the utricle is faintly rugulose.

This plant clearly belongs in the Hymenogastraceae in the obvious origin of the
tramal plates from the peridium. Of the genera in this order it seems closest to Hy-
menogasler in the absence of fibrils from the sides and top of the plants and in the
structure and shape of the spores, a utricle being present in several species of that
genus. However, the appearance of the gleba is more like that of Hysterangium, with
its long, radially elongated chambers, stuffed or nearly so at maturity, radiating in
many cases from a distinct, subgelatinous, central column. The complete continuity
of the tramal plates with the peridium, however, is so striking that the plant cannot be
placed in the family Hysterangiaceae.

We have carefully examined the descriptions of the thirty-four species of Hy-
menogaster listed in Saccardo, Vol. 19, and find that our plant can be separated from each
of them by obvious characters. In the color of the gleba, the shape of its cavities and
the shape and size of the spore body, our plant resembles H. ulriculatus Harkness, but
that can be separated easily by the chocolate brown peridium and the fact that the
utricle of the spore is winged or angled. Moreover, no mention is made by Harkness
of any odor, while that of our plant is so strong and distinct as to be easily noticed
by the most casual observer. So penetrating in fact is the odor that the plants, if
partly uncovered by rain, can be located in the woods before being seen (10 or 15 feet
away). A pitcher of water put in the ice box with a dish of these plants was so im-
pregnated with the odor as to be nauseating. The present species is apparently nearest
H. Thwaitesii Berk, and Br., in that both species are externally white and internally
brown, and the spores of both plants are covered by a utricle. The two plants, how-
ever, can be separated by the distinct difference in the shape and structure of the spores.
Those of H. Thwaitesii are larger and the proximal end is without a cup. Moreover,
the utricle is attached at both ends of the spore instead of only one end as in our plant.
Also, no mention is made of odor in H. Thwaitesii. The New York plants which we
are referring to H. Thwaitesii differ strikingly from the present species in the thinner
peridium, the empty chambers of different shape, and much lighter color of the gleba.

Among our plants the present species can be easily recognized when fresh by the
peculiar and intense odor and in all conditions by the distinct utricle covering the small
spores. Hymenogaster decorus Tul. also has spores with a rounded distal end or very
obscurely pointed and surrounded by a utricle, but it is a much larger plant, has much
larger spores and is nearly odorless. Dendrogaster connectens Bucholtz (Hedwigia 40:
316. 1901) is easily different, as shown by the brown surface, yellow gleba, more
extensive sterile tissue, and much larger spores. No odor is mentioned.

7467. Partly buried in soil by branch under beech trees, by old Hillsboro road about 4 miles from Chapel

Hill, July 30, 1924.
7473. Buried in soil under humus or partly unearthed by heavy rain around the base of a tulip poplar,

by branch behind athletic field, August 4, 1924.

CM

W
H
<
h-1

o

«

HYMENOGASTRACEAE 47

7469. On damp soil by Battle's Rranch. August 4, 1"24.

7580. By a tulip poplar, Nov. 27, 1924. One of the plants in section showed a distinct, hyaline,
central columella.

Hymenogaster Thwaitesii Berk, and Br.

Plate 109

A plant collected by Mr. A. H. \V. Fovah among roots of sedges at Syracuse, New
York, has been determined and reported by Lloyd as Hymenogaster decorns (Myc.
Notes, p. 1166). We have a bit of this collection and on comparing it with Tulasne's
original description we are convinced that it is not that species but rather an American
form of II. Thuaitcsii, a plant described from England and which has apparently not
been found since.

The New York plants are very small, in the dry state about 3 mm. or less. The
interior is not at all purplish but sordid ochraceous. The chambers are comparatively
large, about 3-4 to a mm. and rounded and largely empty. Surface glabrous, pallid
ochraceous or paler, much wrinkled on drying.

Spores oval to subelliptic, with distal end rounded or somewhat pointed, surrounded
by a hyaline sheath, the body of the spore 9.2-11 x 11-17.3/u.

The spores agree very well both in size and structure with those of the type of H-
Tkumiesii, a preparation of which was kindly sent us by the Director of Kew Gardens.
We find the latter to be 8.5-11 x 14-20/z, not counting the irregularly crumpled, hyaline
sheath surrounding them, more or less pointed at each end. The English plant has
spores prevailingly more pointed than the American, as will be seen from our figures,
but the general characters are so similar that the American plant had best be considered
only a form. The original description of H. Thwaitesii gives the spores as globose,
but this is not the case. Tulasne's figures also taken from the type are drawn somewhat
more globose than we find them to be. The spores of decorus are much larger and the
gleba is purplish. Also the plant itself is much larger.

Illustrations: Tulasne. Fungi Hypogaei, pi. 10, fig. 11.

New York. Syracuse. Povah, coll. "Among the roots of sedges in oak woods within an inch of the
surface of the ground."

OCTAVIANIA Vitt.
(Including HYDNANGIUM Wallr.)

Fruiting bodies subglobose, oblong or lobed; exposed or underground. Peridium
thin, nearly smooth, not netted with veinlike strands. Gleba fleshy, elastic, intimately
connected with the peridium and not separable from it; cavities rather large, hollow
(at least when young) ; sterile base present or absent. Basidia 1-4-spored; spores spheri-
cal or subspherical, with warted to reticulated walls.

Two species have been reported from the eastern United States and we are adding a
new one. To separate this genus from Hydnangium has proved practically impossible.
We agree with Lloyd (Myc. Notes, p. 269; also p. 1139) that it is best to unite them and
have done so in the above description. In this last publication Lloyd reunites the
genera and treats all the species known by him.

48 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

For cytology of spore formation in Hydnangium carneum see Rhuland (Bot.
Zeit. 59: 187. 1901). In this plant Rhuland finds basidia in a single fruit body to
have 2, 3 or 4 spores. There are 4 nuclei in each basidium. In the two-spored cases
each spore may contain two nuclei or one may contain one and the other three. In the
four-spored cases each one contains a single nucleus. In the spores further division
of the nuclei may take place producing nuclei up to six. Bausch (Bot. Zeit. 18: 352.
1926) states that this behavior of nuclei, that is more than one entering a single spore
from the basidium is the only case proven. For Hydnangium carneum also see further
Bambeke, Mem. Acad. Roy. Sci. Belgique 54: 1903 and Klika, Mykologia 2:
143, 3 text figs. 1925. See also Patouillard, Note sur trois especes d 'Hydnanghim de
la flore du Jura, Bull. Soc. Myc. France 26: 199, figs 1-3. 1910.

Key to the Species

Plants bay or red-brown O. Ravenelii (p. 48)

Plants purplish on exposure O. purpurea (p. 50)

Plants gray to bluish green 0. asterosperma (p. 51)

Octaviania Ravenelii (Berk.) Lloyd

Hydnangium Ravenelii (Berk.) Curtis
Hydnangium Stepkensii var. Ravenelii Berk.
Octaviania Stephensii var. Ravenelii (Berk.) De Toni

Plates 29 and 110

Plants superficial or partly underground, gregarious and often crowded, subglobose
to oblong-lobed, usually flattened when large, 1-3 cm. wide, 0.6-1.5 cm. high; bay red
to chestnut or Sanford's brown when fresh, paler and more or less plicate underneath;
surface quite smooth, dull (not flocculent) ; rooting by a small white myceloid base,
no fibrils present; peridium wall simple, 130-150/j thick, its context reddish brown under
microscope, consisting of much tangled, closely packed hyphae. Gleba creamy white,
becoming pale rose when cut, exuding a white, tasteless milk which also becomes faintly
rose colored and more watery upon exposure to air; cavities of the gleba large, about
0.3-1.5 mm. long or wide, labyrinthiform, quite hollow until deliquescence and easily
visible without a lens; septa 110-190/* thick, consisting of (a) closely packed, irregular
hyphae, 3.7-6.2/* thick, (b) elongated, laticiferous sacs, and (c) the two conspicuous
hymenial layers which are 40-50/i thick and without cystidia. Odor when fresh rather
faintly aromatic and very like that of a certain group of Lactarias, e. g., L. theiogalus,
as also noted by Berkeley for H. Stephensii. When rotting the odor is mildly fetid and
moderately strong, not at all like the pleasant odor of Rhizopogon rubescens.

Spores spherical, 11.8-16/* thick, with a short mucro and large oil drop, the surface
set with irregular, imperfectly reticulated warts and ridges almost exactly as in certain
species of Russnla and Lactarius. Basidia clavate or spindle-shaped, 8-13 x 45-65/1
counting the sterigmata, which are 7.5-14/* long and only two in number.

Not rare in Chapel Hill on sandy soil mixed with moss and grass in low open woods
among pines. The plants are seated on the surface and never more than about one-
third submerged. After maturity the plants do not deliquesce as in Rhizopogon, but
begin slowly to decay, the gleba and peridium becoming water-logged, the former getting
much darker and usually spotted or covered with a blackish brown mold, the gleba not
changing color, slowly the tramal plates break down and the gleba becomes a tough and
elastic jelly, the disorganization proceeding gradually from the top and sides. Fissures

PLATE 30

f

9

Octaviania purpurea. No. 7456 [top] ; No. 7468 [below]

HYMEN'OGASTRACEAE 49

next appear in the top of the peridium and collapse begins as the jelly is slowly dissolved
out by the rains. Even when the plant is half rotted off above and shrunken like a
dried plum, the structure of the gleba is still obvious below and all the plant remains
tough and elastic. Weeks and probably months are required to complete the dis-
organization.

Reported from North and South Carolina, Florida and Alabama.

Though Ravenel's plant was described as white by Berkeley (Grevillea 2: 33.
1873, as a var. of //. Stephensii) we have no doubt that our plant is the same. We have
compared a collection labelled Octaviania Ravenelii from Alabama by Earle (N. Y. Bot.
Gard.), who notes that the color was "uniform terra cotta red, after exposure the tops
bleach to a dirty white." He also notes that the plants were growing singly on the
ground in a pasture and were usually 1-2 cm. broad (up to 3 x 4 cm.). Our plants are
the same as these in all characters, including the spores which we find to be in Earle's
plants 11— 14/u, rarely up to 16.6/i, including the tubercles. Hydnangium Stephensii
Berk, and Br. (Ann. & Mag. Nat. Hist. 13: 352. 1844) was transferred to Octaviania
by Tulasne, but the characters that distinguish these two genera are very obscure.
(See note under the genus.)

The American plant is closely related to H. Stephensii, but the spores easily sep-
arate the two species. The spores of H. Stephensii are short-elliptic and with slender,
sharper spines and are not reticulated. Our fig. 7 on pi. 110 is drawn from a slide,
which was generously prepared for us by Miss Wakefield from the type atKew. Tulasne
and Massee draw the spores as elliptic, but with the spines more prominent than they
appear to us. Quelet (Champ. Jura et Vosges, pt. 3, p. 18, pi. 1, fig. 9) shows the spore
as subspherical (13/u thick) with small spines. It is interesting that he gives the color
of the plant as whitish, the very character that made Berkeley describe the American
plant as a variety of his "dark rufous" H. Stephensii. This suggests a similar fading
in age. Corda shows the spores of 0. Stephensii as quite spherical and with short,
sharp spines, not reticulated (6: pi. 7, fig. 67). Patouillard's drawing of spores (Bull.
Soc. Myc. France 26: 201, fig. 1. 1910) of what he calls a form of Stephensii {Hyd-
nangium galathejum) are as he notes just like those of the type of Stephensii. He
further notes that in the type as in the form the basidium has but one sterigma.
This makes a further distinction between Octaviania Ravenelii and 0. Stephetisii.
In the fresh state a section of our plant showed a narrow, plug-like sterile base
about 1-2 mm. thick. From this base two or three lines (in section) radiated and
were plainly visible for about half way to the surface. When cut the milk exuded
much more abundantly along these lines and from the sterile base. There is no
sign of splitting in the tramal plates. In his original description of H. Stephensii,
Berkeley does not describe the juice as turning red then yellowish on exposure
as stated by Rea (British Basidiomycetae, p. 28) but says, "substance when cut and
exposed to the air soon acquiring a red tinge, which is not however permanent and in
young specimens vanishes almost entirely in drying, in which state the hymenium is
cream colored." This is exactly the color change in our plants.

Since writing the above we have seen notes on this species in the fresh condition
by Prof. Povah of Alabama (Lloyd Myc. Notes, p. 1140. 1922). His observations
are in close agreement with ours in regard to milk and color changes. The odor he
describes as "suggesting to some strawberries, to another the odor of a freshly dug

50 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

sweet potato." The odor of the sweet potato is not very different from the Lactarius
odor noted by us.

1445a. Beside a low place near Mason Farm, Oct. 28, 1914.

6026, 6042. On damp ground in pine woods on Durham road, Jan. 7 and 12, 1923.

6061. On ground in pasture near pines, Feb. 9, 1923.

Also Nos. 6002, 6009, 7200.

South Carolina. Santee Canal. Ravenel, No. 883; Curtis, No. 2576. (Curtis Herb.)
Society Hill. Curtis, coll. (Curtis Herb.).

Also from South Carolina by Ravenel in Fungi Car. Exs. No. 71 (Philadelphia Acad. Herb., as
H. Stephensii var. Ravenelii).
Alabama. Auburn. Earle. (N. Y. B. G. Herb, and U. N. C. Herb.)

Also collected in Alabama by Povah, as noted above.
New York. Ithaca. Whetzel, coll. (Cornell University Herb, and U. N. C. Herb.). Spores 11-16.6^
thick, with an imperfect reticulum. Basidia 2-spored.

Octaviania purpurea n. sp.

Plates 30, 31 and 111

Fruiting bodies hypogeal, single or cespitose, subglobose, often irregularly lobed ;
fresh plants 0.6-1.8 cm. high by 0.6-2.5 cm. wide, drying to about two-thirds that size,
fleshy; rooting by numerous mycelioid, branched fibrils which extend out from several
points of attachment on the base. Peridium white throughout when young, turning
vinaceous to deep vinaceous when exposed or wounded ; retaining the vinaceous tint on
drying; 1-2 mm. thick when fresh, drying to about one-quarter this thickness; composed
in large part of small thin-walled, densely packed and considerably entangled threads
3-4/x thick, and a few much larger thick-walled threads, 11-16// thick, which often
terminate in large globose cells. Gleba pure white when young, acquiring an olive tint
as the hymenium and spores begin to develop, passing through olive brown to deep
blackish brown as the spores mature. Tramal plates pure white, becoming only slightly
discolored upon maturity and contrasting strikingly with the nearly black spores and
the more or less deep purple peridium; arising from the peridium and composing in the
young state by far the greater part of the gleba, persisting in mature plants as very
thin white byssoid plates. Cavities of gleba first arising as narrow, pouch-like spaces
which by growing and anastomosing, form labyrinthiform spaces, which, in section,
often appear in clusters, 3-8, separated by rather wide tramal plates, thus presenting
the appearance of several small cavities within a large one; cavities becoming only
partially filled with spores. Odor of fresh plants faintly like an Irish potato. Plants
kept covered in the ice box for several days gave an odor strongly suggestive of oysters.
Taste not unpleasant.

Spores subsessile, subspherical, sometimes considerably elongated at the proximal
end, angular-warted, 9. 5-12. 5m thick, with a pedicel about as long as the diameter of
the spore. Hymenial layer, lining the cavities, composed of young and old basidia
which are very peculiar in that in the young state they are thick, rather short cells,
7.4-11.2 x 20-25;ii, which by elongating into a narrow tube, 4.4-5. 5yu thick, more than
double their length; mature basidium 5.5-7.8 x 30-44^, jug-shaped, bearing one apical
spore; collapsing after the spores are formed.

Suitable material being at hand, it was thought of interest to study the nuclear
behavior in this single-spored basidium for comparison with that of the normal four-
spored basidium. The material for this study was prepared as follows: small slices
about a half millimeter thick of the youngest plants were cut and put in Flemming's
weak chrom-acetic acid where they were allowed to remain over night. The material

PLATE 31

■

1

€

■*v

•!>■*

^

**

H

Erf-

$&*& r '" 2

*^BlHH

/ VMH

. j

Octaviania purpurea. No. 7468. X 400

[above]
Geaster velutinus. X 36 [below]

Astraeus hygrometricus. No. 7441. X

100 [above]
Disciseda CANDIDA. No. 7507. X 100

[below]

HYMENOGASTRACEAE 51

was dehydrated and infiltrated in paraffin in the usual manner for very delicate objects.
Sections were cut S microns thick and stained with Haidenhain's iron alum haematoxy-
lin and by Gram's method, the latter giving much better results.

It was found that the nuclear behavior agreed in the essential details with that
described for typical Basidiomycetes with four-spored basidia except that three of the
nuclei remain in the basidium and degenerate, the fourth entering the spore which is
always uninucleate.

In the young basidium, which has not developed the elongated, tubular process,
there were quite often seen two nuclei which might be located near the base or in center
or in position as shown in figure 4. Numerous stages were seen in which the two nuclei
were apparentlv fusing. When the basidium is in a somewhat later stage of develop-
ment (fig. 5) it is usually found to be uninucleate, the nucleus now being considerably
larger than either of those shown in figure 4. This large nucleus soon becomes some-
what vesicular in structure, the linin threads with chromatin granules on them showing
up conspicuously (fig. 6). A comparatively large spindle is formed on which the chro-
matin granules arrange themselves (fig. 7). The first mitosis seems to be followed im-
mediately by the second in which the spindle is considerably smaller and the chromo-
somes much less distinct (fig. 8).

Four nuclei are thus formed from the original fusion nucleus (fig. 9). Meanwhile
the basidium has sent out a tubular process into which the nuclei and some cytoplasm
migrate. At the tip of this tubular process a bud is formed which enlarges to form the
spore. One of the four nuclei passes into the spore (fig. 10). In later stages as shown
in figure 11, the spore is seen to be uninucleate; the three supernumerary nuclei left
in the basidium can hardly be distinguished — so poorly do they absorb the stain. They
are doubtless degenerating.

Our plant is evidently nearest Hydnangium monosporum Boud. and Pat. (See
Boudier; Icon., pi. 193, and Patouillard; Tab. Fung. No. 692) as shown by the strongly
cespitose habit and single-spored basidia with rough spores. That is easily separated
by ochraceous color, larger size, and minutely spiny spores. Gallacea Scleroderma
(Cooke) Lloyd ( = G. violacea (Cooke & Mass.) Lloyd) , a violet plant from New Zealand
and related to Hysterangium, has recently been examined microscopically by Cun-
ningham, and is quite different from our 0. purpurea (Trans. Brit. Myc. Soc. 9: 193,
pis. 9 and 10. 1924).

The genus Leucophleps Harkness {Leucophlebs, as the spelling was corrected by
Roumeguere) contains a species L. magnala which, in its one-spored basidia, resembles
our plants, but which can be easily separated by the partial absence of the peridural
and the smooth spores. Appearances are against Leucophleps being a good genus.

7456. Plants partly or entirely buried in damp soil by Battle's Branch below Forest Theater, July

25, 1924.
7468. Abundant on washed soil by Battle's Branch, Aug. 4, 1924.
7508. On ground and slightly buried under a sweet gum tree near Meeting of Waters, Aug. 27, 1924.

Octaviania asterosperma Vitt.

Plate 110
Dr. Fitzpatrick has kindly sent us a single plant of this species from Ithaca (entered
below), mentioned by Lloyd in Mycological Notes, p. 1141. We find it to agree with

52 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

the descriptions by Tulasne and Boudier so far as can be seen from specimen kept
in formalin. The peridium is irregularly areolate, salmon colored, usually about 100/j
thick, composed of densely packed tangled threads. Tramal plates continuous with
the peridium, considerably thinner than the peridium and much lighter colored, ap-
pearing almost white; glebal chambers rounded or elongated, usually filled with
spores, 0.5-1.5 mm. wide. Spores round, 13-17/* thick, light brown under the micro-
scope, very dark brown in mass, conspicuously set with large pyramidal spines and
often with a short pedicel.

The following description is from Boudier, Icon. Myc. 4: 97:

"Peridia white, rounded, solitary or cespitose, and more or less irregular, covered
with a light tomentum which is separated into small, tomentose, flatfish warts, white at
first, then turning smoky gray; cortex turning more or less bluish green by cold or ex-
posure to air; the peridia are attached to the earth by root-like, mycelial filaments which
form a very visible base. The gleba is full, firm, formed of rounded chambers, filled
with spores and separated from each other by the whitish plates which seem to arise
from the basal portion, and which are colored equally green by cold. The spores are
perfectly round, echinulate, brownish drab when seen one at a time, and purplish brown
when seen in mass, which makes a section of this species appear finely marbled with
white by the plates; they have a large internal oil drop often divided, and are often
furnished with a pedicel; they measure 15— 18/x in diameter with the spiny warts."

"This species seems rare; I have found the specimens which I have figured in damp
woods of the forest of Montmorency in Sept. ; it was there in numbers under the leaves
at the foot of a clump of ash trees."

Illustrations: Boudier. Icon. Myc, pi. 191.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 3 (in error as O. asterospora) .
Lloyd. Myc. Works, pi. 205, fig. 2168 and fig. 2169 (copied from Boudier).
Tulasne. Fungi hypogaei, pi. 11, fig. 1.

New York. Ithaca. Fitzpatrick, No. 9969. (Reported by Lloyd in Myc. Notes, p. 1141.)

SECOTIACEAE

Fruit body above ground and with a short or long stalk which extends as a col-
umella entirely through the peridium and is connected with it above; gleba chambered,
the tramal plates arising like the gills in an agaric and growing into a gill chamber,
anastomosing and crumpling to form numerous irregular cavities which are lined on the
inner side by a typical agaricoid hymenium ; tramal plates persisting after the matura-
tion of the spores; capillitium absent. Dehiscence ventral around the stem with sub-
sequent expansion and in some cases longitudinal cracking, or there may be no true
dehiscence (as in RJiopalogaster), the very thin peridium disappearing more or less by
collapse. Basidia clavate with distinct apical sterigmata.

The family as above described is so modified from that of Fischer as to include
only the genera Secotium and RJiopalogaster (possibly also Macowanites). We are
following Fischer in not considering Elasmomyces as distinct from Secotium. This is
about as suggested by Cunningham (1. c, p. 223). A very interesting family with
characters suggesting the agarics and apparently connecting them with the Hymeno-
gastraceae, which they resemble in later stages of glebal development. Only two
genera have been found in the eastern United States. It is interesting to compare
here the peculiar genus RJiacophyllus B. & Br. (Trans. Brit. Myc. Soc. 11 : 238. 1926)
from the tropics, which externally has the exact appearance of a small, delicate agaric
with conic cap. Petch has recently been able to study the plant in all stages. Instead
of gills there are borne on the underside of the cap more or less linear groups of bulbils
which germinate easily and are the only reproductive bodies. These bulbils arise
separately inside a delicate fundamental tissue. They may be at first hollow, soon
becoming solid through growth of the palisade-like central tissue. This tissue produces
on the inside subspherical cells in rows. It is not stated from what cells of the bulbils
hyphal threads originate in germination.

Key to the Genera

Peridium tough and persistent Secotium

Peridium very delicate and disappearing in part at maturity Rhopalogasler

SECOTIUM Kunze

Peridium above ground with a long or short stalk, which extends entirely through
the sporiferous portion as a stout columella which is continuous above with the pe-
ridium. Gleba spongy, cellular; tramal plates arising from the peridium wall and also
connected with the top or greater part of the columella and filling the space between
them, somewhat lamellate, very sinous with a more or less horizontal direction sug-
gesting an unopened agaric with sinuous gills; capillitium not present. Dehiscence
basal and longitudinal, the peridium usually separating slightly from the stem below
and in some cases expanding more or less, after the manner of a young agaric. Basidia
clavate with 2-4 apical, stipitate, smooth or rough spores.

For a good drawing of the hymenium of 5. krjukmvense, a European species with
rough spores, see Bucholtz, 1. c, pi. 3, fig. 4.

53

54 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

The genus seems to be particularly abundant in Australia and New Zealand, from
which Cunningham (1. c.) reports 16 species. He adds the following facts to our knowl-
edge of the cytology:

"The hyphae of the columella, stipe and peridium are invariably binucleate.
The basidia at first are binucleate, these nuclei fuse, and a slightly larger fusion-nucleus
is formed. This takes up a position in the distal end of the basidium; there it divides
twice. The first division precedes the formation of the sterigmata, the second succeeds
their appearance. When the sterigmata are about half their normal length, spores
begin to appear on them, and when they are full length, the spores are about half size.
When the spores are about one-quarter their normal size, a nucleus migrates into each,
divides mitotically, and the spores become binucleate, a character constant in each
of the numerous species examined. The spore attains its full size before it changes
colour ; at maturity the epispore becomes coloured some shade of brown, the depth of
colour depending on the species.

"Clamp connections are abundant in the tissues of the stipe and partial veil

"On germination a germ tube protrudes, usually from the end of the spore oppo-
site to that by which it was attached to the sterigma; this branches repeatedly to form
a mycelium, the cells of which are septate and binucleate."

Conard and Cunningham, working on different species disagree as to the presence
of a veil, Conard claiming the presence of universal veil and absence of a partial one
and Cunningham stating just the reverse.

LITERATURE

Conard. The Structure and Development of Secotium agaricoides. Mycologia 7: 94, pi. 157 and 1

text fig. 1915.
Cunningham. A Critical Revision of the Australian and New Zealand Species of the Genus Secotium.

Proc. Linn. Soc. of New South Wales 49: 97, pis. 12-15. 1924.
Cunningham. The Structure and Development of Two New Zealand Species of Secotium. Trans.

Brit. Myc. Soc. 10: 216, pis. 11 and 12. 1925.
Lohwag. Entwicklungsgeschichte und systematische Stellung von Secotium agaricoides (Czern.) Holl.

Osterr. Bot. Zeitschr., 1924, p. 161, pi. 2.
Setchell. Two New Hypogaeous Secotiaceae. Journ. Myc. 13: 236. 1907.

Secotium agaricoides (Czern.) Hollos
S. acuminatum Montagne
S. Warnei Peck
S. rubigenum Harkness

Plates 32 and 111

Fruit bodies epigeal, solitary or gregarious, usually distinctly heart-shaped, rarely
subglobose, with a distinct conical or rounded stalk which tapers into a thick strand
and anchors the plant in the ground; very variable in size and shape, 1-5.5 cm. wide by
1.5-6 cm. high. Peridium nearly pure white when young and fresh, turning straw color
to cinnamon-buff or leather color upon maturing and drying; single-layered, 1-3 mm.
thick, nearly smooth in very young specimens, covered at maturity with inherent scales
which may resemble fish scales or may be very irregularly arranged; fleshy and tender
when young, becoming rather tough and fibrous upon drying. Gleba nearly pure white
when young, becoming yellow and finally brown; glebal chambers somewhat labyrinthi-
form, varying in width from a fraction of a millimeter up to 1 mm. Tramal plates

PLATE 32

Secotium agaricoides. Xo. 7091 [two small plants] ; No. 7119 [others]

SECOTIACEAE 55

about 75m thick, persisting in the mature and dried specimens, forming lamella-like
folds. Odorless when young; taste sweet and nutty like that of Boletus edulis. Edible
when young.

Spores (of Xo. 7091) smooth, ovate, yellow under the microscope, 5.5-6.8 x 6.6-8/1,
a few irregular, larger ones. Hymenium 25-30/n thick; basidia club-shaped, 4-spored,
7.4-9.3|i thick, no cystidia.

This is the first record so far as we can find of this fungus in the southeastern states,
but at the New York Botanical Garden we find a collection from Virginia by Dodge
(as 5. acuminatum). The only record we find of its occurrence in the Atlantic states
is from Pennsylvania (Barnett, as 5. Warnei; Peck, 1. c, 1882). It is apparently not
rare in the Mississippi Valley and westward, and is reported from Alabama and Texas.
This is the same as the plant described by Peck from Wisconsin as Lycopcrdon Warnei
(Bull. Torr. Bot. Club 6: 77. 1876) and later transferred to Secotium (Ibid. 9: 2.
ISSJi.

We have examined and compared specimens and spores of S. Warnei from Ne-
braska (X. Y. Bot. Gard. Herb.; H. J. Webber, coll.) and specimens and spores of S.
agaricoides from Hungary (N. Y. Bot. Gard. Herb., determined by Hollos) and find
that they both agree with our Chapel Hill plants. We agree with Hollos that S.
rubigenum Harkness (Bull. Cal. Acad. Science, 1886, p. 257) is the same as the present
species. A specimen at the New York Botanical Garden from California, collected by
Harkness himself, has spores smooth, oval-elliptic, variable in size, 4.8-6.8 x 7-10.5^.

Secotium arizonicum Shear and Griffiths (Bull. Torr. Bot. Club 29: 450. 1902)
seems to be a good species, as shown by the much larger spores (7.2-12 x 8.5— 15m)
of the type material in Washington (pi. Ill, fig. 14), and by the short, often aborted
columella (see note by Lloyd, Myc. Notes, p. 149). Secotium tenuipes Setchell (Journ.
Myc. 13: 239, pi. 107, figs. 4-8. 1907) and S. russidoides (Setchell) Lloyd (published
as Elasmomyces) are also good species, growing entirely concealed under leaves in
California. At the New York Botanical Garden is an abundant collection of the former
by Harper from Berkeley, California (type locality) . The plant (when dry) is small,
blackish red with a slender stem and a deep reddish brown gleba. The spores are
deep brown, smooth, ovate, 8.5-10.2 x 12-16//. Secotium australe Lloyd (Myc. Notes,
p. 788) is also apparently a good species, as the spores are said to be only 4-5/x thick.
Lloyd has seen the types of S. decipicns Peck (Bull. Torr. Bot. Club 22: 492. 1895)
and thinks it the same as the European Gyrophragmium Delilei (Myc. Notes, p. 150).
Later he uses the name G. decipiens and calls it the western form of the European
species (Myc. Notes, p. 196).

Illustrations: Bucholtz. Beitr. Morph. und Syst. der Hypogaeen (Tuberaceen und Gasteromyceten),
pi. 4, fig. 19.
Fries, Th. C. E. Sveriges Gasteromyceter, fig. 7.
Hollos. I.e., pis. 3, 4, 5, 6, and pi. 29, figs. 33-35.
Lloyd. Myc. Works, pi. 13 (as 5. acuminatum).

Lloyd. The Genera of Gasteromycetes, pi. 3, fig. 25 (as 5. acuminatum).
McDougall. Trans. 111. St. Acad. Sci. 15: fig. 3 (of McDougall article).
Peck. Bull. Torr. Bot. Club 9: pi. 9, figs. 6-11 (as S. Warnei). 1882.
Shear. Asa Gray Bull. 6, Xo. 6: 95. fig. 1, A-E. (asS. Warnei). 1898.
Sorokine. Rev. Myc, 1890, pi. 27, fig. 367; pi. 31.
Trelease. Morels and Puff-Bails of Madison, pi. 8, fig. 7 (as S. acuminatum).

56 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

7091. On ground in a pasture near Chapel Hill, August 24, 1923.
7119. Same location as above, September 11, 1923.

Virginia. C. F. Dodge, coll. (N. Y. Bot. Gard. Herb.).

Indiana. Lafayette Co. J. C. Arthur, coll. (N. Y. Bot. Gard. Herb., as 5. Warnei).

Dakota. October, 1908. Braendle. Fungi Dakotensis No. 49. (N. Y. Bot. Gard. Herb., as S.

acuminatum.)
Wisconsin. Madison. Cheney, coll. (U. N. C. Herb.). Spores 5.5-6.6 x 8-9. 7/j.

For other localities see Lloyd, Myc. Notes No. 14: 139. 1903.

RHOPALOGASTER Johnston

Johnston has shown (Proc. Am. Acad. Arts and Sciences 38: 61, one plate. 1903)
that the following species, commonly known as Cauloglossum transversarium, cannot be
retained in that genus which is really a synonym of Podaxon. He proposes for the
species the monotypic genus FJiopalogaster. Johnston defines the genus as follows:

"Fruiting body clavate, stipitate, traversed by a firm subgelatinous axil columella
continuous with the stipe. Stipe firm, erect with naked base. Peridium simple,
continuous with the stipe below and with the columella at the apex, more or less eva-
nescent-indehiscent. Gleba persistent. Tramal plates extending from the columella
toward the peridium. Basidia clavate, in groups, 4-spored, spores simple, borne on
well-developed sterigmata."

Rhopalogaster transversarium (Bosc) Johnston.
Cauloglossum transversarium (Bosc) Fr.

Plates 41 and 111

This plant of the southeastern Gulf states was first collected in the Carolinas but
was not well known structurally until the work of Johnston (1. c). It has been found
at Wilmington in abundance by Curtis and by Wood, and we have a collection from
Nag's Head, N. C. As our dried plants are not in good condition we copy the
following description from Johnston's paper, which see for much fuller structural
detail and discussion of synonomy:

"Narrowly to broadly club-shaped, 3-7 cm. high; the distal end of the columella
appearing at or near the apex, either as a shallow orbicular depression, or a slight pro-
tuberance. Peridium dirty brownish or buff yellow. When young the gleba is dirty
gamboge yellow; when exposed by injury becoming dirty olive brown, eventually dark.
Stem nearly white when fresh. Spores ovate-elliptical, 3.6-4.3 x 5.8-7.2^, yellow
brown, borne on long slender sterigmata. The plants grow out of the bases of living
or dead trees or upon rotten wood, stumps or fallen logs, or among rubbish on the ground
close by in wet pine lands. September-November. S. Carolina (Bosc, 1811, Curtis,
1857?); Santee Canal, S. C. (Ravenel); Wilmington, N. C. (Wood, November, 1880,
Curtis, November, 1846); Carolina (Berkeley, 1873); Gainsville, Fla. (Ravenel, 1878);
Eustis, Fla. (Thaxter, October, 1897); Tuskegee, Ala. (Beaumont, aestate, 1853)."

The specimen labelled Cauloglossum transversarium in the Philadelphia Herbarium
(Ravenel, Fung. Car. Exs. 79) is not this species but Sphaeria herculea Schw., according
to Ravenel.

PLATE 33

Calvatia rubro-flava. Nos. 7154, 7163 and 7107

SECOTIACEAE 57

Illustrations: Johnson. As cited above.
Lloyd. Myc. Works, pis. 12 and 40.

North Carolina. Wilmington. (CurtisHcrb.)

Nag's Head. Around roots of black gum at edge of fresh water, Sept. 18, 1927. Coker and

Braxton, colls. (U. N. C. Herb., No. 8261).
South Carolina. Santee Canal. Ravenel, coll. (Curtis Herb.).
Alabama. Tuskegee. Beaumont, coll. (Curtis Herb.). Also from Alabama (in sphagnum swamp)

by Earle. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.)

LYCOPERDACEAE

Fruit bodies exposed at all stages or subterranean until just before maturity.
Peridium consisting of more than one layer (very obscure in Calvatia rubro-flava, and in
some Lycoperdons where the outer layer may consist of only scattered particles or
flakes). Gleba at maturity breaking down into a dry powder which contains the spores.
Capillitium always present but sometimes (as in Disciseda) obscured by fragmentation
after maturity. Basidia borne in a distinct hymenium surrounding small chambers
or (in Disciseda) scattered in irregular groups without a distinctly organized hymenium
or chambers. Basidia club-shaped with four (rarely fewer) spores borne on apical
sterigmata; spores growing to full maturity on the basidia.

It will be noticed that this description does not include the genus Arachnion. We
have found it impossible to retain it and have any character left to determine the
Lycoperdaceae.

The lower part of the fruit body is often sterile and persistent and composed of
larger compartments than the fertile part above. The former is called the subgleba
and from it may extend up into the fertile tissue a more or less well developed column of
sterile tissue called the columella. In most species the spores escape from a single,
definite apical pore in the inner peridium. In Myriostoma there are several mouths,
in Disciseda a basal mouth and in Calvatia no definite mouth, the whole upper part
of the peridium flaking away. The family includes all the well known puffballs and
earth stars. The larger puffballs when young and white (before the spores begin to
ripen) are not only edible but very palatable and they should be much more used as
food.

For literature see under the order (p. 194) and genera.

Key to the Genera

Fruit bodies small, densely clustered on a common, leathery subiculum covering the substratum

Diplocystis (West Indian) (p. 143)
Not as above

Outer peridium (cortex) thin, mostly peeling off in flakes or wearing away; inner peridium thick,
firm and corky when dry, opening from above by irregular flaps and fissures, capillitium
of separate threads with thorny prickles and a few short branches . .Mycenastrum (p. 101)
Outer peridium thin, mostly peeling off in flakes; innerperidium opening by an apical pore, firm
and resilient, retaining its form for some time after maturity; capillitium of smooth,
separate threads with slender, pointed branches; mature plants loosened from place

of growth and rolled about by the wind (the tumblers) Bovista (p. 97)

Outer peridium thick or thin, scaling off in flakes or particles or wearing off by degrees or more or
less persistent; inner peridium usually flaccid and collapsing or breaking up as the spores
emerge; plants normally remaining attached to place of growth

Capillitium of separate threads with slender, pointed branches Bovistella (p. 94)

Capillitium of long interwoven threads that are not separable into unbroken units (in the
case of Calvatia breaking up into short pieces except in some specimens of C. data)

Peridium opening by a definite mouth Lycoperdon (p. 69)

Peridium irregularly ruptured by the scaling off of fragments above; plants large (at

times small in C. rubro-flava) Calvatia (p. 59)

58

LYCOPERDACEAE 59

Outer peridium thick, splitting at maturity into star-like rays from above downward and remain-
ing attached to the inner peridium by their united bases

Inner peridium with a single, apical mouth Ceaster (p. 102)

Inner peridium with several mouths and several pedicels Myriostoma (p. 138)

Outer peridium splitting horizontally around the middle, the upper part with the inner peridium
attached breaking away and leaving the lower part in the ground as obscure fragments
or an obvious cup Disciseda (p. 139)

CALVATIA Fries

Plants large to very large, globose, flattened or pyriform, etc., with or without a
thick, stalk-like, sterile base; cortex thin, even or areolated (absent in C. rubro-flava) ;
inner peridium thin, delicate, not opening by a pore but falling away irregularly in
scales and plates from above downward and exposing the densely woven capillitium
composed of long, much branched threads which at maturity in most species are easily
broken into short pieces; sterile base (nearly absent in C. maxima) concave above,
persisting a long time as a cup-shaped remnant, easily distinguished from Lycoperdon
by the irregular scaling away of the peridia at maturity, and in most species by the
fragility of the capillitium.

The sterile base is in most species sharply delimited by a diaphragm, and in this
case is obviously cellular. In C. rubro-flava and C. maxima there is no diaphragm
and the sterile base is homogeneous at maturity.

The genus includes the large puffballs, one species (C. maxima) reaching the enor-
mous width of three feet or even more. They are important sources of food and should
be used whenever found in the young condition with white interior, but note that the
edibility of C. rubro-flava is not known. A neighbor of ours had some cooked and
reported that they were too bitter to eat. They may be poisonous. In regard to the
edibility of puffballs, it is interesting to note that Porcher (Resources of Southern Fields
and Forests, 2nd ed., p. 699. 1869) quotes a letter from Ravenel as follows:

"It has been mentioned by medical writers that the spores of the puffballs have
narcotic properties, and it is an anaesthetic agent, acting somewhat like chloroform
when inhaled, but I have never experienced any effects of the kind from its use as a
vegetable. However, Dr. Harry Hammond, of Beech Island, S. C, writes to me,
'since writing to you, I and a number of others have made several meals on Lycoperdon,
and I think I have discovered in myself well marked evidences of a narcotic influence —
and two other experimenters have described similar sensations to me. I recollect also
to have heard from Mr. Mahan that a friend of his, a physician in Georgia, had been
seriously affected in this way by too large a meal on Lycoperdon.'1'

From the context it would seem that the puffball referred to is not C. maxima,
as stated, but C. cyathiformis, as the spores are spoken of as purplish. This note need
not deter one from eating these puffballs, as they are among the most esteemed of edible
fungi. The ripe contents of the larger puffballs have long had a reputation for stopping
the flow of blood (see, e.g., Lind, Danish Fungi as Represented in the Herbarium of
E. Rostrup, p. 401).

Literature

Cunningham. Gasteromycetes of Australasia. The Genus Cahaiia. Proc. Linn. Soc. N. S. W. 51:

363, pis. 23, 24. 1926.
Lloyd. French Collections of Calvatia. Myc. Xotes, p. 264. 1906.

L LIBRARV

60 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Lloyd. Japanese Collections of Calvatia. Myc. Notes, p. 548. 1916.
Morgan. Journ. Cin. Soc. Nat. Hist. 12: 165. 1890.
For other literature, see p. 194.

Key to the Species

Peridium apparently single, the outer peridium being represented by only a delicate furfurescence,
very thin; subgleba homogeneous at maturity; all parts turning yellow when bruised before

maturity; spores yellow-ochre in color C. rubro-flava (p. 60)

Peridium single, or at least not separable into two distinct layers, thick; gleba bright olivaceous

C. pachyderma*
Peridium double, outer layer thick, inner layer thin

Gleba olivaceous yellow C. lepidophorum*

Gleba purplish brown C. cretacea (p. 66)

Peridium double and separable at maturity into a thin outer and a thicker inner layer; subgleba, if
present, chambered at all ages; not turning yellow when bruised; spore color purplish or brown
or olivaceous yellow
Plants very large, almost entirely filled with the gleba; sterile base nearly absent; mature gleba

greenish yellow C. maxima (p. 62)

Plants rather small; subgleba nearly absent; mature gleba dark purplish; spores nearly smooth

C. fragilis (p. 64)
Plants large or moderately small; sterile subgleba large and persistent

Spore powder dark purplish; plants large; spores strongly warted C. cyathiformis (p. 63)

Spore powder brown; plants rather small; spores minutely warted C. data (p. 65)

Spore powder dull or bright olivaceous yellow; spores smooth or nearly so

Capillitium threads about as thick as the spores and with circular holes in them; spores

very small C. craniformis (p. 67)

Capillitium threads much thicker than the spores and with long linear pits; spores larger

C. caelata (p. 68)

Calvatia rubro-flava (Cragin) Lloyd
C. aurea Lloyd

Plates 33, 34 and 112

Plants subglobose, flattened on top, 2-10 cm. wide by 1.5-5 cm. high, strongly
plicate beneath, usually with a short, abruptly pointed stalk which is centrally attached
by one or more slender root-like strands; usually nearly white when fresh and immature
with a faint tint of pink or lavender but varying to pinkish leather color, all parts
turning at once to brilliant yellow (about chrome yellow) upon rubbing or cutting,
drying orange or yellow orange to orange red or bay, in age becoming a metallic brown.
Peridium about 125-160/x thick when fresh, becoming very thin and naoery on drying,
practically single layered, the outer layer being represented by only a delicate fur-
furescence, becoming smooth and shiny on exposure to weather, cracking up into irreg-
ular areas and falling away after drying. Gleba pure white when young and lined
with a hymenium as in Lycoperdon; the cavities very minute, up to about 0.2 mm.

• Calvatia pachyderma (Peck) Morgan and C. lepidophorum (Ellis) Lloyd are two western species which are sometimes
confused. Both have thick peridia but may easily be distinguished by the following characters: In the former the pale peridium
is smooth or with thin adherent, white patches and there are no separable layers, although the entire peridium is apt to be scissile
like short-cake; spores (in plant from type locality, collected by Pringle and determined by Peck) oval to elliptic, very finely
warted under high power (x 2160), 3.5-4 x 4.2-6/x, with a mucro or short pedicel; capillitium threads fragmented, strongly oliva-
ceous yellow under microscope, irregular in thickness, rather frequently branched, pitted, up to llji thick in swollen places;
mature gleba bright olivaceous. In C. lepidophorum the outer peridium has an extremely thin, glazed surface marked with
inherent patches or rounded "bosses" but this is not separable. Beneath the thick outer peridium, unlike any other American
Calvatia, the gleba is covered by a thin, papery membrane and this is the character which most easily determines the species.
Spores spherical, distinctly warted, 4.2-c3/i. Capillitium threads slender, fairly even, up to 5/i thick, brown, occasionally branched,
and not so sinuous as in C. pachyderma. The plant Morgan described as C. pachyderma is C. lepidophorum, as noted by Lloyd,
and not Peck's species. Lloyd proposed the genus Hypoblema for Ellis's Lycoperdon lepidophorum (Myc. Notes, p. 140) but later
reduced it to a subgenus of Calvatia (Footnote 13 to Index to vol. 2).

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LYCOPERDACEAE 61

long in section, labyrinthiform, becoming larger and more indistinct as they pass into
the subgleba, tie cavities of which are up to about 1 mm. long, empty when young,
but becoming loosely filled with soft, woven threads toward maturity so as to obscure
their outlines. At full maturity this growth has in most cases entirely obliterated
the outlines of the chambers and the subgleba is homogeneous. At times the orig-
inal chamber wall may still be noticed, although the cavity is stuffed. This last con-
dition is shown in one of the plants on pi. 33. Subgleba occupying a comparatively
small part of the peridium, about 3 mm. to 2.5 cm. thick below, depending on the
size of the stalk, but running as a thin layer far up around the sides of the gleba.
Gleba and subgleba changing when cut or wounded from a pure white to a brilliant
yellow, the color change appearing first in the subgleba and slowly spreading over its
surface and then over the surface of the gleba. Gleba changing to honey yellow or
Isabella color when fully ripe, the subgleba more brownish and often with a tint of
chocolate. Glebal chambers usually incompletely breaking down, particularly in the
region just beneath the peridium so that erosion is very slow, the gleba remaining as a
soft, spongy mass like a powder puff for a long time after exposure. Taste pleasant and
odor weak when fresh ; a decidedly strong odor of old ham on drying.

Spores (of No. 7163) nearly yellow ochre, between that and buckthorn brown of
Ridgway, spherical, very minutely rough even under high power (x 2160) with a hyaline
outer membrane and a colored inner one, 3^1.4^ thick, with a short pedicel. Basidia
subglobose, about 6-8/j. thick, with 2-4 sterigmata. Capillitium threads up to 4.4/i
thick, considerably septate, moderately thin-walled and remarkable in the frequent
occurrence of distinct holes, 1-3/x in diameter, in the walls, sinuous, usually breaking
into short pieces when put under a cover glass.

This is certainly C. aurea Lloyd (Myc. Notes, p. 11) and it is probably true as
Lloyd thinks that C. rubro-flava is the same (Myc. Notes, pp. 32, 90, 149). We have
not seen any authentic material of the latter, but the description agrees. The species
differs sharply from all other Calvatias in the single layered peridium and homogeneous
subgleba at maturity. These characters are probably of enough importance to entitle
the plant to a genus of its own. The long persistence of the soft gleba makes these
very clean puffballs to handle, and is in sharp contrast to the easily fragmented and
dirty habit of C. caelata, etc. A homogeneous subgleba at maturity is, so far as we
know, found in no other puffball except C. maxima, C. Candida and Lycoperdon poly-
morphic (L. cepaeforme). The present plant is in fact very closely related to C.
Candida, and Cunningham reduces it to a variety of that species. The striking color
and the remarkable color change of fresh plants when injured lead us to retain it as a
species, at least until more is known of C. Candida in the fresh condition. The latter
is not known to occur in America. A peculiarity of the color change when a fresh young
plant is rubbed is that the bright yellow color persists only for a few minutes, then
fades away to the original tint.

The species seems confined in our country to cultivated soil, and this suggests that
it is an introduction, probably from South America. It is reported by Lloyd from
Alabama, Missouri, Ohio, Indiana, Washington, D. C., and Connecticut, and we are
adding several other stations. Cragin's plant was found in Kansas. Our record is the
first for this state. It is also reported from Brazil, Bolivia, Australia, and Japan.

Illustrations: Lloyd. Myc. Illustrations, No. 22 (asC. aurea).

7107. On a lawn, Sept. 2, 1923. Spores minutely rough, 3.5^4.4^.

7154. Under an arbor among flowers, The Rocks, Sept. 26, 1923. Spores 3-4.2/j thick.

62 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

7163. On the ground in a corn field, Sept. 27, 1923.

7242. Same location as No. 7163, Feb. 15, 1924. This winter collection was less red than the others.

Louisiana. (N. Y. B. G. Herb. No data or name) .

Virginia. Blacksburg. Murrill, coll. (N. Y. B. G. Herb.).

New York. New York Botanical Garden, in a dahlia bed. Miss Eaton, coll. (N. Y. B. G. Herb.).

Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).

Missouri. St. Louis. Trelease, coll. (N. Y. B. G. Herb., as Lycoperdon delicahim) . Spores nearly

smooth, 3.5-4^ thick, often with a short pedicel. Capillitium threads slender, 2-3. 7\i thick

with holes in the walls.
Jamaica. Cinchona. On a dry bank. Murrill, coll. (N. Y. B. G. Herb.).
Brazil. Rick, coll. (U. N. C. Herb.). Spores apparently smooth, 3.4-4^, with a mucro.

Calvatia maxima (Schaeff.) Morgan
Lycoperdon giganteum Batsch
Lycoperdon bovista Linn.

Plate 112

We take the following description from Morgan (1. c, p. 166):

"Peridium very large, globose, depressed-globose or obovoid, with a thick cord-like
root. Cortex a flocculose or nearly smooth continuous layer, very thin and fragile,
white or grayish, changing to yellowish, drying up and becoming brown, remaining
closely adherent to the inner peridium or sometimes peeling off in patches ; inner perid-
ium thin and very fragile, after maturity gradually breaking up into fragments and
falling away. Subgleba very shallow or quite obsolete, when present said to be compact
and not cellular; mass of spores and capillitium greenish yellow, then brownish oliva-
ceous; the threads very long, frequently septate, branched, the primary branches much
thicker than the spores, the ultimate ones more slender; spores globose, even or some-
times very minutely warted, 3.5^1.5^ in diameter, often with a minute pedicel."

We have examined a specimen from Madison, Wisconsin, and find the spores to be
smooth, subspherical, 3.5-4ju thick, with a short pedicel and a distinct oil drop; capil-
litium threads easily fragmented, occasionally branched, about 5.5yu thick.

It is surprising that we have not found this huge puffball in Chapel Hill, nor has
Beardslee found it at Asheville. Both Schweinitz and Curtis report it, the former say-
ing that it may reach a breadth of three feet, but his finding it in this state is doubtful
as the Bovista giganlea of his herbarium is Calvatia cyathiformis (also noted by Lloyd,
p. 396). It occurs in meadows and is excellent food. In American Naturalist (18:
530. 1884) Professor C. E. Bessey records a specimen of the giant puffball found by
Professor R. E. Call in Herkimer County, N. Y., in 1877, that was 5 ft. 4 in. x 4 ft.
6 in. x 9| in. This, says Professor Bessey, is by far the largest on record. Buller
(Researches on Fungi, p. 85) has estimated that a large fruit body of this species
(40 x 28 x 20 cm.) contained over seven trillion spores.

The species is said by Dearness to be fairly common in Ontario.

Illustrations: Batsch. Elench. Fung., pi. 29, fig. 165.

Berkeley. Ann. Sci. Nat., 2nd ser., 12: pi. 2, figs. 7-14. 1839.
Boudier. Icon. Myc, pis. 188 and 189 (as Lycoperdon bovista).
Gibson. Our Edible Toadstools and Mushrooms, pi. 34. 1895.
Hard. Mushrooms, pi. 57 and fig. 455.

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Calvatia cyathiformis. No. 7162

LYCOPERDACEAE 63

Hollos. L c, pi. 12, figs. 4-6.

Kriegcr. Xat. Geog. Mag. 37: 415. 1920.

Murrill. Mycologia6: pi. 126. 1914.

Nees von Esenbcck. Syst. Tilzc Schw., pi. 11, fig. 124C.

Patterson and Charles. U. S. Dept. Agric. Bull. 175: pi. 38, fig. 2 (as C. gigantea). 1915; also

Farmers' Bull. No. 796, fig. 18 (as C. gigantea). 1917.
Schaeffer. Fung. Bavar., pi. 191.
Sorokine. Rev. Myc. 1890, pi. 26, fig. 360.
Taylor. Food Products I, fig. 11. 1894.

Wisconsin. (Univ. Wis. Herb, and U. N. C. Herb.) Spores smooth, subspherical, 3.5-4^, with a short
pedicel. Capillitium fragmenting, occasionally branched, frequently pitted, up to 5.5^.

Calvatia cyathiformis (Bosc) Morg.
Lycopcrdon cyathijorme Bosc
Bovista lilacina Berk. & Mont.

Plates 35, 36 and 112

Plants nearly globose or turbinate, 7.5-15 cm. thick, attached below by a short
root; cortex smooth or slightly scaly, the upper part often cracking into areas. Inner
peridium thin and delicate, at maturity scaling away by degrees together with the
cortex to expose the dark purple interior; lower, sterile part (subgleba) remaining intact
as a persistent, dark, cup-like base which may remain in place over winter. Gleba at
first white, then changing through yellow to deep purple brown as the plant matures.

Spores (of No. 550) globose, with numerous distinct spines, 4.4-6. 5ju thick, ex-
cluding the spines. Capillitium threads up to 4.3/i thick, very long, sparingly branched,
interwoven, considerably septate and breaking into sections at the septa, walls with
minute pits which are not easily visible except under high power.

The species is common in Chapel Hill from July to late autumn in open places,
uncultivated fields, lawns and orchards. When it is young and the inside white it is
excellent food when fried like an egg plant.

Some interesting facts concerning this Calvatia have been brought out in a study by
Shantz and Piemeisel (Journ. Agric. Research 11, No. 5: 191. 1917). They find that
the so-called "fairy rings" caused by this fungus advance about 24 cm. a year. One
ring has been found which is estimated to be 420 years old. The grass is stimulated
just behind the fruiting bodies of the ring, and forms a deeply colored, luxuriant growth,
whereas it is dry, often dead over the mycelium of Agaricus tabularia and Marasmius
oreades and shows a luxuriant growth on both sides. When disturbed, as by being
plowed up, the rings cease to fruit, and cannot be detected in vegetation, while in the
closely related species, C. polygonia, they continue to grow and fruit, as is also the case
with Agaricus tabularia.

Cunningham considers C. novae-zelandiae Lev. an additional synonym for this
species (Trans. N. Z. Inst. 57: 191. 1926).

Illustrations: Cunningham. Trans. N. Z. Inst. 57: pi. 2, fig. 3 (as C. lilacina). 1926.
Hard. Mushrooms, pi. 58 and fig. 457 (as C. lilacina).
Hollos. 1. c, pi. 13, pi. 14, figs. 1-5, pi. 29, fig. 7.
Krieger. Xat. Geog. Mag. 37: 416. 1920.
Lloyd. The Lycoperdaceae of Australia, etc., pi. 35, fig. 1.
Murrill. Mycologia 1: pi. 15, fig. 1. 1909.
Sweetzer. Univ. Oregon Leaflet, Bot. Ser. No. 9, fig. 3.

64 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

550. On campus, Oct. 10, 1912.

556. On a lawn, May 18, 1912. Spores distinctly warted but still immature, 4-5.5/u thick.
672. Among broomsedge on a hillside, Sept. 18, 1908.
3229. In a pasture, Feb. 25, 1919. Plant practically gone except for the sterile base. Spores dis-
tinctly spiny, 4.4-6ji.
6044. In open, grassy place, Jan. 10, 1923. Plant with no sterile base. Spores with distinct warts,
4.8-6.2m thick.

Asheville. Beardslee.

Florida. Weber, coll. (U. N. C. Herb.).

Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb.).

Kentucky. Crittenden. Lloyd, coll. (N. Y. B. G. Herb.).

Virginia. Blacksburg. Murrill, coll. (N. Y. B. G. Herb.).

District of Columbia. Cook, coll. (U. S. Nat'l. Herb.).

New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb.).

New York. Peck, coll. (N. Y. B. G. Herb.).

Wisconsin. Vilas Woods. (U. N. C. Herb.) Spores apparently immature, some distinctly warted

others less so, 4.4-4.8j», often up to 6.8p.
South Dakota. Huron. Miss Crouch, coll. (N. Y. B. G. Herb.). Spores distinctly warted, 4.5-6.2/i

thick, sometimes pedicellate. Capillitium threads up to 5.5ju thick, sometimes irregular and

up to 10m thick in swollen places, fragmented with ends often closed.
Canada. Melbourne, London, Vauxhall Bridge. Dearness, coll. (Dearness Herb.). Said by Dearness

to be common in Ontario.

Calvatia fragilis (Vitt.) Morgan

Plates 37 and 112

Fruit body obovate, up to 5.5 cm. thick (in our collections), the base abruptly
contracted to a blunt point; cortex thin, separable, brownish to straw color, the surface
minutely felted to smooth and usually areolated or pitted (in a form quite smooth),
the larger depressions usually with a central flat, purplish brown scale with the edges
more or less free; inner peridium deep purplish brown in section, about 0.7 mm. thick,
brittle and breaking away by degrees in irregular fragments from the top downward.
Gleba at maturity deep purplish brown, soft but not very powdery and not falling out
easily as in C. caelata. Subgleba often almost absent, at times more obvious when the
base is pointed, brown, the cavities very small and obscure.

Spores (of No. 4783) dark brownish purple, spherical, smooth or minutely warted,
3.7-4.6yu, rarely 5.4ju thick. Capillitium of short segments, irregular, sinuous, 3.5-6/x
thick, pits very minute and not easily visible except under high power (just like the
capillitium of C. cyathiformis).

This species is distinguished from C. cyathiformis by its small base with very little
subgleba, the smaller size of the plants, and the nearly smooth spores which are of
smaller average size. Calvatia caelata is of course separated by its gleba color and by the
much thicker and peculiarly pitted capilhtium. The surface appearance of the typical
form of the present species is almost exactly like Lloyd's figs. 2 and 4 and, in part, fig.
3 of plate 36 of C. caelata and form, but that is, of course, excluded by the purplish
spores of our plants.

Calvatia pachyderma differs distinctly in the olive brown or buffy brown gleba,
homogeneous but scissile peridium of uniform color throughout and without a
sharply contrasting cortex, and in the average larger size of the capillitium.

PLATE 37

Calvatta fracilis. Urbana, Illinois [above] ; London. Can. [below]

PLATE 38

Calvatia elata. Connecticut

LY( OPERDACEAE 65

Our description is from the Canadian plants listed below. A plant from Colorado
is quite similar to these, except that the areolations above are very pronounced, so that
the tendency is for each areolation to break away separately, bringing its part of the
inner peridium with it. The smooth form mentioned above is represented by plants
from Kansas and Nebraska entered below. The peridium is quite smooth, tan to
brownish tan; all forms are otherwise the same.

Illinois. Urbana. McDougall, coll. (U. N. C. Herb.). Spores minutely waited, 3.5-4.5ju.
Kansas. Rooks Co. On open prairie. Bartholomew, coll. (U. N. C. Herb.).

Also a Kansas plant from Cragin, No. 522. (N. Y. B. G. Herb, and U. N. C. Herb.) Spores

minutely rough, 4.5-5.8/1.
Nebraska. Long Pine. Bates, coll. No. 486. (N. Y. B. G. Herb, and U. N. C. Herb.) Spores

smooth with a mucro, 4-5.5/1. Capillitium threads sinuous, 2-4/i thick.
Colorado. Bethel, coll. (N. Y. B. G. Herb., as C. pachyderma, and U. N. C. Herb.). Spores purple,

minutely warted, 4.5-6/1 thick. Capillitium threads about 5m thick, in fragments with ends

usually closed.
Wyoming. Pitchfork. Davis, coll. (U. N. C. Herb, from N. Y. B. G. Herb., labelled C. lilacina var.

occidentalis). Spores nearly smooth, 4-5.5/1, rarely up to 7/x, no pedicel.
Canada. London. Deamess, No. 4783 Font. (Dearness Herb, and U. N. C. Herb.) Also another

collection by Dearness from London. (N. Y. B. G. Herb, and U. N. C. Herb.) Spores nearly

smooth, 3.7-4.5/1 thick.

Calvatia elata (Massee) Morgan

Plates 26, 38 and 112

Plants small for this genus, with a distinct, usually long, subcylindrical, often
pitted stalk up to 4 cm. thick and 8.5 cm. long (rarely longer) and a subglobose head
about 3-6 cm. thick; color when mature pale brown to leather color; cortex only a very
thin granular or powdery, persistent layer that is almost absent on the stalk below;
inner peridium thin and at maturity very fragile, soon cracking up into fragments and
falling away or adhering for awhile to the capillitium. Gleba brown; the subgleba of
the stalk with distinct empty chambers of fair size.

Spores (of plant from Litchfield, Conn.) spherical, very minutely warted, short-
pedicellate, 3.5-4.4/z. Capillitium threads much fragmented, moderately pitted, up
to 5m thick.

The species was described by Massee as Lycoperdon datum from New England
plants in the Berkeley Herbarium, where it was included as L. saccatum. We have
seen a good example of C. saccata from England (N. Y. Bot. Garden, from Cooke)
and find that the spores are larger and rougher than in our C. elata (distinctly warted,
4.5-5.5/0, thus confirming Morgan's statement to that effect. As in so many other
cases, these differences are hardly more than in regional forms of the same species.
Hollos considers C. elata a variety of C. saccata.

The two Canadian plants entered below are like each other in every particular
except that the capillitium in one plant was unbranched, in the other freely branched.
In both plants the capillitium was less fragmented than in other specimens we have
seen. One of the plants had previously been sent to Lloyd and determined by him
as C. saccala.

This species is in the cranijormis group, and seems nearest that species. It
differs in smaller size, a proportionately longer stem, darker color of mature gleba,
and in larger and minutely roughened spores. It seems also near L. muscorum and in
certain respects resembles L. gemmation. See these species for further discussion.

66 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustrations: Hollos. I. c, pi. 17, figs. 3-5.

Lloyd. Genera of Gasteromycetes, pi. 10, fig. 48.

Massee. Journ. Roy. Mic. Soc, 1887, pi. 13, figs. 13-15.

Morgan. Journ. Cin. Soc. Nat. Hist. 12: pi. 16, fig. 7.
Illustrations of C. saccata: Hollos. 1. c, pi. 16.

Krombholz. Abbild., pi. 30, figs. 11,12.

New York. Peck, coll. (N. Y. B. G. Herb., as L. saccatum).

Connecticut. Litchfield. Miss White, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.).

Massachusetts. Waltham. Reported by Lloyd in Genera of Gasteromycetes, p. 12.

Ontario. Byron. Dearness, coll. Two plants. No. 2843 H. (U. N. C. Herb.). Spores minutely
warted, short-pedicellate, 3.7-5^. Capillitium threads not breaking up into short pieces,
pitted, sinuous in places, up to 9m thick. No. 4825/2843. (Dearness Herb.).

Calvatia cretacea (Berk.) Lloyd
Lycoperdon cretaceum Berk.
C. borealis Th. Fries

Plate 112

Peridium subglobose to flattened, plicate below, the base pinched to a small point
which is lightly attached by white mycelium, 2.5-5 cm. broad; cortex conspicuous, white
then buffy to ochraceous, composed of a continuous coat which consists above of thick
compound warts with connivent tips, becoming smoother downward and fading to a
mealy furfurescence below, or in some cases much smoother above and resembling the
smoother forms of Scleroderma aurantium; in all cases cracking into irregular areas or
groups of warts much as in Lycoperdon marginatum and falling away by degrees to expose
the thin, shining, silvery brown to chocolate colored inner peridium which falls away in
flakes from above downward. Gleba passing through golden olive to chocolate brown,
the subgleba small, chocolate brown or the basal part white, the chambers obvious,
but not large.

Spores (of plant from Lapland) spherical, greenish yellow under the microscope,
minutely but distinctly warted, the warts embedded in a hyaline coat, 4.2-5.5/^, rarely
6.8ju, with one strong oil drop and sometimes a mucro. Capillitium threads fragmented,
color of spores, often wavy, rarely branched, much pitted, 3-7ju thick.

This arctic species is strongly characterized by its thick, deciduous cortex and
purplish brown gleba. It was first described from Bellot Island, and has since been
reported from Lapland by Th. C. E. Fries (as C. borealis; Zur Kenntnis der Gastero-
myceten flora in Torne Lappmark, p. 238. 1914) and from Herschel Island, Kay
Point, Mackenzie River Delta, and Bernard Harbor by Mr. John Dearness (Rept.
Canadian Arctic Exp. 1913-18, 4, Part C: 17. 1923). We have two good collections
from Lapland made by us in 1921, and these show the variation in the cortex mentioned
above. Lloyd (Myc. Notes, p. 650) suggests that C. arctica Ferdinandsen and Winge
(Meddelelser om Groenland 43: 144, pi. 9. 1910; see Saccardo, Sylloge 21: 480)
may be the same and in this he is followed by Fries (Arkiv f. Botanik 17, No. 9: 23.
1921). We think it very probable that this supposition will finally prove to be correct.
The illustrations of C. arctica show stouter warts than are recorded for C. cretacea
(up to 8 mm. thick at least in one plant) and little or no chocolate color is shown in the
gleba. However, in the discussion Ferdinandsen says the gleba is "grayish chocolate-
brown." The larger warts may easily be a variation within the species. The long
sterigmata described in C. arctica are a remarkable peculiarity, but this does not

>

LYCOPERDACEAE 67

exclude C. crelacca, as its basidia have not been studied. Our material does not show
them.

Illustrations: Fries. As cited above.
Lloyd. Myc. Notes, p. 650, fig. 929.

Canadian Arctic. Bernard Harbor. Johansen, coll. (Dearness Herb, and U. N. C. Herb.) Spores

distinctly warted, 4.2-6.5m- Capillitium threads up to 1 l/i thick.
Swedish Lapland. Abisco. Coker, coll. (U. N. C. Herb.). One of these two collections had spores

very minutely warted, less so than in the other, apiculate, spherical to subsphcrical, 4.5-6ju.

Capillitium threads up to 7.5m thick.

Calvatia craniformis (Schw.) Fr.

Lycoperdon delicatum B. & C. (Not L. delicatum Berk.)
L. Missouriense Trelease

Plates 39 and 112

Plant large, up to 9 cm. thick, obovoid or top-shaped, the thick, stalk-like base
rounded below and attached by fibrous strands; cortex very thin, papery, smooth,
nearly glabrous to delicately furfuraceous, pale tan or grayish in color, at maturity
scaling off in areas; inner peridium also thin and fragile, yellowish to reddish, the upper
part cracking into plates and scaling off after the cortex peels. Subgleba occupying
the stalk-like base, of honeycomb structure and quite spongy, concave above and long
persistent. Gleba white, then greenish yellow or soiled ochraceous in color.

Spores (of No. 538) spherical, smooth, with a distinct mucro or short pedicel,
2.8-3.9/z thick. Capillitium slightly branched, sparingly septate, walls with numerous
pits, possibly holes in them which are 1-3^ in diameter, easily breaking up, usually at
these pits; average thickness of main threads 3.5ju.

Unlike C. cyathiformis, this species inhabits wooded places or shrubby borders of
paths or walls, and is much less abundant, though not at all rare. It, too, is a palatable
and valued food. It is easily distinguished by the greenish yellow rather than purple
spore mass. The type in the Schweinitz Herbarium is a fine example of the species.
The spores are smooth, 3.2-3.6/* thick.

In referring to C. craniformis on p. 100, Hollos is confused. The L. delicatum
that Lloyd says is the same as C. craniformis is evidently from the context the delicatum
of Berkeley and Curtis from Pennsylvania, not of Berkeley from India, etc. (L. Berkeleyi
De Toni). In his reference to L. delicatum on p. 153 of Myc. Notes, Lloyd makes the
double error of confusing Morgan's treatment of what he took to be L. delicatum Berk,
with the B. & C. species and further of accusing Morgan of having no conception of
L. delicatum B. & C, when Morgan stated thirteen years earlier (Journ. Cin. Soc. of
Nat. Hist. 12: 171. 1890) the same thing that Lloyd here announces in italics, i.e.,
that L. delicatum B. & C. is the same as C. craniformis. It seems well established that
L. Missouriense Trelease (see Sacc. Syll. 9: p. 277) is also the same as C. craniformis,
(Morgan, 1. a). For explanation of Cooke's mistake in thinking it C. lilacina see
Lloyd, Myc. Notes, p. 308.

Illustrations: Hard. Mushrooms, pi. 60 and fig. 461.
Krieger. Xat. Geog. Mag. 37: 419. 1920.
Marshall. Mushroom Book, pi. opposite p. 126.

68 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

538. Near rock wall in Episcopal churchyard, October 9, 1912.
676. Battle's Park, just east of campus, September 19, 1908.
1535. In woods, Battle's Park, summer of 1915. Spores smooth, 3-3.8^.

Alabama. (Curtis Herb.)

Auburn. Earle, coll. (N. Y. B. G. Herb.).
Texas. Long, coll. (N. Y. B. G. Herb.).
Virginia. Blacksburg. Murrill, coll. (N. Y. B. G. Herb.).
Maryland. Agric. College. James, coll. (U. S. Nat'l. Herb.).
Delaware. Mt. Cuba. Commons, coll. (N. Y. B. G. Herb.).
Pennsylvania. (Curtis Herb.)

Also Gentry, coll. (N. Y. B. G. Herb.).
New Jersey. Ballou, coll. (N. Y. B. G. Herb.) .
New York. (Curtis Herb.)
Wisconsin. (Curtis Herb.)
Ohio. Morgan, coll. (N. Y. B. G. Herb.).
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.). Spores smooth, 3. 2-3.8/i.

Calvatia caelata (Bull.) Morg.

Plates 40 and 112

Plant large to very large, sterile base ample, either long and cylindrical or shorter
and tapering to the pointed base; cortex a flocculent layer which is thicker above and
there usually areolated into more or less prominent warts, thinner downward, but with a
tendency throughout to form more or less stellate, flattish areas; color whitish, then
pallid yellowish to brownish. Inner peridium thin, breaking up into fragments at
the top, then downward, exposing the bright olivaceous gold to brownish olivaceous
gleba, which is very fragile and powdery. Sterile base persistent, up to 9.5 cm. thick
and 10 cm. high, furrowed below and pinched to a point; the chambers of the subgleba
distinct, empty and moderately large, extending up the sides for several centimeters
as a tapering margin to the cup.

Spores (of the plant from Pawling, New York) spherical, smooth, with a short
mucro and distinct oil drop, 3.8-4.5ix. Capillitium threads up to 17/* thick, usually
7— 11/i, somewhat branched, easily breaking up at maturity; walls with narrow (about
lfi wide), linear, sinuous pits.

This is a northern and middle western species and so far as we know has not been
collected in the southeastern states. Specimens we have seen are from New York and
Canada. Curtis reports Lye. caelatum from this state (Cat'l., p. 110), but there is no
reason to think that his plant was correctly determined. Calvatia caelata is not rep-
resented in his herbarium, but C. craniformis, which he does not report, is found there
under the name of L. delicatum B. & C. The species is well marked and very easily
recognized by the fibrous or flocculent covering, separated into warts or stellate areas,
and by the very characteristic thick capillitium with its conspicuous, linear pits. For
treatment of this species see Morgan, Journ. Cin. Soc. Nat. Hist. 12: 169; also Lloyd,
Myc. Notes, p. 166 and Lycoperd. Austr., p. 35. Lloyd's magnified figure of the
cortex is good.

The plant is most like C. craniformis in size, shape, and color of gleba, and in the
northern states it seems largely to take the place of that species; in the south C. crani-
formis is very common, while in the north it is certainly rare.

Cunningham gives the following list of synonyms for this species: L. Fontanesii
Dur. et Mont.; L.favosum Bon.; L. Sinclair ii Berk.; C. favosa (Bon.) Lloyd. (Trans.
N. Z. Inst. 57: 190. 1926.)

PLATE 40

Calvatia caelata. Pawling, N. Y.

LYCOPERDACEAE 69

For cytological information by Maire covering this and other species, see under
Scleroderma ourantittm.

Illustrations: Bulliard. Herb. Fr., pi. 430.

Cunningham. Trans. N. Z. Inst. 57: pi. 1, fig. 1. 1926.

Hollos. 1. c, pi. 15, tigs. 1-12.

Krombholz. Abbild., pi. 30, figs. 7-10.

Lloyd. Myc. Works, pi. 36.

Lloyd. Lycopcrdaceae of Australia, etc., fig. 39.

Lloyd. Photogravure of Am. Fungi, No. 22.

Schaeffer. Fung. Bavar., pi. 189.

New York. Pawling. Dr. H. S. Robinson, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.).

Wisconsin. Sheboygan. Brown, coll. (N. Y. Bot. Gard. Herb.).

Dakota. Huron. Miss Crouch, coll. (N. Y. Bot. Gard. Herb.).

Colorada. Denver. Bethel, coll. (N. Y. B. G. Herb.).

Canada. Ontario. Ottawa Valley. Dearness, coll. (Dearness Herb., No. 4783 od'l).

LYCOPERDON Tourn.

Plants comparatively small, globose, obovoid, top-shaped or pyriform; the base in
nearly all of our species filled with a sterile, honeycomb tissue (subgleba); cortex some-
times smooth but usually composed of a coat of spines, scales, warts, or scurf, the spines
frequently converging at their apices to form stellate groups. Inner peridium thin,
papery and flaccid, opening by a definite apical mouth and collapsing more or less as the
spores escape. Gleba composed of minute chambers which are lined with the hyme-
nium, white when young, changing as the plant ripens through yellowish or olive to
brown or purplish. Capillitium well developed, composed of long, branched or un-
branched interwoven threads which are continuous with the inner peridium and the
subgleba. From the center of the subgleba the threads may extend upward in such a
thick fascicle as to form a more or less conspicuous column called the columella. Sub-
gleba in most species obvious and "cellular" in structure, that is, composed of small or
rather large empty chambers. Spores globose or rarely elliptical, commonly warted or
spiny, in most species with a short or long pedicel. Basidia short and plump with four
slender, apical sterigmata that are usually of unequal length (2-3-spored forms are said
to occur).

In L. polymorphum, the subgleba is not cellular but formed of homogeneous tissue.
In L. pusillum it is lacking. In some species a distinct homogeneous diaphragm very
similar in structure to the inner peridium separates the subgleba from the fertile tissue;
in other cases the two may merge imperceptibly into each other. For the develop-
ment of Lycoperdon depression in the diaphragm group, see Rabinowitsch (I.e.); for
the group without diaphragm, see Bonorden. In his paper on the development of L.
depressum (1. c.) Cunningham says:

"About the time of the commencement of the breaking-up of the tramal plates,
hyphae from the endoperidium (of the capillitium type, being thick-walled and spar-
ingly branched) grow out from the endoperidium into the cavity of the gleba, following
for the most part the inner portions of the glebal plates. They develop in large
numbers, so that soon the whole cavity of the gleba becomes filled with them."

Though he does not say so in so many words, it is evident that he means that these
are the capillitium threads and that such is their origin. So far as we know this has
not been brought out before. As to the cytology, he finds that the cells of the fruit body

70 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

are probably binucleate, that there is a nuclear fusion followed by two divisions in the
basidium, one going into each of the spores and there dividing, so that the mature spores
are binucleate.

A genus of numerous species of small plants that grow on earth or rotten wood, or
rarely on moss or the bark of living trees. Unless found in large groups, which often
happens with L. pyriforme, the plants are too small to be of much value as food. In
addition to the species treated, L. muscorum, which Hollos thinks is the same as L.
pseudoradicans Lloyd, should be found here in dense moss beds, as Lloyd's plant has
been collected in Virginia. In a group by itself is the very remarkable Lycoperdon
sculptum found by Harkness in the mountains of California. It is covered with massive
pyramidal warts up to 4 cm. broad and 3 cm. high (see Bull. Cal. Acad. Sci. 3 : 160,
pi. 1. 1885).

Literature

Bambeke. De la valeur de l'epispore pour determination et Ie groupement des especes du genre Ly-
coperdon. Bull. Soc. Myc. Fr. 22: 1. 1906.

Bonorden. Die Gattungen Bovista, Lycoperdon u. ihr Bau. Bot. Zeit. IS: 593, 609, 625. 1857.

Cunningham. Development of Lycoperdon depressum. New Zealand Journ. Sci. and Tech. 8: 228,
figs. 1-7. 1926.

Cunningham. Lycoperdaceae of New Zealand. Trans. N. Z. Inst. 57: 187. 1926.

Cunningham. The Genus Lycoperdon. Proc. Linn. Soc. N. S. W. 51 : 627. 1926.

Hollos. I.e., p. 90.

Lloyd. The Genus Lycoperdon in Europe. Myc. Notes No. 19. 1905.

Lloyd. The Lycoperdons of the United States. Myc. Notes No. 20. 1905.

Massee. Monograph of the Genus Lycoperdon. Journ. Roy. Mic. Soc, p. 701, pis. 12, 13. 1887.

Massee. 1. c, p. 66.

Morgan. Journ. Cin. Soc. Nat. Hist. 14: 5. 1891.

Peck. New York Species of Lycoperdon. Rept. N. Y. St. Mus. 32: 58. 1879.

Peck. United States Species of Lycoperdon. Trans. Albany Inst. 9: 285. 1879.

Petri. Flora Italica Cryptogama (Gasterales), Fasc. 5, p. 34. 1909.

Rabinowitsch. Beitrage zur Entwickelungsgeschichte der Fruchtkorper einiger Gastromyceten.
Flora 79: 385, pis. 10 and 11. 1894.

Trelease. Morels and Puffballs of Madison, Wis. Trans. Wis. Acad. Sci., Arts and Letters 7: 105.
1888.

Tulasne, L. R. and Ch. De la Fructification des Scleroderma comparee a celle des Lycoperdon et des
Bovista. Ann. Sci. Nat., 2nd. ser., 17: 5, pis. 1, 2. 1842.

Key to the Species

Mature spore-mass with a purplish tint (grayish purple or purplish brown)

Cortex composed of very slender hairlike spicules among granules L. atropiirpnreum (p. 71)

Cortex composed of very long spines convergent in groups at their apices

Spines pale at maturity and, on falling off leaving the inner peridium smooth and unreticulated

L. pulcherrimum (p. 72)
Spines dark brown at maturity and, on falling off leaving the peridium reticulated with minute

granules L. echinatum (p. 73)

Cortex a smooth, continuous layer which breaks up into very thin scales . . . .L. rimulatum (p. 75)
Cortex thick with spongy patches on a granular membrane which peels off at maturity

L. subvelatum (p. 76)
Cortex a scurfy or granular, persistent coat, at times mixed with very fine spines

L. umbrinum (p. 76)
Mature spore-mass olive-brown

Growing on the bark of living trees or rarely on logs; plant very small L. acuminatum (p. 78)

LYCOPERDACEAE 71

Growing on dead wood (rarely on carthl

Flants pyriform, usually crowded, surface nearly smooth L. pyriforme (p. 79)

Plants subglobose, rarely touching; the surface of the pcridium becoming marked with distinct

pits like a thimble L. subincarnatum (p. 80)

Growing among mosses in meadows and pastures

Plants distinctly stalked, cortex of minute granular warts and particles. ,L. muscorum (p. 81)
Growing on the ground or rotting leaves (rarely on rotten wood)

Plants of medium size, rarely below 2 cm. thick; subgleba usually well developed; plants not
crowded
Subgleba at maturity apparently homogeneous, the minute chambers invisible except

with a lens L. polymorphum (p. 92)

and L. coloratum (p. 93)
Subgleba with obvious chambers

Cortex of stout, terete, separate spines, mingled with shorter wart-like ones; surface

reticulated when larger spines fall L. gemmation (p. 82)

Cortex of pyramidal spines up to 1.5 mm. long which are arranged in groups with the
tips united, and which in our form are lavender when fresh; spores with-
out a long pedicel, 3.7-4.4/j thick L. Peckii (p. 84)

Cortex of very slender spines which are single or grouped and convergent at the

apex, and which fall away with age and leave a smooth, shining surface to

the inner peridium

Spines up to 1 mm. long, rather pale; spores with a long persistent pedicel,

4-5 times as long as the diameter of the spore ... X. pediccllalum (p. 85)

Spines up to 1 mm. long, brown to deep brown at maturity; spores elliptic,

4.3-5 x 5-6.5^ L. eximium (p. 86)

Spines up to 0.5 mm. long, deep brown to black at maturity; easily falling off;

spores spherical, 4-4.8*i thick L. fuscum (p. 87)

Cortex a dense coat of short, sharp spines which largely cohere in groups at their
tips, the soft tissue supporting them breaking up at maturity into large
flakes which fall away above, exposing the inner peridium, which is
dull and furfuraceous with minute scales until weathered

L. marginatum (p. 87)
Cortex as above, but each fascicle of spines falling off singly or in small groups from

above downward L. marginatum, form of woods (p. 87)

Cortex of minute spinules and granules or scurfy scales which slowly wear away

L. umbrinum (p. 76)
Plants small, rarely up to 2 cm. thick; subgleba scanty or almost absent

Cortex of rather persistent, crowded spines; plants often crowded L. Curtisii (p. 89)

Cortex a thin, whitish, scurfy coat, remaining as persistent scales or minute warts upon
the pale brown surface of the inner peridium

Spores short-elliptic to subglobose, smooth L. oblongisporum (p. 91)

Spores globose, minutely asperulate or varying to smooth L. pusittum (p. 91)

Mature spore-mass brown or gray-brown, etc., with no distinct tint of purple or olive (this includes
certain specimens of species included in the other two sections) .

Growing on the ground; cortex of minute, soft granules and spicules L. umbrinum (p. 76)

Growing on wood (very rarely on earth); cortex of minute, harsh warts L. pyriforme (p. 79)

Lycoperdon atropurpureum Vitt.

Plates 42 and 112

Plants up to 3.5 cm. in diameter, depressed-globose or nearly pyriform, plicate
below in large squat individuals, and often with a distinct, pointed root. Subgleba
large, occupying a third or sometimes nearly a half of the body. Surface light tan,

72 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

turning brownish tan, covered with very delicate, hairlike spicules and granules with the
spicule tips usually free from each other. These are quite persistent but in old plants
are partly worn away at the top, thus exposing the light brownish gray shining surface
below. When the plants are exposed to the weather after maturity the granules and
spicules may be worn away over the entire plant, leaving it perfectly smooth, as in our
No. 3230. Pore apical and small.

Spores (of No. 498) deep smoky purple when fully mature (but apparently going
through an olivaceous stage which may be found in herbaria, with the possibility of
confusion as to the real spore color), spherical, 4.4-5.5// thick (counting spines) , covered
with small blunt spines about 1m long, pedicellate, the greater length of the pedicels
being broken off and intermixed with the spores, the part remaining attached to a spore
about half as long as the diameter of the spore. Capillitium composed of long, brown,
unpitted, branched fibers attenuated at the ends and about 5-7. 5/u thick in the larger
parts.

The typical form of this species as described above seems to run into forms ap-
proaching L. echinatum with longer, stellate spines, and Peck treated the latter as var
stellate of atropurpureum.

Illustrations: Lloyd. Myc. Works, pi. 42; pi. 57, figs. 1-6; pi. 123, figs. 7-12.
Peck. Bull. N. Y. St. Mus. 150: pi. 121, figs. 6-10. 1910.
Petri. 1. c, figs. 19, 20.

Quelet. Champ. Jura Vosg. 3: pi. 2, fig. 10. 1875.
Vittadini. Monog. Lycoperd., pi. 2, fig. 6. 1842.

498. Near Battle's Branch, October 4, 1912.

934. Under cedars east of campus, Oct. 17, 1913. Spores 3.7-4.8/t thick, not counting spines which

are up to 1.5// long.
1019. Among leaves in woods, Nov. 26, 1913. Spores 4.4-6// thick, covered with blunt spines which

are surrounded by a hyaline substance which gives the appearance of a halo.
7143. On ground in mixed woods, Sept. 26, 1923. Spores densely covered with blunt spines, 4.4-5.5//

thick.
Also Nos. 498a, 1005, 3230, 7147, 7430.

Asheville. Beardslee, coll.

Blowing Rock. Coker, coll. (U. N. C. Herb.).

Nantahala. Alma Holland, coll. (U. N. C. Herb.). Spores 3.8-5.5//.

Wisconsin. (Univ. Wis. Herb, and U. N. C. Herb.) Spores rather strongly warted and with hyaline
material between the warts, 5.5-6.5//, with many long pedicels broken off and mixed with the
spores.

Canada. London. Dearness, coll. (Dearness Herb.). Spores distinctly warted, often mucronate,
3.7-5//.
Ottawa. Macoun, coll. (N. Y. B. G. Herb.).

LycoperdonpulcherrimumB. &C.
L. Frostii Pk.

Plates 43, 44 and 112

Plant 1.8-4.5 cm. broad and about the same height, tapering below to a slender
root, covered at first with very long, slender, pure white then pale tan spines, which are
united into groups by their tips and which finally fall away from the upper part of the
plant, leaving the smooth shining deep brown, purplish brown or silvery brown surface
of the inner peridium exposed and not reticulated. Gleba white then passing through

PLATE 41

Rhopalogaster transversarium. Alabama [upper left]
Lycopf.rdox SUBVBLATUM. No. 7263. Florida [upper right]
Lycoperdox rimllatlm. No. 7264. Florida [below]

PLATE 42

Lvcoperdon fuscum, cupricum form. No. 5719 [above]
Lycoperdon atropurpureum. No. 498 [below]

LYCOPERDACEAE 73

olivaceous to dark purple-brown; subgleba occupying about one-third or slightly more
of the plant. Odor when maturing distinctly aromatic, somewhat like that of L.
marginatum, but not nearly so offensive.

Spores (of No. 477) with a large oil drop and no obvious stalk, globose, set with
short blunt spines. 4-4.8/1 thick, not counting the spines, average about 5/x with spines.
Capillitium much branched, long and tapering, the lesser branches about the size of the
spores, the main branches up to 8.6/u in diameter in places.

This is a beautiful species and notable for its very long white spines. While hardly
rare, we never find it in abundance, and often only a single specimen at a time. For a
comparison with L. cch'niatum see that species.

Illustrations: Hard. Mushrooms, figs. 463 and 464.
Hollos. 1. c, pi. 17, figs. 15-17.
Lloyd. Myc. Notes No. 20: pi. 55. 1905.
Lloyd. Photogravure of Am. Fungi, No. 19.
Morgan. Joum. Cin. Soc. Nat. Hist. 14: pi. 1, fig. 3.

477. At edge of woods, October 2, 1912.

979. Under cedars in woods, October 21, 1913. Spores covered with blunt, colorless spines which are

less than 1m long, 3. 7-5. 5m thick, omitting spines.
1683. On soil under white oak tree, September 14, 1915.

7560. On campus under a tree, Oct. 8, 1924. Spores minutely but distinctly warted, 4-5m, usually
without a pedicel. Capillitium threads irregular, frequently branched, pitted, about 4m thick.

Asheville. Beardslee.

Maryland. Agric. College. James, coll. (U. S. Nat'l. Herb.).
Pennsylvania. Bethlehem. (Schw. Herb., No. 2259, as L. echinalum.)
Vermont. Brattleboro. Frost, coll. (N. Y. B. G. Herb., as L. Frostii).
Ohio. Morgan, coll. (N. Y. B. G. Herb., as L. Froslii).
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).
Kansas. Stockton. Bartholomew, coll. (N. Y. B. G. Herb.).

Lycoperdon echinatum Pers.
L. constellatum Fr.

Plates 45 and 112

Plants about 2.5-4 cm. broad not counting the spines, usually flattened, the base
pinched to a slender root; cortex of long (2-5 mm.) fascicled spines, with the tapering
points approximated; when young creamy white, in age deep brown and falling off,
leaving the shining, light or dark brown inner peridium reticulated with minute scurfy
particles. Gleba purplish brown to dull brown; subgleba small, with very small
chambers, brown to golden. Odor in drying like that of old ham.

Spores (of No. 7421) spherical, distinctly and rather regularly warted, with hyaline
material between the warts, 4.8-6/*, often with long pedicels attached but these are
usually broken off. Capillitium threads up to 7/t thick, rarely or rather frequently
branched, set with rather numerous knobs, pitted, slightly sinuous and irregular.

The species seems to be confined to deciduous woods and is apparently rare in this
section. It was not reported from North Carolina by Schweinitz or Curtis. In
addition to our Chapel Hill plants, we have examined plants from Asheville, N. C,
and from Wisconsin, and plants from Scarboro, England (New York Botanical Garden),

74 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

also plants from Watlington, England, from Denmark, and from France. These last
three were kindly furnished by Kew Gardens, Romeh, and Patouillard, respectively.

The spines vary considerably in length and delicacy : in our No. 7421 they are much
longer than in the Asheville plants or in any others we have seen. The spines of the
European plants are much shorter than in the Chapel Hill ones, and are stouter than
in any American specimen met with. Lloyd also notes the more slender spines of the
American form (Myc. Notes, p. 208). When the spines of our plants were fresh they
also were stout below, but in drying they shrank greatly. The spores of the plant
from Scarboro (Massee, coll.) are distinctly warted, 4.2-5. &fi; those of the plant from
Montmorency, France, 4.2-6. 2/j, with a distinct oil drop; those of the Asheville plants,
4.2-5. 5/i thick with pedicels usually broken off but sometimes with long ones present.

In regard to L. echinatum, Romell writes that since Fries gives the spores of the
plant from Denmark as "6-7^" thick, he thought it might be different, but on receiving
a Denmark plant from Rostrup he found the spores to be no larger than in other speci-
mens, for example, one from Persoon's herbarium which were 4.5-6/*. These measure-
ments we can confirm for the Rostrup plant, which is the one sent us by Romell.

The gleba color and the spore characters are about the same as in L. pulcherrimum,
but from that the present species is easily separated by the dark brown to blackish
spines at maturity, and the paler color and reticulated surface of the inner peridium
when the spines fall off, the reticulations being due to the minute granular particles
left around the bases of each fascicle of spines. Lycoperdon echinatum has also a strong
tendency to a more flattened shape. The color of the immature plant has rarely been
noted. Rea says that the young plants are white. In our collection the spines on the
basal hah of freshly opened plants were pale creamy brown at the tips, cream colored
at the base, and those at the very top dark brown.

There is an odd contrast in the color changes in the present species and in L.
pulcherrimum. Both are light colored when young; in the last species as the plants
mature, the inner peridium soon becomes deep brown long before the spines fall off,
the bases of the spines in some areas becoming brown at times, but the tips remaining
white. In L. echinatum the opposite is the case, the tips of the spines soon become
brown, the bases and inner peridium remaining cream colored, in fact the peridium
remains cream colored for some little time after the spines fall off, turning brown
gradually on exposure, in some cases remaining very pale. The scurfy dots that later
form the reticulum are deep brown even in young stages.

Peck seems to have treated the present species as L. atropurpureum var. stellare,
as shown by a plant so determined by him at the New York Botanical Garden. A good
collection from New Zealand, labelled L. echinatum (gift of Mr. G. H. Cunningham),
looks exactly like our plants, but has distinctly smaller spores (3.5-4/x thick) and less
warted than in ours.

Lycoperdon compactum Cun. is a New Zealand species with similarly reticulated
peridium, but differing otherwise (Lycoperdaceae, etc., p. 195).

Illustrations: Dufour. Atlas Champ., etc., pi. 74, No. 169.
Fries. Th. C. E. Sveriges Gasteromyceter, fig. 14.
Hollos. 1. c, pi. 17, figs. 11-14 and 18-24.
Leuba. Champ. Comest, pi. 52, figs. 1-6.

PLATE 43

... . \ *.„

LYCOPERDON PULCIIERRIMl'M. No. 477

5

Ah

6

LYCOPERDACEAE • 75

Lloyd. Myc. Works, pi. 41.

Morgan. journ. Cin. Soc. Nat. Hist. 14: pi. 1, fig. 5. 1891.

Rolland. Champ. France, pi. 110, fig. 251.

Trclcase. Morels and Puffballs of Madison, Wisconsin, pi. 9, fig. 4 (as L. eonslellalum).

7421. On deciduous leaf mold, upland woods near Fern Walk, July 18, 1924.

Asheville. On mats of old leaves. Beardslee, coll.

New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb., as L. atropurpurcnm var. stcllare).

New York. Newcomb. House, coll. (Herb. N. Y. St. Mus. and U. N. C. Herb.) Spores appa-
rently not mature.

Wisconsin. Madison. Aug. 29, 1924. (Univ. Wis. Herb, and U. N. C. Herb.)

Manson. July, 1909. (Univ. Wis. Herb., as L. cepaeforme Bull., and U. N. C. Herb.) Spores
distinctly warted, 5-6.5/*, no pedicel.

Reported also from Kentucky, Ohio, Michigan, New York. Schweinitz's record from Pennsylvania
(No. 2259) is L. pitlcherrimum.

Lycoperdon rimulatum Peck.

Plates 41, 46, 47 and 112

Plants 1-5 cm., generally 2-3 cm. broad, depressed-globose, conspicuously plicate
below and narrowing quickly to a slender pointed root. Cortex light gray at first,
turning a deeper gray and often in age with reddish brown colors. Almost as smooth as
kid at first, but sooner or later the cortex becomes minutely dotted and cracked and
in age wears away irregularly and incompletely as very thin scales. When young the
surface may appear minutely fibrous-granular or almost completely smooth. In
exposed areas the inner peridium is seen to be smooth and shining, brownish gray and
often tinted with the spore color. Gleba changing from light yellow to gray, purplish
gray and then deep ferruginous brown or chocolate brown (auburn to chestnut brown of
Ridgway). Subgleba small, occupying only the lower fifth or sixth of the plant and
composed of large papery cells. When fully grown but still immature plants are cut,
water exudes from the surcharged peridium and escapes as drops, and this excess water
often water-soaks areas of the surface when the plants are brought in to dry; in these
respects behaving exactly like Bovistella radicata, which it further resembles in the long
pedicellate spores. Lloyd does not place this species in Bovistella, but it would seem to
belong there if his extension of the genus is adopted.

Spores (of Xo. 715) when mature a deep chocolate brown with faint tint of purple,
globose, covered with warts embedded in a clear, amorphous substance, some of the
warts extending slightly beyond this substance, giving the appearance of an outer wall
with radial pits, 5.5-6.8// thick, with one conspicuous oil drop and a stalk about twice,
or rarely nearly thrice, the length of the spore diameter. Frequently the stalk is broken
off. When KOH is applied the clear material surrounding the spore swells consider-
ably. Capillitium of simple or moderately branched, somewhat irregular and knotted
threads tapering at both ends and about 4-5^ wide in middle, occasionally 7/x wide at
knotted places. Rarely a short branch is observed on a thread. At a certain stage it
can be seen that threads are formed from the sclerotic thickening of certain selected
parts of the trama. These were not simple threads at first, but the branches that came
from them did not become hard and went to pieces with the spore mass at maturity.
The points where these branches joined the capillitium may be detected as knots or
short stubs.

Illustrations: Hollos. 1. c, pi. 18, figs. 36-38.

Lloyd. Myc. Notes No. 20: pi. 56, figs. 1-9. 1905.
Morgan. 1. c, 14: pi. 1, fig. 6.

76 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

715. On D -. WagstaS's lawn, September 8, 1913.
811. In old road to Rocky Ridge Farm, September 10, 1913.

7059. On wet ground in deciduous woods in Emerson's swamp, August 5, 1923. Spores dull brown
under the microscope, spherical, warted, 5.8-7.2m thick, often with a very long pedicel, up to 3
times the length of the spore. Capillitium threads 3.5^.8^ thick, pitted.
7160. On lawn among elms, September 27, 1923. Smallest plants about 1 cm. broad. Spores

distinctly warted and with hyaline material, 5.5-7^, often with a long pedicel.
7165. On ground in swamp of Emerson Farm, September 28, 1923. Largest plants up to 5 cm. wide.

Florida. Alachua Co. Couch, coll. June 24, 1924. (U. N. C. Herb., Nos. 7261, 7252, 7262, 7264.)
New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb., as L. cupricum var.).

Lycoperdon subvelatum Lloyd

Plates 41 and 112

Plant depressed-globose, 3 cm. broad and about 1.5 cm. thick in the dried state;
base pinched to a short stalk, ending in a stout cord which in our one collection branches
under ground to bear two fruit bodies; cortex thick, superficially areolated into appres-
sed, spongy patches on a granulated membrane, the whole peeling back from the top
in strips, some of which have fallen off. The inner peridium is glabrous, shining, light
grayish brown, without the scales which form the conspicuous furfurescence on the
denuded inner peridium of L. marginatum. Mature gleba deep brownish purple;
subgleba nearly white, occupying only the stalk, the chambers obvious.

Spores (of No. 7263) distinctly warted, 4.8-6// thick, furnished (in the dry state)
with long pedicels up to three times the diameter of the spores, but these pedicels fall
off when put into water. Capillitium threads irregular in size, wavy, encrusted in
places, up to 6.5/i thick.

This agrees in every respect with Lloyd's species (Myc. Notes, pp. 224 and 274).
It differs from L. rimidaium in the thick, continuous cortex which peels off in pliable
patches and strips. This peeling off in good sized patches recalls L. marginatum, but
the resemblance between the two is entirely superficial. We have four collections of
typical rimulatum from the same section of Florida, which is good evidence that the
present plant is not merely a regional form of that species.

Illustrations: Lloyd. Myc. Notes, p. 274, figs. 120 and 121; and pi. 56, figs. 10 and 11.

Florida. Alachua Co. Couch, coll. On sandy soil, July 28, 1924. (U. N. C. Herb., No. 7263.)

Lycoperdon umbrinum Pers.
L. hirtum Mart.
L. glabellum Peck

Plates 48 and 113

Plants depressed-globose with an abrupt base or pear-shaped with a tapering base;
up to 3.5 cm. broad and 4.5 cm. high, usually about 3 cm. high; white or light brownish
gray before maturity, turning to tawny buff or brownish buff or light grayish tan, or
rarely golden yellow. Surface lightly covered with a fine scurfy or granular stuff mingled
with minute, slender spines that are separate and erect or, especially on the lower half,
flattened toward each other in patches. These spines are often absent in places. This
superficial material is very persistent and is only partly worn away in old age to show
the shining inner peridium. Subgleba occupying the stalk-like or tapering base and
extending in a thin layer up to the widest part of the plant, with chambers of moderate

PLATE 45

'I

1

I

Lycoperdon echinati/m, Asheville, N. C. [upper four]

Lvcoperdon muscorum. No. 676 [lower left]. West Albany. Type [other two]

I.WOPERDACEAE 77

size. Gleba pale olivaceous then darker olivaceous brown at maturity, varying con-
siderably in color in different collections. It may be Isabella color (nearly old gold) or
Brussel's brown with a faint tint of purple or rarely a distinct purplish brown.

Spores (of No. 679) very small, spherical, about the color of the mature gleba,
3.7-4.5/1 thick and with a short pedicel, minutely asperulate; after soaking in water
appearing to have a thick, hyaline wall, the spines showing as darker strialions in the
hyaline material. Capillitium threads more or less uneven, with numerous or scattered
distinct pits in the walls, frequently branched, tapering to a point, averaging about
3.9-4.3ju in the larger threads, in places rarely up to 7/i thick.

Our collections have been compared with a good plant of L. umbrinum from Bresa-
dola's herbarium (labelled L. hirtum). The spinules on the top of this specimen are
somewhat larger than on any American plant we have seen, but otherwise the differ-
ences are trivial The spores are minutely but distinctly warted, 4-5.5/1, rarely with a
mucro; capillitium threads knobbed and strongly pitted. Our plants also agree well
with descriptions of L. glabellum by Peck and Morgan except that the spore color is
given as purplish brown and the spores as described are too large. However, we have
examined a plant from the type locality (West Albany), collected by Peck and possibly a
part of the type, and find that it agrees in these spore details and not with published
descriptions. Peck gives the spores as purplish brown, 5.1-6.3/x thick, but in his plant
we find them to be brown with little or no purplish tint and 4-4.8/1 thick, including warts.
Peck's specimen agrees further in showing the distinct pits in the capillitium, a character
that we find most useful in determining this species. The threads are also of the same
thickness and irregularity, up to 4.2/1 thick.

We cannot find any good difference between the present species and L. Turneri
as treated by Morgan (1. c, p. 17) and Lloyd (Myc. Notes, p. 236). The original
description of L. Tumcri E. & E. (Journ. Myc. 1: 87. 1885) was unfortunately
based on two different lots of plants, one, the true type from Labrador, and another
smaller and different plant from the eastern United States. This last we cannot dis-
tinguish from L. glabellum. The type from Labrador is represented in the New York
Botanical Garden and is a larger plant with much larger and rougher spores and with
the capillitium threads unpitted (see pi. 113, figs. 4 and 5). One supposed difference
between L. glabellum and L. Turneri is the purplish spores of the former, but this does
not hold, as most specimens of L. glabellum found in herbaria, including some of Peck's
own, as stated above, have brown to olivaceous gleba.

Lloyd considers L. glabellum a synonym of L. umbrinum (Myc. Notes, p. 225),
while Hollos treats it as a variety. The species is treated as L. glabellum by Morgan
(1. c, 14: 10). Lloyd thinks that L. elegans Morgan is the same. This is the plant
determined by Ellis as L. molle Pers., and Lloyd also considers the latter the same as
L. glabellum (Myc. Notes, p. 209). In a plant of L. molle from the Persoon Herbarium,
now at the New York Botanical Garden, the gleba is Isabella color and the spores
3.4-3. Sfi thick, faintly warted to smooth, smaller and less rough than in American
L. umbrinum (pi. 113, fig. 6).

Illustrations: Fries, Th. C. E. Sverigcs Gasteromyceter, fig. IS.
Hollos. 1. c, pi. 18, figs. 23-29; pi. 29, fig. 5.
Lloyd. Myc. Works, pis. 43 and 58.
Morgan. Journ. Cin. Soc. Xat. Hist. 14: pi. 1, fig. 7.

78 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

679. In grass on shaded lawn, Oct. 4, 1908.
1001. In pine woods, Sept. 15, 1913. Spores olivaceous, 3.7-4.6m thick. Subgleba in some plants

purple.
1027. Under cedars near campus. The plants in this collection were fully mature and were a pale

earth color (gray brown) outside; the spores and capillitium somewhat deeper earth color

within, and with a slight tint of olive. Spores 3.7-4/1 thick, smooth, spherical. Capillitium

delicate, branched, diameter of spores and tapering at the ends.
7215. On ground in pine woods, Nov. 17, 1923. One plant in this typical lot was strong golden yellow

on one side shading to brownish yellow on the other.

Also Nos. 1003a, 1004, 1026, 1762, 3233, 5921, 7103, 7148.
Asheville. Beardslee, coll.
Blowing Rock. Coker, coll.
Hendersonville. Alma Holland, coll. (U. N. C. Herb.). Spores 3.5-4/t.

Virginia. Warrenton. Coker, coll. (U. N. C. Herb.)

Florida. Alachua Co. Couch, coll. June 22, 1924. (U. N. C. Herb. Nos. 7253 and 7254.)

New York. West Albany. Peck, coll. (Albany Herb.). Also other New York plants from Peck

at N. Y. B. G.
Wisconsin. (U. N. C. Herb, from Univ. Wisconsin Herb.) Spores minutely but distinctly warted,

3.7-4.2/1, no pedicel.
Eagle Heights. Sept. 1903. (Univ. Wis. Herb, and U. N. C. Herb.) Spores distinctly rough,

4.2-5n, no pedicel.
Canada. London. Dearness, No. 915. (Dearness Herb, as L. molle.) Spores nearly smooth, with a

short pedicel, 3.4-4^.

Lycoperdon acuminatum Bosc

L. leprosum Berk. & Rav. in Peck
L. calyptrijorme Berk.

Plates 49 and 113

Plants very small, 3-9 mm., shaped like an inverted top or long, pointed egg, with a
small apical pore in age; cortex composed of a thin, spongy or granular scurf which very
slowly wears away exposing the dull inner peridium or in rare cases the cortex may as-
sume the form of short fascicled spines up to 1 mm. long above (as in plants from Cin-
cinnati and Newfield entered below) ; color whitish or pale tan, inner peridium yellowish
to gray or pale gray-brown in color. The entire peridium wall is very thin and delicate.
Gleba pale brown with a slight olivaceous tint. Subgleba none.

Spores (of No. 680) pale olivaceous, then dirty gray, globular, 3.3-4a< thick, smooth,
walls fairly thick, a large oil drop and a short mucro. Capillitium simple, about the
color of the spores, up to 10.8^ in diameter, averaging about 5.5-6^.

This is the smallest known puff ball and it is remarkable in its habit of growing on
mossy places on the bark of living tree trunks at some distance (usually 4-15 feet)
from the ground. Others give the habitat as on old logs or around the mossy bases of
living trees, but with us it always grows on living trunks. We have never found it
near the ground. It has exactly the same habits and preferences for cold weather that
Geaster leptospermus shows, and may often be found associated with it. This is the
same as L. leprosum in the sense of Ravenel, Peck, and others, as shown by plants from
their herbaria. The capillitium of Peck's plants shows slight branching, whereas in our
plants it is simple or very rarely branched, agreeing with Lloyd's description.

PLATE 46

JO

I

•*

Lycoperdon rimllatum. No. 715

W

<

a.

6

P

5

s

2

|v>

LYCOPERDACEAE 79

Illustrations: Hollos. 1. c, pi. 22, figs. 1-7.
Lloyd. I.e., pi. 64, tigs. 1-4. 1905.
Morgan. 1. c, 14: pi. 2, fig. 8.

680. On oak trees among moss about 5 or 6 feet from the ground, September 19, 1908, and on Cupres-

sus trees in a yard, Oct. 9, 1908.
°75a. In moss on a cedar tree, Dec. 9, 1913. Spores as usual, 3.2-3.6ji thick.
3221. On trunks of oak and hickory, Jan. 22, 1919. Spores smooth, spherical, 3-3. 7m, with a distinct

oil drop and often a short mucro.
7513. On bark of living white oak, Sept. 18, 1924.

Asheville. Beardslee.

Yadkin College. On branch of limits alata, Sept. 16, 1922. Coker, coll. (U. N. C. Herb.).

Hillsboro. (Curtis Herb.)

South Carolina. Aiken. Ravenel, Fungi Amer. Exs. No. 14 (N. Y. B. G. Herb, as L. leprosum).
New Jersey. Newfield. On mossy stump. Ellis, coll. (N. Y. B. G. Herb., as L. leprosum?).
Ohio. Cincinnati. (N. Y. B. G. Herb, from Morgan Herb., as L. calyptriforme Berk.)
Missouri. Perryville- Demetrio, coll. Rabenhorst-Winter, Fungi europaei, No. 3535, as L. leprosum
(N. Y. B. G. Herb.).

Lycoperdon pyriforme Schaeff.

Plates 49, 50 and 113

Plants pyriform, often crowded, up to 4.5 cm. broad and 4.5 cm. high, usually
much smaller; color red-brown when dry or old, paler when fresh; cortex composed of
minute warts, spines and granules which are very persistent and rough to the touch like a
file; sometimes cracked into distinct areas. The apical pore is very slow to form, often
taking three weeks or a month to appear after full size is reached. The plants are con-
nected with the wood and with each other by conspicuous, pure white strands of my-
celium. Gleba changing through greenish yellow to deep olive-brown at maturity;
subgleba rather small, occupying only the constricted base, the chambers very small.

Spores spherical, smooth, with a large oil drop, deep olive-brown when fully mature,
3.3-4.3/* in diameter. Capillitium branched, brownish olivaceous, main threads from
3.5-5.5^ in diameter, averaging about 3.7/z, tapering to the tips.

The spores and capillitium of our plants have been compared with those of Peck's
collections, and were found to agree. The species is often found in immense quantities
on rotten logs and stumps and rarely in the ground. The entire plant is very persistent
and after maturing in the fall may pass the winter with little change, and be found in
good condition in the spring. It is edible and when found is abundant enough to be
of use.

Illustrations: Dufour. Atlas des Champ. Comest. et Yen., pi. 74, No. 170. 1891.
Gillet. Champ. Fr. (Gasteromycetes), pi. 14.
Hard. Mushrooms, pi. 62 and fig. 470.
Hollos. 1. c, pi. 20, figs. 4-12.
Krieger. Nat. Geog. Mag. 37: 418, 419. 1920.
Lloyd. I.e., pi. 48. 1905.

Marshall. Mushroom Book, pis. opposite pp. 125 and 134.
Michael. Fuhrer f. Pilzfreunde 2: No. 200. 1918 (2nd ed.).
Murrill. Mycologia 6: pi. 127. 1914.
Patterson and Charles. U. S. Dept. Agric. Bull. 175: pi. 34, fig. 1. 1915; also Farmers' Bull.

796, fig. 17. 1917.
Yittadini. Monog. Lycoperd., pi. 2, fig. 9. 1842.

80 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

633. On stump of dead beech tree in Battle's Park, October 24, 1912.

917. Scattered in large numbers over several yards on earth and in and by the brick drain and in grass
just east of the Alumni Building, October IS, 1913. This colony of plants was interesting in
being apparently unconnected with wood. The mycelium could be traced as white strands for
some distance into the rather poor earth and then was dissipated into invisible fibers. All
characters exactly as in plants on wood. The plants were fully grown a month before the
apical pores appeared.
1046. On exposed dead root in Bowlin's Creek swamp, December 6, 1913.
Also Nos. 665, 935, 998, 1469.

Asheville. Beardslee.

Bldwing Rock. Coker and party, August 1922.

South Carolina. Ravenel. Fungi Car. Exs. No. 72. (Phil. Acad. Herb, and N. Y. B. G. Herb.)

Aiken. Ravenel. Fungi Amer. Exs. No. 469. (N. Y. B. G. Herb.)
Georgia. Harper, coll. (N. Y. B. G. Herb.).
Florida. Clark's Run. Calkins, coll. (N. Y. B. G. Herb.).

Marion County. On dead wood. Couch, coll. (U. N. C. Herb.). Spores smooth, 3.4-3.&H
thick.
New York. Alcove. Shear, coll. Ell. & Ev., Fungi Columb. No. 106 and N. Amer. Fungi No. 2906.

(N. Y. B. G. Herb.).
Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.)
Ohio. Cincinnati. Lloyd, coll. (N. Y. B. G. Herb.).
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).
New Zealand. Cunningham, coll. (U. N. C. Herb.). Spores smooth, 3.5-4.2ju thick, often with a

mucro.
Also from Venezuela, Ecuador, and Europe in N. Y. B. G. Herb.

Lycoperdon subincarnatum Peck

Plates 51, 57 and 113

Plants small, about 1-2.5 cm. thick, depressed-globose, almost or quite sessile,
growing on logs or stumps and attached by byssoid strands; color when young reddish
brown, at maturity becoming dark brown or dull leather brown; covered with minute,
separate, brown warts or with little pointed spines in groups with united tips, their
bases connected by a smooth thin tissue of the same color; after full maturity these
warts or spines and tissue slowly wearing away and leaving exposed the paler brown,
dull inner peridium, which is very firm and tough and is marked all over with small pits
outlined by acute ridges, appearing under a lens exactly like a thimble. This re-
ticulated surface is assumed after maturity, in any case, whether the outer peridium
falls away or not and is obvious through the outer peridium which accommodates itself
to it. Subgleba scanty or practically absent, composed of small pale chambers.
Gleba peculiar in the distinctly radiating tramal plates which, before maturity, give
a section very much the appearance of a Geaster; color at maturity grayish purple.
In some specimens there is a very distinct columella, thus increasing the resemblance
to Geaster.

Spores (of No. 3171a) dull brown, spherical, when dry appearing distinctly
warted, when in water appearing spiny under high power, 3.8-5/i thick, counting the
spines. Capillitium of long, simple or sparingly branched, very pale threads (much
paler than the spores) up to 6.5ju thick, the walls irregularly thickened in places, some-
times closing the lumen.

This species is a strongly marked one and is easily distinguished by the small deep
pits, growth on wood, sessile habit and peculiar capillitium. In most of our collections

PLATE 48

J 4

*■&&

Lycoperdox OMBBINUM. No. 1004 [above] ; Xo. 679 [below]

LYCOPERDACEAE 81

the peridium is much more rigid and leathery than in most puffballs, but in our No.
3171a from Chapel Hill it is more pliable, though still thick and leathery. In this lot
too, the outer peridium is more scurfy than warty and has not become at all worn off
though the plants are old, and the spores have slightly longer spines. The surface has,
however, assumed the pitted appearance in about the same fashion as other lots. All
other characters are the same.

Illustrations: Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 46, figs. 4, 5. 1926.
Lloyd. Myc. Works, pi. 63, figs. 5-9.
Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 2, fig. 6.

3171a. On very' rotten wood, Aug. 13, 1918.

7030. On rotten wood, Handcock's swamp, Aug. 3, 1923. Spores spherical, spiny, 4-4.8ai with a
distinct oil drop and a pedicel usually about one-half as long as the diameter of the spore.

Blowing Rock. Coker and party, Aug. 1922. No. 5556. On small pieces of rotten deciduous
wood and bark. No. 5641. On rotting deciduous log. Spores minutely spiny, 3.7-4.6m
thick. No. 5775. On very rotten chestnut stump. Also collection by Coker, Sept. 3, 1924.
Spores minutely warted, 3.5-4.2pi thick.

Linville Falls. Coker and party, Aug. 1922. No. 5754. On a rotting log in rich deciduous woods.

Asheville. Beardslee, coll. (U. N. C. Herb.). Spores spiny, 3.7-4.6ju.

Buncombe Co. Harvey, coll. (U. N. C. Herb.). Spores minutely warted, 3.5-Aft.

Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.).
New York. Sand Lake. Peck, coll. (N. Y. B. G. Herb.).

Arkville. Murrill, coll. (N. Y. B. G. Herb.).
Maine. White, coll. ( N. Y. B. G. Herb.).

Wisconsin. Devil's Lake. (Univ. Wis. Herb, and U. N. C. Herb.) Spores minutely but distinctly
warted, 3.6-4.2/1, with no pedicel.

Lycoperdon muscorum Morgan
L. pseudoradicans Lloyd
L. polytrichum Lloyd

Plates 45 and 113

We copy the following from Morgan's original description (Journ. Cin. Soc. Nat.
Hist. 14: 16. 1891):

"Peridium turbinate, globose or depressed-globose above, contracted below into a
stem-like base, with a filamentous and fibrous mycelium. Cortex a thin white or yel-
lowish coat of minute spinules with intermingled granules, which are coarser towards
the apex; these wither or shrivel with age and are mostly persistent on the smooth
olive-brown, shining surface of the inner peridium. Subgleba occupying little more
than the stem-like base; mass of spores and capillitium greenish yellow, then brownish
olivaceous; spores minutely warted, 4-4.5^ in diameter.

"Growing among mosses, especially Polytrichum, in old meadows and pastures.
New York, Peck. Peridium one-half to one and one-third inches in diameter and 1-3
inches in height. This is L. molle of Peck's U. S. Species of Lycoperdon."

Peck calls attention to the resemblance of this plant in the immature state to
Cahalia data. The mature plant is easily distinguished from the latter by the apicaJ
mouth, and the average size is smaller.

82 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Dr. House has kindly sent us two plants of the type collection of L. muscorum from
West Albany (Peck, as L. molle, 1. c, p. 310). The spores are spherical, minutely
warted, 4^-5.5/z with a short pedicel; capillitium threads about 3-4ju thick, pitted, easily
broken up.

Hollos considers this the same as L. polytrichum Lloyd and L. pseudoradicans
Lloyd. Lloyd states that his L. polytrichum grows only in hair cap moss, but he refers
to that species a collection from Maine by Beardslee, a part of which at least was not
growing in Polytrichum. Three plants of this collection seen by Lloyd and returned to
Beardslee show no Polytrichum, but a much smaller moss attached, and Beardslee
writes that the plants did not all grow in Polytrichum. The individual of this lot
figured by Lloyd (pi. 67, fig. 7) has a moss plant attached that looks like Polytrichum.
We find on examination of Beardslee's plants that the spores are like those of the type,
faintly warted or many smooth, usually with a mucro, 3.8-5//, rarely 6/*; capillitium
easily fragmented by slight pressure on the cover glass, pitted, often encrusted, up to 5/x
thick. It is to be noted that the resemblance to Calvatia data in general appearance
is further emphasized by the breaking up of the capillitium into short joints under slight
pressure. This fragmentation is found in all or nearly all Calvatias. It does not occur
in any other Lycoperdon we have seen. As to L. molle of Persoon, this cannot be that
species, as the plant in his herbarium, which may be considered the type, has smaller
spores, and according to the original description grows in oak woods. Fries says it
grows in mossy fields.

Illustrations: Hollos. 1. c, pi. 29, figs. 1 and 2.

Lloyd. Myc. Notes No. 9, figs. 50 and 51 (as L. pseudoradicans); also pi. 67, figs. 1-5 (as L. mus-
corum) and figs. 6-10 (as L. polytrichum) .
Lloyd. Genera of Gasteromycetes, pi. 10, fig. 45 (as L. muscorum, later changed to polytrichum) .

Connecticut. Miss White, coll. (N. Y. B. G. Herb.). Two plants on Polytrichum, one on Leucobryum

glaucum (det. by Mr. R. S. Williams).
Maine. West Harpswell. Eeardslee, coll. No. 676. (U. N. C. Herb.)
Ohio. Foerste, coll. (N. Y. B. G. Herb., as L. glabellum). On Polytrichum.

Lycoperdon gemmatum Batsch
L. excipuliforme Scop.
L. macro gemmatum Lloyd

Plates 52, 53 and 113

Plants 3-7.5 cm. high and 2.5-6 cm. broad, obovoid to turbinate with a distinct
base which is comparatively large and often elongated and stalk-like; growing on mouldy
woods earth or from decaying wood or leaves. Cortex made up of intermingled long
and short terete spines or warts, the short ones being considerably more numerous than
the long ones and all becoming smaller and more uniform downward and fading to
scurfy granules toward the base; the color grayish tan to buff or leather color, the
longer spines often darker. The large spines fall off first and leave pale smooth spots
that are very obvious and characteristic; the other warts fall very slowly, if at all, and
even in quite old collapsed plants only the top is free from them in areas so as to entirely
expose the smooth, straw-colored inner peridium. Gleba at maturity dull ochraceous
brown with a tint of olive or in some old plants brown with a tint of purple. Subgleba
occupying the stem-like base which is usually a third to more than a half of the whole,
its chambers up to 1 mm. wide.

PLATE 49

Lycoperdon acuminatum. No. 3221 [above]

Lycoperdon pyriforme. On chestnut log at Blowing Rock, N. C. [below]. Much reduced

LYCOPERDACEAE 83

Spores (of Xo. 3159) color of the gleba, usually smoky brown with a tint of
olive, spherical, very minutely asperulate, 3.7-4.4^ thick. Capillitium of long, spar-
ingly branched, brownish threads, about 5.5/u in thickest places, tapering to about 2/j,
heavily encrusted in places, frequently pitted, the color about that of the spores, ap-
pearing lighter when the spores are knocked out.

The plant is peculiar in the variation shown in fragility in various collections.
Some are as fragile as Cahatia data, and the peridium easily breaks up into small
fragments; in others the peridium is pliable and toughish as is usual in Lycoperdon.
Except for the characteristic warts, the more fragile collections could easily be referred
to Cahatia data. Furthermore, in some specimens of that species the capillitium does
not fragment so readily as in other Calvatias. For cytological information by Maire
covering this and other species, see under Sderoderma aurantium.

Morgan (1. c, p. 13) treats L. perlatum as distinct, but Lloyd thinks he misunder-
stood that species. Hollos considers them the same. Figures by Tulasne (Ann. Sci.
Nat. 17: pi. 2, fig. 11), as L. perlatum, do not show any roughness on the spores. Under
the name L. nigrescens, Lloyd treats an American plant that he considers "really a
form of gemmation excepting that the stiff spines are not consolidated" (Myc. Notes,
p. 229). The European plant with blacker warts he refers to on pages 212 and 338
(see his plates 47, 60, and 123), and in his last reference he considers it less like L.
gemmatum than he had formerly supposed. The American plants above mentioned as
L. nigrescens may be the same as those we refer to L. Peckii. It seems clear that the
original L. excipuUforme Scop, is only a form of L. gemmatum, as treated by Fries and
Lloyd. The name has also been applied, apparently without reason, to a form of L.
pyrijorme and apparently to a form of Cahatia saccata. A European plant (Herbarium
of Klotzsch), received from Ryks Herbarium at Leiden as Lycoperdon gemmatum (No.
58), has the external appearance of that plant, but is peculiar in having very small,
smooth spores, 2.3-3.2/x thick. We have found one cespitose group of plants very
much like this: spores nearly as small (2.8-3. 7/x) and nearly smooth (Linville Falls,
No. 8221).

Illustrations: Dufour. Atlas des Champ. Comest. et Ven., pi. 74, No. 167. 1891.
Fries. Sverig. Atl. Svamp., pi. 73, fig. 2. 1861.
Gibson. Our Edible Toadstools and Mushrooms, pi. 34.
Gillet. Champ. Fr. (Gasteromycetes), pi. 12.
Hard. Mushrooms, pi. 61.
Hollos. 1. c, pi. 19, figs. 14-25.
Krieger. Xat. Geog. Mag. 37: 414. 1920.

Lloyd. Myc. Works, pi. 46; also Lycoperdaceae of Australia, etc., fig. 35. 1905.
Micheli. Nova Tlant. Gen., pi. 97, fig. 1.
Michael. Fiihrerf. Pilzfreunde 1: Xo. 79. 1918 (2nd ed.).
Sweetzer. Univ. Oregon Leaflet, Bot. Ser. Xo. 9, fig. 1.
The figure by Murrill in Mycologia 1: pi. 15, fig. 3, 1909, is not this species.

500. On a piece of rotting wood, Oct. 4, 1912. Spores minutely warted, 3.5-4.4m thick.

502. Around and on roots of a cedar tree, Oct. 5, 1912. Spores in this collection slightly rougher than

in others.
1403. In rich damp places in the edge of woods, Oct. 22, 1914. Spores 3.7^1.5^ thick, rarely with a

short pedicel.
1784. Among dead leaves on Lone Pine Hill, Sept. 14, 1915. Spores 3.5-4.2m thick.
Also Xos. 554, 677, 678, 731, 2990, 3159.

84 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Asheville. Beardslee, coll.

Blowing Rock. Coker and party, August 1922. Common. No. 5669. On chestnut leaves.
Stalk 5 cm. long, 2.5 cm. thick above; head 4.5 cm. wide. 5670. Among frondose leaves in
woods. No. 5806. In leaves in deciduous woods. Spores very minutely rough, 3.4-3.9*1.

Linville Falls. Coker, coll. (U. N. C. Herb., No. 8221).

Salem. Schweinitz. (Schw. Herb., as L. excipuliforme.)

Winston-Salem. Schallert, coll. Oct. 1, 1922. (U. N. C. Herb.)

Alabama. Earle, coll. (N. Y. B. G. Herb.).

Virginia. Mountain Lake. Murrill, coll. (N. Y. B. G. Herb.).

Pennsylvania. Bethlehem. (Schw. Herb., as L. perlatum.)

Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.). Spores minutely rough, 3.5-4ji

with a distinct oil drop and often a short pedicel.
West Chester. Jackson, coll. (N. Y. B. G. Herb.).
New York. Alcove. Shear, coll. (N. Y. B. G. Herb.).

Osceola. House, coll. (Herb. N. Y. St. Mus. and U. N. C. Herb.). Spores 3.5^tM.
Massachusetts. Jamaica Plains. (N. Y. B. G. Herb., as L. perlatum.)
Wisconsin. (Univ. Wis. Herb, and U. N. C. Herb.) Spores faintly warted, 3.2-3.8*1.
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).
Washington. Langley. Grant, coll. (U. N. C. Herb.). Spores very minutely warted, 3.6-4.2/1,

no pedicel. Capillitium threads up to 6n thick, strongly pitted.
Canada. Manitoba. Victoria Beach, May 24, 1926. Bisby, coll. (U. N. C. Herb.).
Swedish Lapland. Abisco. Coker, coll. (U. N. C. Herb.). Spores olivaceous brown, minutely
warted, 3.5-4m rarely with a mucro. Capillitium threads irregular, strongly pitted, about 5m
thick.

Lycoperdon Peckii Morgan. A form

Plates 53 and 113

Plants shaped as in L. gemmatum, the distinct stalk up to 3 cm. long; the mycelium
ropy; peridium about 1.5-3.8 cm. broad, covered with tapering spines about 1-1.5 mm.
long, which as the plant grows split at base into 2-4 parts which remain united at their
tips and are easily rubbed off, leaving pale, smooth, circular spots which are surrounded
by minute granular warts and dots; color of the longer spines in youth and until near
maturity a delicate and pretty purplish lavender, at maturity changing to buffy brown.
Toward the base the spines become more slender and hairlike and are more obviously
intermingled with granular matter. Except for the lavender color of the upper half,
the plant until near maturity is nearly white. Subgleba composed of rather small cells
and occupying about a third of the plant.

Spores (of No. 5749) olivaceous brown, spherical, 3.6-4.4/j thick, with a small oil
drop and a short pedicel, the wall obscurely dotted (pitted or minutely warted) and
appearing faintly striate radially in optical section; surrounding the spore is the thin
hyaline layer that is found in most Lycoperdon species. Capillitium threads very
slightly branched, up to 6/x thick and tapering to delicate tips.

We have compared our plants with a collection in the Albany Herbarium from
Forestburg, N. Y., collected and determined by Peck and find them the same in appear-
ance, surface characters, spores and capillitium. The species (at least in so far as our
collections go) is easily distinguished in the fresh state before maturity by the lavender
color, a character not mentioned by others; in the mature and dry state it is distin-
guished by the ochraceous or buff color, by compound spines with flaring bases and
delicate, cohering tips, by granular matter between the spines, and by the small, dotted
(not obviously warted) spores with distinct pedicels. When the spines fall away they

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LVCOPEBDON SUBINCASNATUM. Ashcvillc, N. C. [top row], I

No. 5754 [all others]

LYCOPERDACEAE 85

leave smooth circular spots surrounded by minute granules, thus giving a spotted or
reticulated appearance to the surface as in L. eckinatum and L. gcmmatum. According
to -Morgan (Journ. Cin. Soc. Nat. Hist. 14: 15. 1891) this is the L. eckinatum of Peck's
United States Species of Lycoperdon, while the true L. eckinatum is treated as L. con-
stcllatum by Peck and by Trelease. In the Ellis collection at the New York Botanical
Garden are plants from Peck (N. V.) labelled L. eckinatum which are L. Peckii.
(See Morgan, above cited, and Lloyd, Myc. Notes, p. 223.) Lycoperdon Peckii is
not recognized by Lloyd, but we cannot reconcile our plant with any other species
and refer it to C. Peckii as it seems to agree well with Morgan's description and with
the above-mentioned plants from Peck. To all appearances this might represent the
L. nigresccns of Europe to which it might be referred except for the light color which
is retained at all ages (see Lloyd, Myc. Works, pi. 123, figs. 1-6). It differs from L.
eckinatum in the shorter pale spines and much smaller olivaceous spores; from L.
pidcherrimum in the much shorter spines, the pale surface of the inner peridium and the
smooth, olivaceous spores; from L. eximium in stouter, longer spines and the smaller,
spherical spores; from L.fuscum in the paler, stouter, longer spines with granular or
scurfy material between. The most important differences between the present species
and L. gcmmatum, which is nearest, and of which this might be treated as only a variety,
is the tapering fascicled spines with tips united, and the smooth spores. In these
characters it agrees with the plants from Peck above-mentioned, which are also similar
in the small, dot-like warts between the spines. As L. Peckii is described as having a
smooth surface (not spotted) after the spines fall off, a divergence from our plant in
this respect is indicated; Peck's specimens above-mentioned are, however, like ours.
A difference in age of plants might easily account for such a discrepancy. Our oldest
plants were found soon after opening and only a part of the upper surface was denuded.

North Carolina. Blowing Rock. Coker and party, August 1922. No. 5654. On rotten chestnut
wood. No. 5655. Among frondose leaves in woods. Spores smooth or nearly so, 3.5-4.2m
with a short pedicel. No. 5801. Under white pines. (U. N. C. Herb.)
Linville Falls. Coker and party, August 1922. No. 5749. Among leaves under rhododendron,
chestnut, etc. (U. N. C. Herb.)

Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. HertO.

Canada. Ontario. Dearness, No. 3026 c. (U. N. C. Herb.) Spores olivaceous in mass, smooth
or nearly so, 3.2-3.8/x.
Newfoundland. Waghorne, coll. (N. Y. B. G. Herb., det. by Peck). Spores smooth or nearly so,
3.4-3.8/1 thick.

Lycoperdon pedicellatum Pk.
L. caudatum Schroet.

Plate 113

Plants depressed-globose, plicate below, sessile or with a small stem, attached by
delicate fibers. Cortex pale, grayish then ochraceous to rather light ochraceous brown,
composed above of rather stout spines about 0.5-1 mm. long, single or arranged in
fascicles with the incurved tips united and the bases separated by a thin, glabrous
cortex of the same color. The spines fall off singly and each carries its own area of the
thin cortex, thus exposing the pale brown, smooth and shining, inner peridium which is
faintly pitted or wrinkled, or in a form distinctly reticulated. Subgleba very scanty
in nearly sessile forms or more obvious when there is a distinct stalk.

Spores (of No. 5668) dull brown at full maturity, subspherical to short oval,
3.7— 1.8/i thick, minutely rough, furnished with a long persistent pedicel which is about

86 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

4-5 times as long as the spore. Capillitium threads up to 6.6m thick, pitted, sparsely
branched, ochraceous under transmitted light.

Growing in humus or on logs in deciduous woods, and apparently northern in its
range except for the higher southern mountains (reported by Atkinson from Alabama).
It has not before been reported from our state. The species is easily recognized by the
rather long, deciduous spines, the pale, shining, inner peridium and particularly by the
very characteristic smooth spores with very long persistent pedicels. Lloyd includes
this species in his expansion of the genus Bovistella (Myc. Notes, p. 282), but we are
not including in Bovistella plants with the capillitium of Lycoperdon.
Illustrations: Hollos. I. c, pi. 20, figs. 44-47.

Morgan. 1. c, 14: pi. 2, fig. 2.
North Carolina. Blowing Rock. Coker and party, August 1922. No. 5668. On rotting wood of

a deciduous tree.
New York. Center. (N. Y. B. G. Herb.) Spores oval, smooth, 3.2-3.6 x 3.5-4/j, with slender, per-
sistent pedicels up to 3(V.
Wisconsin. Parfrey's Glen. (U. N. C. Herb, and Univ. Wis. Herb.) Spores oval, smooth, 3.4-3.8 x
3.7-4.5/x, with slender pedicels up to 35/i long.
Homewood. (Univ. Wis. Herb, and U. N. C. Herb.) Spores oval, smooth or nearly so, 3.5-4 x
4-5/i, with slender pedicels up to 35/i long.

Lycoperdon eximium Morgan

Plates 54 and 113

Plants up to 4 cm. broad, turbinate, distinctly stalked, even or plicate beneath.
Cortex composed of slender spines about 1 mm. long which often curve and converge
at the apex, and gradually decrease in length downward until they are mere granules
at the base of the stalk; color tan, brownish tan or deeper brown, and in age the spines
become very deep brown. After maturity the spines begin to wear away, so that in
age they are reduced to granular dots, or even these may be worn off in areas, so as to
expose the tan or light brown shining layer beneath. The mouth is a torn, irregular
opening of rather tardy appearance. Gleba greenish yellow, then brownish olivaceous
and at full maturity deep brown with a tint of purple, about bister of Ridgway. Sub-
gleba of honeycomb structure, occupying all of the stalk and extending with a thin
expansion to the broadest part, occupying in all about one-third to one-fourth of the
body.

Spores (of No. 1002) purplish brown, elliptic or oval, with a thicker, clear inner wall
and a darker outer wall which is dotted with minute warts, surrounded by a hyaline
coat, distinctly stalked, 4.3-5 x 5-6. 5m. Capillitium of long, sparingly branched,
deep brown threads that are almost the diameter of the spores, and taper to slender tips.

Recognized by its densely set, regular spines, dark color in age and rather large
oval spores. It does not seem to have been reported from this state before, and has
been known only from South Carolina and Alabama. The species is described by
Morgan as having brownish olivaceous spores, but in our fully mature plants with open
mouths the spores are deep purplish brown. Only a slight difference in maturity would
make this difference in color.

Illustrations: Lloyd. Myc. Works, pi. 59, figs. 6-8.

Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 2, fig. 3.
1002. In a pine grove, Sept. 28, 1913.

1762. In pine woods, Sept. 12, 1915. Spores minutely punctate, 4.4-4.8 x 5.5-6.3ju.
7144. On the ground in mixed woods, Sept. 26, 1923. Spores oval, minutely warted, 3.8-4.2 x4.4-5.4ji,

all with a short pedicel.
Also Nos. 501, 553, 588, 7150.

Asheville. Beardslce.

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LYCOPERDACEAE 87

Lycoperdon fuscum Bonorden

Plates 42 and 113

Plants, in our collections, broadly top-shaped or pyriform or subglobose, about
1.8-3.5 cm. broad, with a short stalk; covered with slender, delicate, dark brown to
blackish spines about 0.5 mm. long which are either single or grouped in fascicles of
several with the tips usually incurved, and which are long persistent; they are not
intermixed with other granular or scurfy material and on falling away finally they leave
the dark brown inner peridium quite smooth and shining. Subgleba small, in the pres-
ent form occupying only about a sixth of the body, in one form nearly absent (see below) ;
chambers of moderate size.

Spores (of No. 5525) dusky olivaceous, spherical, 4-4.8m thick, distinctly dotted
with warts which project through the hyaline coat. Capillitium threads slightly
branched, up to 3.7m thick, with thick, pitted walls.

The plant described above is the dark-colored, often large form with dark brown
inner peridium and persistent spinules. We also have the form known as L. dryinum
and one approaching the form L. cupricum. The former is a small plant with very short
black spinules, silvery brown inner peridium, and little or no subgleba (No. 5777).
The latter is a plant of moderate size with slender, rather scattered, black spinules and a
golden brown to coppery brown inner peridium (No. 5719). All forms are separated
from related species by the very slender spines which soon turn dark brown to black
and which are so openly placed as to clearly show the smooth surface of the inner per-
idium between them. These spines are the only covering of the inner peridium and are
not intermingled with scurfy or granular material. The resulting appearance in the
fresh plant is remarkably like that of a baby's head, set with thin dark hair. All
forms grow in deciduous woods on leaves or rotting wood.

Illustrations: Hollos. 1. c, pi. 21, figs. 1-7 and 18.
Lloyd. Myc. Works, pi. 45 (back), figs. 1-7.

North Carolina. Blowing Rock. Coker and party, August 1922. No. 5525. On bark of dead fron-
dose tree near ground. No. 5642. In thick humus on a mossy rock. Spores minutely warted,
pedicellate with ends of pedicels broken off and mixed with the spores, 4.4-5. 5m- No. 5777.
On leafy mold, deciduous woods (L. dryinum form). No. 5779. Among frondose leaves.
Typical; spores pedicellate, faintly dotted, 4.4-5m. (U. N. C. Herb.)
Linville Falls. Coker and party, August 1922. No. 5719. Among leaves in deep frondose woods
(L. cupricum form). (U. N. C. Herb.) Spores pedicellate, 4-4.8m, faintly warted and sur-
rounded by a hyaline coat.

Washington. Langley. Grant, coll. (U. N. C. Herb.). This is a form with minute dark warts instead
of spines. Spores minutely warted, 4.2-5. 5m, with a small, dark mucro. Capillitium
branched, up to 5m thick, distinctly pitted.

Lycoperdon marginatum Vitt.

L. papillatum Schaeff . in sense of Hollos
L. cruciatum Rostk.
L. separans Peck
L. caivescens B. & C.

Plates 55, 56 and 113

Plants 1.2-4.5 cm. broad and nearly always broader than tall; narrowing below to a
short stem and root. Cortex pure white, then a little discolored or pinkish, becoming
clay colored or deep brown in age, composed of densely set, sharp, erect warts which are

88 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

short, thick and simple or more slender and fascicled with their tips cohering, at matur-
ity breaking up into large or small flakes (see below), usually beginning in a median
line between the stem and top, but often rupturing at every point or at many points at
once (see below for woods form). The lower third, covering the stem and lower part
of the body, does not fall away. Inner peridium strong to pale olivaceous brown when
first exposed, even, or in one of our collections (Warrenton, Va.) obviously pitted, much
as in gemmatum. It is peculiar in having beneath the spines a covering of a minute
brown furfurescence which disappears rather slowly and leaves the surface paler and
shining. This furfurescence is different from that of any other plant we know. It is
composed of little flat scales, the prevailing shape of which is spherical to oblong, vary-
ing to irregular and angled, the majority elongated and up to 60 x IOO/j. Subgleba
ample, composed of distinct, rather large compartments, occupying the stem and lower
part of the body and persistent after the top has disappeared, remaining as a dark brown
pedicel with a thin, collapsed margin.

Spores (of No. 482) spherical, smooth, 3.6-4. lju thick, with one oil drop. The
spores have mixed with them about an equal number of small rods like bacilli, that
average about twice the length of the spore's width. Basidia (of No. 5225) 6.6-7.4,u
thick, usually 4-spored. Capillitium composed of yellowish brown fibers very little
branched, about as thick as the spores as a rule, but up to 5/x in diameter, attenuated
at the ends.

The spores and capillitium of our plants have been compared with Peck's collec-
tions of separans and are the same. One of the most characteristic qualities of the plant
is its strong odor when ripening or fading like that of horse urine, a distinction it shares
with L. gemmatum. It is a very common plant with us in open places and cultivated
ground, and less often in woods. The spines are very variable, in some cases very thick
and short and not split up into fascicles, in others slender and moderately short or long,
and in groups of several with their tips united. In Chapel Hill there are two rather
distinct forms; in one (typical) of cultivated or open ground the cortex falls off in larger
flakes and more irregularly and the spores have a little mucro left after the pedicel
breaks off; in the other one of woods each group of spines with its underlying tissue is
apt to fall off singly or with a small group of others, and the denudation is more regular,
proceeding from the top downward. In this form the spores have no mucro. The
scales on the denuded surface are just like those of the typical form. This character
alone is sufficient to distinguish the species. For general discussion of the species see
Lloyd (Myc. Notes, pp. 214 and 231).

It seems certain that L. cruciatum is L. marginatum, as Rostkovius's good figure
could not be referred to anything else. Spores of three specimens labelled cruciatum
in Bresadola's herbarium (sent by Romell) are not those of the present species. This
is true also of a specimen labelled marginatum in the same herbarium We have seen
European material of true L. marginatum, as L. cruciatum (Herb. Hasskarl & Herb.
Klotzsch), received from the Ryks Herbarium in Leiden. They show the same spores
and peridial scales as in our specimens (spores 3. 6-4. 5m). The capillitium is not color-
less but of about the same tint as in ours. We have compared capillitium of plants
just maturing in our No. 482 with other mature ones of the same collection and find that
the capillitium does not change after the spores are mature.

A bit of the inner peridium and spores and capillitium of L. cahescens B. & C.
from the Curtis Herbarium (Wright, No. 6366, co-type), kindly sent us by Dr. Thaxter,
shows it to be marginatum. The peridial scales and the spores are the same as in the
typical form of L. marginatum. Lycopcrdon pratense Pers. is also considered a synonym

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LYCOPERDACEAE 89

by Hollos, but Lloyd thinks that the typical pratense docs not occur in the United
States. A specimen from New Zealand determined as L. praiaisc by Lloyd docs not
look like our L. marginatum, as the spines are much shorter and the denuded pcridium
shows no obvious furfurescence. The spores are slightly smaller and both they and the
capillitium are paler than in our plants.

From the description and illustration, L. abscission R. E. Fries from Bolivia and
Argentina is nearest L. marginatum. It is remarkable in the comparatively large size
of the subgleba which occupies most of the fruit body.

Illustrations: Hani. Mushrooms, fig. 467.

Hollos. 1. c, pi. 20. tigs. 17-22; pi. 29, fig. 6 (as L. papulation).

Lloyd. Myc. Works, pi. 51 (as L. cruciatum). 1905.

Lloyd. Photogravure of Am. Fungi, No. 3 (as L. separans).

Morgan. 1. c, 14: pi. 2. fig. 1.

Murrill. Mycologia 1: pi. 15, fig. 7 (as L. Wrightii). 1909. The three plants on right may

be true L. Wrightii.
Rostkovius in Sturm. Deutsch. Fl. Tilze 1: pi. 8 (as L. cruciatum). 1813.
Vittadini. Monog. Lycoperd., pi. 1, fig. 11. 1843.

99. On ground in cemetery, October 27, 1911. Spores spherical, smooth, 3.6-4m thick, with a
very distinct oil drop.

482. Glen Burnie meadow and on campus, October 3, 1912.

922. In mixed woods, September 13, 1913.

1762b. In pine woods, September 12, 1915.

5225. In a lowground pasture, June 23, 1922.

7194. In mixed woods, October 14, 1923. Spores 3.7-1.2m thick, with pedicels entirely broken off.

Haywood County. In mold in semi-open place by Pigeon River, Aug. 9, 1926. Coker, coll.
(U. N.C. Herb., Xo. 8131).

South Carolina. Ravcncl. Fungi Car. Exs. No. 73. (Phil. Acad. Herb, and N. Y. B. G. Herb., as

L. gemmation .)
Florida. Couch, coll. (U. N. C. Herb., Nos. 7266-7272).
Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb.).
Virginia. Mountain Lake. In moist woods. Murrill, coll. (N. Y. B. G. Herb.). This is exactly

like our woods form No. 7194.
Warrenton. Coker, coll. (U. N. C. Herb.). Spores faintly rough, 3. 5-4.2^1. Woods form.
District of Columbia. Cook, coll. (U. S. Nat'l. Herb., as L. cruciatum).

New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb., as L. Wrightii and as L. pcdiccllatum) .
New York. Sand Lake. (N. Y. B. G. Herb., asL. separans.)

Newcomb. House, coll. (Herb. N. Y. St. Mus. and U. N. C. Herb.). Spores smooth, 3.7^.2^.

Scales on inner peridium as usual.
Connecticut. Miss White, coll. (N. Y. B. G. Herb.).
Ohio. Lloyd, coll. No. 2655. (Bres. Herb., as L. separans, and U. N. C. Herb.)

Cleveland. Beardslee, coll. (Bres. Herb., as L. separans, and U. N. C. Herb.). Spores 3.6-4.2^.
Wisconsin. La Crosse. (Univ. Wis. Herb, and U. N. C. Herb.). Spores smooth, 3.5-4/x, sometimes

with a mucro.

Lycoperdon Curtisii Berk.
L. Wrightii B. & C.

Plates 57 and 113

Plants small, globose or depressed-globose, 0.7-2.3 cm. (rarely 3.5 cm.) thick,
averaging about 1 cm., with a distinct tendency to cespitose growth, generally sessile

90 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

or with a very abrupt, short stem; surface thickly set with white or whitish, rather
persistent spines, which are usually convergent at their tips in stellate groups. With
age these fall away in part from the top of the plant, but the powdery scurf between
them wears away more slowly so that the pale brown inner peridium becomes very
tardily smooth. Gleba greenish yellow, then light brownish yellow to brown with a
tint of olive. Subgleba quite small but distinct, the compartments of moderate size.
Spores (of No. 552) small, spherical, minutely asperulate, 2.6-3.7ju in diameter,
averaging about 3.3,u. Capillitium of almost hyaline, sparingly branched, flaccid,
septate threads, 3-7// in diameter, averaging about twice the diameter of the spores.

This little plant has almost the external appearance when immature of L. margi-
natum, but averages much smaller and also differs in a strong tendency to crowded
growth. The cortex does not flake off as in L. marginatum, but this difference cannot
be made out until full maturity. The plants of No. 539 were thought at first to be
L. marginatum and were watched for two weeks. As no cracking occurred they were
brought in and found to differ from L. marginatum in the color of the spores and the
character of the capillitium. In the herbarium at Albany some of Peck's collections
labelled L. Wrightii are L. marginatum. As Morgan points out (as L. Curtisii), the
thin-walled, collapsing capillitium is a peculiar character. We have examined the
co-type of L. Curtisii No. 197 and find the spores to be spherical, minutely rough,
3-3. 7/i thick. Morgan treats both L. Wrightii and L. Curtisii, the latter being the plant
as we have it. What plant Morgan treated as L. Wrightii we do not know (see Lloyd,
Myc. Notes, p. 153). Lycoperdon Wrightii is in reality a synonym of the present
species, as shown by the co-type in the Curtis Herbarium (Wright, No. 7; Curtis, No.
5633). The spores are spherical, minutely warted, 3-3. 6/x thick. Another collection
so labelled in the Curtis Herbarium from New York (Peck) looks like L. pusillum, while
still another from New Jersey (Austin) looks like L. marginatum (L. cruciatum), as
Lloyd says (Myc. Notes, p. 153). For the distribution of this species see R. E. Fries,
I.e., p. 8.

We have examined a specimen of L. candidum Pers. from Persoon's herbarium,
kindly sent us by Miss Cool. The appearance of the cortex is like that of L. Curtisii,
and the spores agree in size (3-3. 8/x) but run smoother (pi. 113, fig. 23). The plant
seems to be not quite mature, and this may account for the difference. The Lye.
candidum of Schweinitz Herbarium as represented in the Michener Herb. (No. 2747)
is Bovista plumbea. Lycoperdon candidum (Rostk.) Bon. is a different plant, now
put under Cahatia.

Illustrations: Hard. Mushrooms, fig. 468 (as L. Wrightii).
Lloyd. Myc. Works, pi. 63, figs. 1^1 (as L. Wrightii).
Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 2, fig. 4.

552. In grass on the campus, Oct. 10, 1912.
1718. On bare earth in a yard, Sept. 10, 1915.
7546. On bank of a ditch in the Arboretum, Oct. 1, 1924.
7549. In manured grass on campus, Oct. 2, 1924.
AlsoNos. 539, 984, 7151.

Asheville. Beardslee.

Linville Falls. Coker and party, Aug. 1922. No. 5760. On ground in a pasture. (U. N. C.
Herb.)

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LYCOPERDACEAE 91

District of Columbia. Lloyd, coll. (lT. S. Natl. Herb.).
New York. DnionporL Haiper, coll. I X. V. B. G. Herb.).

Wisconsin. Eagle Heights. (Univ. Wis. Herb, and U. N. C. Herb.) Spores subsphcrical, nearly
smooth, 3-3.$^. Capillitium very pale and easily collapsing.

Lycoperdon oblongisporum B. & C.

Plates 58 and 113

Plants globose or depressed-globose, with a distinct cord-like root. Cortex of
minute appressed fibers arranged in groups so as to form whitish, scurfy spots on the
pale, dull brown surface of the inner peridium, which is tardily exposed. Gleba oliva-
ceous to dark olive brown; subgleba none.

Spores (of Xo. 497) oval to subspherical, deep olive brown, smooth, 3-4 x 4.3-5.5//,
with a large oil drop and a prominent mucro about 1/u long. Capillitium threads long,
delicate, branched, up to 6.5/z thick, considerably pitted in places.

This little plant was originally described from Cuba, and is quite rare in this
country, only a few scattered collections having been noted before and none from this
state. However, it agrees so well with the descriptions of both Morgan and Lloyd for
this species that we feel quite sure our determination is correct. The plants look like
Lloyd's figures, and the spores are shaped like those shown on his plate 65, figs. 9-12.
The species shades into L. pusillum by intermediate forms. We are entering No.
5312 as this species on the ground of its absolutely smooth spores that are not perfectly
spherical. Xo. 5211, which we are entering under L. pusillum, is also intermediate.

Illustrations: Hollos. 1. c, pi. 21, figs. 36-41.
Lloyd. As cited above.

497. In poor soil on campus and in Arboretum, Oct. 1, 1912.

5312. In a field of Sudan grass, July 8, 1922. Plants up to 2.3 cm. thick; subgleba none; gleba gray-
brown; the spores themselves brown with a faint olive tint, smooth, spherical to slightly sub-
spherical with the proximal end pointed, 2.8-4m thick. Capillitium threads pitted, up to 6/i
thick.

7116. On ground in a pasture, September 9, 1923. Spores 3.4-3.7 x 4-4. 4^.

Asheville. Beardslee, coll.

Lycoperdon pusillum Batsch

Plates 58 and 113

Plants subglobose or flattened, small, rarely up to 2 cm. broad, pinched to a cord-
like root; surface variable, usually almost smooth to naked eye, but with a lens it is
seen to be covered with a fine fibrous flocculence that seems to flatten down like drying
suds and separates into very small patches, giving a maculated or areolated appear-
ance, or rarely this flocculence takes the form of small soft spines; again the surface
may be entirely covered with a delicate powder or more rarely minute compact warts.
Color when young pure white, then clay color or dull olivaceous yellow, and often
dark brown in age. Gleba passing through yellow to greenish yellow, becoming at full
maturity brownish olive or grayish olive-brown to deep coffee-brown without a tint of
olive, filling the entire peridium; that is, there is no subgleba. The peridium before
maturity is thick, up to 15 mm., but rapidly becomes extremely thin as it dries during

92 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

maturation. There is no breaking away of a superficial part. At maturity a small
apical pore is formed.

Spores color of the gleba, spherical, 3.2-4.3/j. thick, with rather thick wall, a distinct,
small mucro and a large oil drop, quite smooth in some plants, in others with the walls
faintly warted or pitted, appearing radially striate. Outside of the main brownish
wall a faint hyaline coat may be seen under high power. Basidia (of No. 5211) short-
clavate, stout, 4.5-5.5 x 9-1 In, with 4 or in many cases only 2 long sterigmata visible
at one time. CapiHitium color of the gleba and composed of long, delicate, branched
threads that are attenuated at the ends and are about the width of the spores, some up
to 6.5^i thick; walls more or less pitted, the larger threads usually more pitted than the
smaller ones.

This is a rather common little plant in Chapel Hill in open places, fields and yards.
It is a puzzling species due to variation in the spores which is not associated with any
other character we can make out, such as size, subgleba or surface. The capillitium is
the same in all. None of our plants has a subgleba.

Some collections that we are retaining in L. pusillum are intermediate between that
and L. oblongisporum, as No. 5211, which has spores that are mostly oval and pointed
toward the mucro, smooth to very minutely punctate.

98. In grass on ground in cemetery, Nov. 10, 1911. Spores minutely asperulate.
2209. On bare soil under cedars, June 23, 1916. Spores 3. 2-4.3^.
5202. On ground among grass and moss, June 19, 1922. Spores 3.2-4.2^ thick, all minutely spiny

and only a few with a mucro.
5390. Sandy old field, July 23, 1922. Spores appear smooth in water even when magnified 2160 times;

warted, however, under same magnification when nearly dry.
Also Nos. 1003, 1025, 3268, 5211.

Asheville. Beardslee, coll.

South Carolina. Ravenel, coll. (N. Y. B. G. Herb.). Spores smooth, 3.4-4^ thick.

Florida. Gainesville. Couch, coll. (U. N. C. Herb., Nos. 7256-7259, 7264, 7265). Spores (of No.
7259) smooth or nearly so, 3. 5-4. 2m thick, with a mucro.

New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb, and U. N. C. Herb.). Spores spherical,
minutely warted, 3.5-3.8/1.

New York. Newcomb. House, coll. (Herb. N. Y. St. Mus. and U. N. C. Herb.). Spores minutely
rough, 3.6-4/i.

Wisconsin. Blue Mounds. (Univ. Wis. Herb, and U. N. C. Herb.) Spores smooth or nearly so,
3.4-3.8/1.

North Dakota. Kulm, Aug. 1909. Brenckle's Fungi Dakotenses No. 93 (Path, and Myc. Herb., as
L. cepaeforme). In size some of these specimens approach L. cepaeforme, but there is no sub-
gleba. Spores spherical or subspherical, 3.5-3. 8^, smooth, usually with a mucro. Capil-
litium rather pale and irregular, encrusted, up to 7.4ju thick.

Pacific Coast near Canadian boundary. Dearness, No. 5550. (U. N. C. Herb.) Spores minutely
warted, 3.5-4.2/i.

Lycoperdon polymorphum Vitt.
L. cepaeforme Bull.

Plate 113

Plants subglobose, often contracted into a stalk-like base, and characteristically
with a conspicuous, stout root which may ramify in the earth and connect individuals
of a group; size variable, 1.3-3.5 cm. broad and up to 4.5 cm. high. Peridium in the

PLATE 57

«/

Lycoperdon* subincarnatum. No. 5556 [above]
Lycoperdon- Curtisii. No. 552 [below]. X 1 1/3

PLATE 58

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Lycoperdon oblongisporum. No. 5312 [above]
Lvcoperdon pusillum. No. 3268 [below]

I u OPERDAI I M 93

dry state tlull brown to olivaceous brown, varying to a brighter reddish yellow, thin
and papery, in the Wyoming plain noted below, really divided into two extremelj thin
layers, the outer of which ma] 01 may not scale ofi in pan; the surface dotted with
minute, scattered, persistent Seeks <>r nodules or varying to more closely plat ed granular
particles. Gleba dull olivaceous brown at maturity; Bubgleba obvious to large, filling
the stalk-like base, about the same color as the gleba and remarkable in iis compai I
structure with chambers so small as to be invisible, thus easily separating this species

from all other American species of /.v. ojnrdoti (Calvotia ruloo flaw has no chambers

at maturity).

Spores (of plant from Wyoming) smooth or rarely faintly dotted, spherical, 3.4 4/i,
with a conspicuous "il drop and sometimes a short pedicel. Capillitium threads

branched, averaging about T7^ thick.

Gregarious on earth in open places. We have not found it in North Carolina, but
it is reported from a number of eastern stales, and the < rjuiclonv.c form is said by l.loyd
to be common. This last form is the more spherical one without a stalk like base. In
plants we have from Wyoming, kindly sent us by Mr. Simon Davis, both forms are
well represented and apparently from the same collection. One of the plants has as
distinct a stalked lease as thai from Kurope shown by Lloyd on his plate 52, fig. 4. We

have a good European specimen from Rome]] (Sweden) which agrees with the Wyoming

plants in every particular except that the peridium does not divide into two papery
layers in age (spores smooth, id 4/j); also one from Ton cud ( Portugal) which is similar
(spores smooth, 3.S 4.2/j, rarely with a short pedicel).

This species is closely relai ei I to /.. pusillum, and Lloy.l thinks thai they intergrade
With us I., pusillum is a very common little plant without subgleba, and it doesnot vary

here to the larger forms of the above description. Its root is distinctly more slender

than in L. polytHorpkum of the same size. Its spores are Indistinguishable, l.loyd

shows forms of this species from Australia with several plants of a colony connected by
underground rhizomorphs (Myc. Notes, p. 730, fig. 1096). If is interesting to note
that some of our plants from Wyoming show this character clearly.

Cunningham gives the spores of the present species as 1.5 5.,5/j. Other characters
agree well with the above. It is supposed bv some authors, as Hollos, that /.. poly-
morph urn is the same as L.furfuraccuni SchaelT., a species which is insufficiently known.

Illustrations: Hulliard. Herb. Fr., pi. 435, fig. II (as L. ccparforme).
Cunningham. Trans. N. Z. Inst. 57: pi, S, lie;. 12. 1926.

Hollos. 1. c, pL 21, figs. 19 23{uL.futfvractUM).

Lloyd. My.. Work-, p] 34, figi, l o; pi. 52; pi. S3, figi i i (u L. upotform); pi. 65, tins. 1-7

(as / ■me).

Morgan, Joum.Ciii.Soc. Nat. Hist. H: pi. 2, Mr. ') (as /.. ccimcformt).

Vittadirii. Moii.. ■ I -. , pi. >, lie,'. 7.

Wyoming. Pitchfork. Simon Davis, coll. (U. N. ('. \l- rfa

Reported also from many other states (tee Lloyd, . p. 2.14).

Lycoperdon coloratum Peck

Plants Bubglobose, up to 3 cm. thick, pinched in at base, firmly attached cither by

one conspii UOUI trand or by more numerous white libers which may often be seen to

connect beneath the surface small Bclerotia like initials. Color from lirsi appearance

until near maturity a brilliant golden yellow; surface dotted with minute, harsh, dis-

\J*l\ "*

94 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

crete warts or nodules which during maturity become dark brown to nearly black and
slowly wear away separately, exposing the shining bright brown peridium. Young
gleba white then passing through yellowish to dull dark brown at maturity; subgleba
very small, with small but distinct cavities under a lens. Odor while maturing strong
and fragrant.

Spores (of No. 8044) spherical, nearly smooth but not quite, 3.5-4^ thick. Capil-
litium (of No. 8108) irregular, main branches 4-5. 5m thick; color dull brown with a
tint of vinaceous under the microscope, no obvious pits in the walls.

Until dulled by maturity, this striking plant is by far the most brilliant of our
puffballs. No description by others suggests the brilliant orange of our mountain
plant, though Peck speaks of the species as conspicuous. While the plant may be only a
form of L. polymorphum, it is so conspicuous and distinctive in itself that it will do no
harm to retain Peck's name for it. In the mountains of western North Carolina, L.
piisillum grows in pastures, and L. coloratum in the woods. They are strikingly
different.

Illustrations: ? Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 2, fig. 10.
PTrelease. Morels and Puffballs of Madison, pi. 2, fig. 2.

North Carolina. Haywood County. On ground in mixed woods by Crawford's Creek, Aug. 2, 1926.
Coker, coll. (U. N. C. Herb., No. 7995). Under rhododendron and hemlock by Cold Creek,
Aug. 6, 1926. Coker and Totten, colls. (U. N. C. Herb., No. 8044). Woods mold by Cold
Creek, Aug. 8, 1926. Coker, coll. (U. N. C. Herb., No. 8108).

BOVISTELLA Morgan

Plants subglobose and permanently attached by a rooting base; inner peridium
thin, flaccid, opening by a definite, apical pore, and collapsing as the spores escape;
capillitium as in Bovista, of distinct units with all the ends tapering and not connected
with the peridium; but in some cases very slender and not abundantly branched;
spores oval or subglobose, smooth (at least in American species), and with long pedicels.

Separated from Lycoperdon by the discrete, branched units of the capillitium;
from Calvatia by this character and by the torn mouth ; and from Bovista by the col-
lapsing inner peridium, and by the permanent attachment to the soil. The above is
essentially the conception of the genus by Morgan who established it (see below),
except that we are including species without a sterile base, as does Lloyd. We are not
following Lloyd in including plants with long, intertwined capillitium fibers which arise
from the periphery as in Lycoperdon. Without this character we would not know how
to separate the latter. Lloyd would extend the limits of the genus as follows (Myc.
Notes No. 23: 277. 1906).

"Peridium flaccid, with or without a sterile base, opening by a definite mouth.
Capillitium of short, separate threads or long, intertwined threads. Spores pedicel-
late." Note that the flaccid peridium is the only character in this definition that would
exclude species of Bovista. However, in his discussion, Lloyd makes the further dis-
tinction of the breaking away from the base in Bovista and the permanent attachment of
Lycoperdae, including Bovistella. He says that frequently the characteristic capillitium
and pedicellate spores are associated in the same plant, but not always, and he would
"embrace in the genus Bovistella all plants of the tribe Lycoperdae that have either or
both of these characters."

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LYCOPERDACEAE 95

Literature

Cunningham. The Genera Bm'sta and Borisklla. Proc. Linn. Soc. New. South Wales 50: 367, pi.

37. IMS.
Lloyd. As cited above.

Morgan. Joum. On. Soc. Nat. Bust 14: 141. 1892.
Petri. I"IoraItalicaCryptogama(Gastcrales),Fasc. 5: 58. 1909.

Key to the Species

Plants of moderate size; sterile base obvious B. radhata

Tlants small; sterile base none

Plants epigeal; capillitium units very slender, the main branches only about 6-7 'ji thick. . B. echir.clla
Plants subterranean until maturity; main threads of the capillitium up to 18.5m thick

See Bovista minor

Bovistella radicata (Mont.) Pat.

Bovistella ohiensis Ell. & Morg.

Plates 59 and 114

Plants globose or broadly top-shaped, up to 7 cm. broad usually, but rarely as large
as 14.5 cm., often plicate below; attached by a firm, thick, tapering root; cortex com-
posed over the upper half or more of soft, pyramidal warts which are often fused at
their tips into fascicles as in species of Lycoperdon, these fading towards the base to a
dense soft scurf; white at first, then dull ochraceous when dry and wearing away ir-
regularly and incompletely to expose the thin, papery, inner peridium which is metallic
buff or silvery buff or ochraceous in color, rupturing at maturity by an apical pore or
slit which extends by fission on exposure and finally by collapsing and incurling leaves
the entire subgleba exposed in age. Subgleba large, broad, occupying about one- third to
one-half the plant, the edges extending up the sides of the peridium to form a cup-
shaped base which is long persistent. Gleba at first white and cheesy, then passing
through yellowish to brown (about Brussels brown of Ridgway) ; chambers of the young
gleba 25-250^ wide, hollow. Before ripening begins, a section of the peridium shows
that the scurfy and warty superficial coat is an extension of a thick, compact, pulpy,
tender layer about 450,u thick of bladdery cells beneath which lies the thin, very tough
inner peridium about 130m thick, composed of densely woven, slender filaments about
3.7/i thick.

Spores oval, smooth, 3.5-4.5 x 4.3-5. 2ju, with one large oil drop and long, hyaline,
persistent pedicels which average about 11m long. Basidia short, 6.6/u thick, with 4 long
sterigmata which break off with the spores. Capillitium threads free, short, 3-5 times
branched, the main stem 7-12/x thick, the branches tapering to about 3.5/x thick near
the ends.

At Hartsville, S. C. (see below), we have collected a very robust form reaching
14.5 cm. broad, which is considerably larger than the form figured by Lloyd (cited
below). The species is common in Chapel Hill and prefers cultivated soil and open
places. When quite fresh and white the taste is sweetish and pleasant and the odor
slight, but as the color begins to change the odor becomes rather strong and nitrous.

As represented in Torrend's Fungi Selecti Exs. No. 77, Bovista radicata is the same
as our plant and confirms Patouillard's conclusion (accepted by Hollos) that our Ameri-
can plant is B. radicata. The type was described from northern Africa (Syll. Crypt.,
etc., Paris, 1856). Plants from California distributed by McClatchie as B. ammophila
Lev., a European species, now included in Bovistella, are considered by Lloyd as B.
plumbea (Myc. Notes, p. 88).

96 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustrations: Hard. Mushrooms, fig. 473.
Lloyd. Myc. Works, pi. 86.
Marshall. Mushroom Book, pi. opposite p. 128.

79. In a corn field, October 12, 1909.
3903. In pine woods, Nov. 11,1919.

5200. Strowd's pasture, June 17, 1922. Spores 3.7-4.2 x 4-5m, with pedicels about 12m long.
5919. On ground in pasture, Nov. 22, 1922. Spores 3-4 x 4-5.5/i. Capillitium threads up to 5.5m thick.
Also Nos. 537, 1018, 1428, 1520, 2438, 3010, 5361, 6094.

Asheville. Beardslee.

Flat Rock. Memminger, coll. (N. Y. B. G. Herb.).

South Carolina. Hartsville. Coker, No. 1545. In a sandy cow pasture, June 12, 1915. A very stout
form, up to 14.5 cm. broad, but the spores and all other characters just as in the Chapel Hill
plant.
Georgetown. Coker, No. 6015. In sandy woods pasture near Silver Hill, Dec. 29, 1922.

Florida. Couch, coll. (U. N. C. Herb., No. 7276). Also collections by Weber and Walker.
Clearwater. Rousseau, coll. (U. S. Nat'l. Herb., as B. ohiensis).

Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb., as B. ohiensis).

New York. Botanical Garden. Murrill, coll. (N. Y. B. G. Herb., as B. ohiensis).

Indiana. Gentry, coll. (N. Y. B. G. Herb., as B. ohiensis). Main threads of capillitium about 11m
thick. Spores oval, smooth, 3.4-4.2 x 3.7-5. 5m, with a pedicel up to 10m long-

Bovistella echinella (Pat.) Lloyd

Plate 114

Plants small, subglobose, 5-8 mm. thick, attached by a short, blunt, sandy pad;
when young, white with a thin, continuous, felt-like or granular cortex, at maturity
becoming dark brown and papery with scattered white flecks, in age practically de-
corticated. Mouth apical, definite, becoming fimbriated or toothed. Mature gleba
olivaceous brown, glebal chambers minute and irregular; subgleba practically none.

Spores (of plant from Lapland) subspherical, 4.2-5.5//, apparently smooth but with
radial lines showing in the wall, furnished with slender, tapering pedicels up to 8.5//
long. Capillitium of separate units, but threads long and slender and less frequently
branched than in most species of this genus or of Bovista; main threads about 5.5//
thick.

This little species has been collected in a number of widely separated places. The
original collection was from Ecuador, and we have specimens from North Dakota and
Lapland. Dr. R. E. Fries has also reported it from Lapland, and Lloyd reports it from
Michigan. This last is the only record, so far as we know, from the eastern United
States.

We have examined a part of the original collection from Ecuador, kindly sent us by
Patouillard himself, and find it to be just like our collection from Lapland. The capil-
litium is of long, slender branched units with main threads only about 6.5// thick; spores
subspherical, 4-5.2 x 4.8-5.8// with slender pedicels up to 12.8// long, the walls of the
spores showing faint radial lines. Patouillard describes the spores as echinulate, but
we see only faint lines with no projection in optical section.

Lloyd describes from Massachusetts Bovistella Davisii (Myc. Notes, p. 286, pi.
89), which from the description can hardly be distinguished from the present species.
We have not seen an authentic specimen.

I.YCOPERDACEAE 97

Illustrations: Fries, Th. (". E. Sveriges Gasteromyceter, fig. 21.
Lloyd. Myc. Works, pi. 89, figs. 1 and 2.

North Dakota. Anselm. On mossy sod in sand hills. Brenckle and Stevens, colls. (N. Y. B. G.

Herb, and U. X. C. Herb.). Spores 3.6-4.4/1, with a slender pedicel up to 8.2m long- Capil-

litium of long, branched units, about 4.8^ thick in largest places, olivaceous under the

microscope.
Michigan. Reported by Lloyd.
Swedish Lapland. Abisco. On grassy railroad embankment, summer, 1921. Coker, coll. (U. N.

C. Herb.).

BOVISTA Pers.

Plants spherical, attached centrally at the base, the outer peridium thin, fragile,
flaking off at maturity and leaving exposed the smooth, thin, distended, metallic-looking
inner peridium which opens by an apical pore, becomes detached from the ground, and
is blown about like a light ball, puffing out the spores. Capillitium not connected
with the peridium, but composed of separate units which branch dichotomously or
irregularly and end in tapering points. There is no sterile base. Spores dark brown
or purplish brown, in one of our species with a well developed pedicel.

The genus is remarkable in its tumbling habit (shared by Mycenastrum and to a
less extent by Disciseda*) and the inner peridium is especially adapted to this end by the
retention of its shape as an inflated ball and by its firm, parchment-like, resilient tex-
ture. These balls are rolled about by the wind and the spores shaken out by degrees
through the apical mouth or slit. They are very durable and in mountain pastures old
plants of the preceding year may still be found in good condition.

Lloyd considers B. montana Morgan the same as B. pila.

Literature

Bonorden. Die Gattungen Bovista, Lycoperdon u. ihr Bau. Bot. Zeit. 15: 593, 609, 625. 1857.

Cunningham. The Genera Bovista and Bovistella. Proc. Linn. Soc. N. S. W. 50: 367, pi. 37. 1925.

Lloyd. The Bovistae. Myc. Notes No. 12: 113. 1902.

Massee. A Revision of the Genus Bovista. Journ. Bot. 26: 129. 1888.

Morgan. Journ. Cin. Soc. Nat. Hist. 14: 144. 1891.

Petri. Flora Italica Cryptogama (Gasterales), Fasc. 5: 60. 1909.

Rolland. Observations sur le Mycenastrum corium Desv. et sur le Bovista plumbea Pers. Bull. Soc.

Myc.Fr.22: 109. 1906.
Tulasne, L. R. and Ch. De La Fructification des Scleroderma compared a celle des Lycoperdon et des

Bovista. Ann. Sci. Nat., 2nd. ser. 17: 5, pis. 1 and 2. 1842.

Key to the Species

Fruit bodies above ground, of moderate to large size

Spores spherical with a short pedicel B. pila (p. 97)

Spores oval with a long pedicel B. plumbea (p. 99)

Fruit bodies subterranean until exposed by rains, etc., small; spores oval with a long, wavy pedicel

B. minor (p. 99)

Bovista pila B. & C.

Plates 60 and 114

Plants globose or subglobose and irregularly pinched and lobed downward, 6-9
cm. in diameter, attached abruptly below by a single, small cord which breaks at

• Disciseda does very little tumbling, the flat shape with one side heavier not adapting it well to that habit. It is, how-
ever, detached from its place of growth at maturity and knocked about by rain.

98 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

maturity; outer peridium thin (0.25 mm.), pure white, tending to become alutaceous
or pinkish when handled, surface delicately furfuraceous, becoming rivulose, flaking off
in patches at maturity; inner peridium when exposed smooth and shining, more or less
bronze colored and splotched with blackish or gray areas, the effect being quite metallic-
looking like burnt copper or tin, soon opening at the top by an irregular pore or by more
extensive fissures and flaps.

Spores deep brown, thick-walled, smooth, spherical with a very short pedicel or
mucro, 3.5-4.3/x thick. Capillitium dark purplish brown at maturity, much branched;
branches long, gradually tapering to a point; main hyphae mostly about 11/u thick,
rarely up to 13/u. Immature gleba white, showing very small irregular chambers
under a lens; basidia (of No. 7999) short-clavate, 8-10.5 x 14— 18/x, with 4, rarely 3,
unequal sterigmata 4-7. 4/i long.

The co-type of B. pila in the Curtis Herbarium agrees with the usual interpretation.
The spores are spherical, smooth, 3.7-4.5/x thick. This species seems to be the American
representative of B. nigrescens of Europe, and Lloyd seems to be right in concluding
that the latter does not occur in this country. Early records in this country, as by
Schweinitz and Curtis, have since proved to be based on B. pila or B. plumbea. The
two species, B. pila and B. nigrescens, can easily be distinguished by the spores (as
noted by Hollos and Lloyd), the latter having somewhat oval spores with a long pedicel.
(In a specimen from Bresadola, Cavelante, August, 1898, the spores are 3.8-4.8 x
4-5. 2yu, with a long tapering pedicel.) The spores of B. nigrescens (pi. 114, fig. 12)
are much more like those of B. plumbea, which are oval with a long pedicel, but those
of the latter that we have seen are distinctly larger. Th. Fries gives the spores of B.
nigrescens as 5-6/u in diameter. If this variation exists, the plants should hardly be
separated. The spores of Schweinitz's "B. nigrescens" are those of B. pila, spherical,
usually sessile, sometimes with a short mucro, 3.7-4.5/j thick.

Illustrations: Lloyd. Genera of Gasteromycetes, pi. 9, figs. 43 and 44.
Lloyd. Myc. Works, pi. 2.

North Carolina. Black Mountain. At summit on grassland frequented by cattle. Holmes, coll.

(U. N. C. Herb., No. 5359).
Boone. Coker, coll. Aug. 22, 1922. The photograph shows this collection but the plants were

lost.
Asheville. Beardslee.
Haywood Co. Piney Mountain (alt. 5700 ft.), open pasture, July 31, 1926. Coker, coll. Also

Cold Spring Mountain, by barn, immature state, Aug. 3, 1926. Totten, coll. (U. N. C.

Herb. No. 7999).
Virginia. Apple Orchard Mountain. Murrill, coll. (N. Y. B. G. Herb.).
Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.). Spores smooth or nearly

so, 3.8-4.4m, with a pedicel and very distinct oil drop. Capillitium much branched, main

threads up to 17^.
New York. Arkville. Murrill, coll. (N. Y. B. G. Herb.).
Vermont. Middleburg. Burt, coll. (Bres. Herb.). Spores spherical, usually with a short mucro,

3.7-4.4/*.
Wisconsin. Madison. Trelease, coll. (N. Y. B. G. Herb.).

Algonia. Dodge, coll. (Univ. Wis. Herb, and U. N. C. Herb.). Spores smooth, 3.7-5/1, often

with a short pedicel.
Wyoming. Pitchfork. Davis, coll. (U. N. C. Herb.). Spores smooth or nearly so, 3.8-5/1, with a

short pedicel.
Washington. Langley. Grant, coll. (U. N. C. Herb.). Spores subspherical, smooth, 4-5.5ju, with a

short pedicel. Main threads of capillitium up to 12/i thick.

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LYC0PERDAC1 U 99

Canada. London. Pcarncss, coll. (U. N. C. Herb.). Spores smooth, spherical, 3.8-4.5/1, with no
obvious pedicel.

Bovista plumbea Ters.

Plate 114

Plants small, 2-3 cm. thick, nearly globose and attached by a clump of fibrous
mvcelium; cortex thin, whitish, at maturity shelling off in plates over the upper half
or over most of the surface. Inner peridium thin, smooth, mouse gray to deep mouse
gray of Ridgway, dehiscing at the apex by a rather large, nearly circular mouth.

* Spores (of No. 5358) oval with long, pointed, hyaline pedicels up to 13^ long, very
minutely rough, with a halo around the spore; wall two-layered, outer layer thinner and
dark, inner one thicker and lighter, 4.5-6 x 5.5-7^, with one large oil drop. Capillitium
ochraceous brown, loose, the threads several times branched, main stems 16-22^ thick,
branches tapering to a point. According to Rolland (see below), the basidia are
4-spored with rather long sterigmata.

Our plant agrees well with Morgan's description. Bovista ovalispora Cooke and
Massee is probably the same plant (Ann. Bot. 4: 62. 1889). Lloyd states (Myc.
Notes No. 12: 115) that in B. plumbea "most spores are not truly globose, but have a
tendency to oval form. There is, however, a wide range in this respect." See under
B. pila for comparison with B. nigrescens.

Illustrations: Berkeley. Ann. Sci. Nat., 2nd ser., 12: pi. 2, figs. 15-18. 1839.
Corda. Icon. Fung. 5: pi. 6, fig. 47. 1837.

Dufour. Atlas des Champ. Comest. et Ven., pi. 75, No. 171. 1891.
Hard. Mushrooms, fig. 472.
Lloyd. Myc. Works, pi. 1.
Massee. Ann. Bot. 4: pi. 4, fig. 63.
Rolland. Bull. Soc. Myc. Fr. 22: pi. 22, figs. 12-15. 1906.
Sorokine. Rev. Myc, 1890, pi. 12, figs. 342, 343; pi. 25, fig. 352.
Tulasne. Ann. Sci. Nat., 2nd ser., 17: pi. 2, figs. 1-3. 1842.

North Carolina. Asheville. Beardslee. (U. N. C. Herb., No. 5358.) Spores oval, with faintly
striated walls, 4.5-6.5 x 5.5-8/*.

South Carolina. Ravenel. Fungi Car. Exs. No. 81, as B. nigrescens. (Phil. Acad. Herb.)

New York. Albany. Peck, coll. (Curtis Herb.). Also others from New York as B. nigrescens.

Massachusetts. Boston. Sprague, coll. (Curtis Herb., as B. nigrescens).

Ohio. Hayden, coll. (Curtis Herb., as B. nigrescens).

Cincinnati. Lloyd, coll. (Bresadola Herb, and U. N. C. Herb.). Spores dark, oval, faintly rough,
4.5-6.8 x 5.5-7.4/1, with a pedicel up to 13/i long. Capillitium as usual, dark, thick-
walled, up to 22m thick.

Wisconsin. Blue Mounds. (Univ. Wis. Herb, and U. N. C. Herb.) Spores oval with long tapering
pedicels, 4.2-5.2 x 5.5-7/1.
Eagle Heights. (Univ. Wis. Herb, and U. N. C. Herb.) Spores oval, with walls faintly lined,
4.8-6 x 6-7.5/1, with pedicels up to 16/i long.

Wyoming. Pitchfork. Davis, coll. (U. N. C. Herb.). Spores oval, minutely rough, 4-5.5 x 5.5-7/1,
with long pedicels.

Bovista minor Morgan

Plate 114

Plant subglobose, about 1.5 cm. broad (1.3-1.8 cm., according to Morgan), sub-
terranean until exposed by rain, etc., the surface at first entirely covered with earth

100 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

held on by a rather inconspicuous mycelial felt, this wearing away slowly and exposing
the pale grayish, then reddish brown, flaccid inner peridium, except for a basal remnant
as an earthy point or disc, opening tardily by a small, apical mouth. Gleba olivaceous,
then brownish; subgleba none.

Spores (of No. 2416) oval, smooth, 3-3.7 x 3.8-5/*, with long, wavy pedicels up to
15// long. Capillitium of branched units as in its relatives; main threads up to 22^u
thick, with thick walls.

This rare plant was described by Morgan from Ohio and Nebraska and is doubtfully
reported from Massachusetts by Lloyd. We have drawn the above description from
two Canadian plants noted below and kindly sent us by Dearness. One of these plants
is still covered all over with the thin sand case except for spots here and there; the other
is clean of dirt, the cortex remaining only as a basal disc. This last has the inner per-
idium deep red-brown, while in the former it is grayish where exposed. The spores and
capillitium in both plants are identical. The thin, papery peridium would seem to
indicate a close approach to the genus Bovistella.

Hollos (1. c, p. 126) reduces both this species and Bovistella dealbata Lloyd (Myc.
Notes, p. 86) to Bovista tomentosa (Vitt.) De Toni of Europe. We have examined an
authentic specimen of Bovistella dealbata (pi. 61 and pi. 114, fig. 14) and find it quite
different from the plant described above. It is an epigean species exposed throughout
the entire growth of the fruit body and in this way differing sharply from what we are
taking to be B. minor and also from Morgan's description of that species. The spores
of the Lloyd specimen are larger than in the Dearness plants and are as Hollos said
furnished with minute warts (x 2160). They are spherical to subspherical, 4.3-5.5 x
5-6. 3p (rarely 6.8//), with pedicels 6-12. Su long, usually about 8ju long. Capillitium
threads up to 10/j. thick, moderately branched into long, slender, tapering branches.
We have also examined a specimen of B. tomentosa from Bresadola's herbarium from
Andola, collected and determined by him, and find that it agrees well with Hollos's
description and figures except that the spores are practically smooth (3.5-4.2 x 4-5. 5/u).
The plants are the same size as B. dealbata, but seem to be darker at all stages, the spores
are somewhat smaller, and the capillitium threads are thicker and more branched.
Without further evidence we prefer to keep the two species separate. As the European
species is obviously epigean we cannot see how it could be Bovista minor. We have
not seen an authentic specimen of the latter, but as both Morgan's (Journ. Cin. Soc.
Nat. Hist. 14: 147) and Lloyd's (Myc. Notes, p. 117) descriptions agree in every respect,
we use the name for the Canadian plants.

Plants in the U. S. National Museum from Nevada de Toluca, Mexico, determined
by Lloyd as B. brunnea Berk., are, in appearance, spores and capillitium, like those of
specimens of Bovistella dealbata from the Lloyd Herbarium, sent us by Lloyd. Bovista
brunnea Berk, was originally described from New Zealand, and plants from that country
sent us by Cunningham under that name are distinctly olive-brown in the immature
condition and otherwise look somewhat different from the others. The spores and
capillitium are, however, the same. Lloyd thinks that B. brunnea is the same as B.
tomentosa and not the same as B. dealbata.

Illustrations: Lloyd. Myc. Works, pi. 3, figs. 3-5.

Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 5, figs. 10-12.

Canada. London. On clay bank by spring. Dearness, coll. (Dearness Herb, and U. N. C. Herb.
No. 2416).

PLATE 61

BOVISTELLA DEALBATA. LLOYD HERB. [above]

Crucibllum vulgare. No. 1534. Magnified

LWOPERDACEAE 101

MYCENASTRUM Desv.

Fruit body epigcal, breaking from its attachment after maturity; peridium thick
and tough, splitting irregularly above. Capillitium of short, free threads which are
furnished with short prickles. Spores brown, spherical (said to be sessile and 2-4 to a
basidium in If. radkattm). According to Hollos there is only one species, though a
good many have been described.

The genus is easily distinguished from all others here included by its prickly
capillitium.

Literature

Desvaux. Sur le genre Myccnastrum, du groupe des Lycoperd£es. Ann. Sci. Nat., 2nd. ser., 17: 143.
1842.

Mycenastrum corium Desv.
M . spimdosum Peck

Plates 62 and 114

Plants seated on the ground, subglobose, depressed or obovate, often irregular,
more or less plicate or pitted below and attached by a point to the fibrous mycelium;
size various, usually about 4-10 cm. thick; cortex a soft, felted, whitish coat which
dries down irregularly and wears away slowly, parts remaining here and there as inherent
scales; inner peridium deep brown on surface and in section, about 2 mm. thick, tough
and corky, splitting tardily from above downward into unequal and irregular flaps
which may separate slightly or at least open widely. Gleba without a sterile base,
dark brown to purplish brown at maturity, not very powdery.

Spores (of a plant from the Lloyd Herbarium) dark brown, spherical, thick-walled,
minutely but, under high power, distinctly marked with a more or less perfect reticulum
of linear warts, 8.5-1 ly. thick. Capillitium of separate, short threads with a few short
branches, and peculiar in the thorn-like spines with which they are more or less
thickly set.

This species is abundant in the west, but rare in the eastern states. Lloyd reports
it from Illinois (Myc Notes, p. 120) and from Florida (Letter 62, note 441). (See
also Myc. Notes, p. 1173, where an error is corrected.)

The plant much resembles a Scleroderma in general appearance, but it is easily dis-
tinguished by the spiny capillitium threads. It differs also in its habit of breaking
loose from its attachment when mature and adopting a tumbling habit like Bovista.
The separate threads arising freely in the gleba also recall Bovista. For numerous
synonyms see Hollos.

Illustrations: Cunningham. Trans. N. Z. Inst. 57: pi. 9, figs. 25 and 26. 1926.
Fischer. Engl. & Prantl. Pflanzenf. I1: fig. 165 B.
Hollos. 1. c, pis. 24-26; pi. 29, fig. 31.
Lloyd. Myc. Works, pi. 5, figs. 1-11.
Lloyd. Photogravure of Am. Fungi, No. 18.
Morgan. Journ. Cin. Soc. Xat. Hist. 14: pi. 5, figs. 13-14.
Petri. 1. c, fig. 35.

Sorokine. Rev. Myc, 1890, pi. 25 (101), fig. 354 fas var. Kara-Kumianicm); pi. 27 (103), fig. 365;
pi. 29-30 (105) (figs. 371-372 as var. Kara-Kumianum).

102 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

New Jersey. Newfield. Ellis, coll. (N. Y. B. G. Herb., as M. spinulosum). Spores dark, reticulated,
9.3-11. 2/i. Capillitium threads of branched, spiny units about 12. 5m thick in main
threads.

Wyoming. Pitchfork. Oct. 1912. Simon Davis, coll. (U. N. C. Herb.).

Canada. Eastern Ontario. Dearness, coll. (N. Y. B. G. Herb.).

Also specimens seen from Kansas, Nebraska, North Dakota, Colorado, Texas.

GEASTER Mich.

Plants at first subglobose and with or without an apical beak; either just buried
in the soil, trash, or moss they grow in and with or without a fine flocculence all over
which binds the substratum to the surface, or in some species seated on the surface of
rotting wood, trash, or earth and attached only at the base (epigeal). Outer peridium
at maturity splitting from the apex into few or many rays which expand more or less in a
stellate way, or even recurve strongly and are composed typically of three layers, the
surface one a delicate, interwoven coat of flocculence or tomentum that in some species
is scarcely obvious, but in others forms a more or less perfect membrane which remains
attached to the next layer on expansion, or becomes more or less detached from it,
and in some cases is completely stripped off and remains as an empty sac in the ground
when the rest of the plant is lifted out by the recurving lobes of the next layer. This
next inner layer, called the fibrous layer, is thicker and tougher and is formed of closely
woven, homogeneous fibers. Lining this layer and splitting with it is a third layer of
turgid, fleshy tissue which dries out and shrinks after opening, either remaining attached
to the inner side of the fibrous layer as a continuous or much cracked cartilaginous-
looking membrane, or peeling off and disappearing in places or throughout. Sometimes
a part is separated and remains as a kind of collar around the center. For the micro-
scopic structure of peridia, see under G. velatinus. Inner peridium (spore sac) thin,
tough, membranous, pliable, stalked or sessile, opening by a single, apical mouth that is
simply a small torn hole or an elevated pore which may or may not be surrounded by a
circular area (peristome) of different texture, and more or less sharply circumscribed
outline. In one group of species this area is prettily and regularly fluted (sulcate).
Gleba pure white when young, composed of tramal fibers and plates which radiate
straight from the columella to the peridium, and which enclose the very narrow, much
elongated, almost tubular cavities which are lined with the basidia. The glebal
chambers are not visible in longitudinal section and barely so in cross section. Basidia
thick, elliptic to pyriform, borne on very slender hyphae, 4-8-spored. Spores spherical
or subspherical, usually warted or asperulate. Capillitium abundant, composed of
simple, rarely-branched threads which arise from the walls and also from the more or
less prominent (in some species vague or absent) columella.

An interesting genus of numerous species that are popularly known as "earth stars."
The dry spores puff out of the mouths as in puffballs, leaving the persistent capillitium
behind. A peculiar, long-stalked species (G. stipitatus Solms) is recorded from Java
(Fischer, Hedwigia 32 : 50, pi. 5. 1893). Before opening it has much the appearance
of Lycoperdon muscorum but has a short point. Another peculiar plant should be men-
tioned here, although not found in America. It is Trichaster melanocephalus from
Europe. It is like a large Geaster fornicatus in general appearance except that the inner
peridium wall very soon disappears. It differs also in the very large, globose, hard
columella around which the capillitium forms a matted mass filled with spores. There
is a specimen in Washington under the name of G. fornicatus (Myc. and Path. Herb.;
Petrak, Fl. Bohemiae et Moraviae exs. No. 1498). It is just like Lloyd's figures
(Myc. Notes, p. 189, pi. 17) and has all the characters of the species. The spores are
about like those of fornicatus, distinctly warted, 3.8-4.5/n. The plant is said to be quite

PLATK 62

Mycenastrum cojuum. Lloyd Herb, [above]
Pitchfork, Wyoming [below]

LYCOPERDACEAE 103

epigean, and this is borne out by the plant just mentioned, as the outer layer has not
been stripped off and is furthermore free of trash.

In attempting to find a natural classification for the species of Geaster, one meets
with difficulties and with differences of opinion among authors. What are the char-
acters expressing the most fundamental relationships; and those of lesser degrees of
importance? All arrangements heretofore proposed have in certain sections obviously
violated, it seems to us, the true relationships. De Toni (Rev. Mycol. 9: 64. 1887.
See also Sacc. Syll. 71: 70, and Hollos, I.e., p. 51) divides the true Geasters first into a
fornicate and a cupulate group (Fornicati and Cupulati) and the remaining species which
are subdivided into the sulcate-striate group (Striati) and the fimbriate group (Fim-
briati). As a result of the first two groupings there are thrown together the very dis-
similar fomicatus and radicans, while the near relatives fomicatus and rufescens, and
coroiiaius and minimus are separated. The third group is a natural one, but the last
should be rearranged.

Morgan's two main groups, DepeUiti and Pelliculosi, are in part natural, but are
unnatural in having species with sulcate-plicate peristomes in each. His two sub-
groups under DepeUiti are unnatural for the same reason. He does not make the
mistake of making the hygroscopic character a leading one. In Lloyd's last outline of
the Geasters (Myc. Notes, p. 317), as well as in The Geastrae, he first divides them into
the Rigidae and the Xon-rigidae, the latter with six groups. This throws together such
distantly related plants as mammosus and Drummondii and separates species with
sulcate peristomes.

Of the six groups of Lloyd's Non-rigidae, the first and last are natural in their in-
clusions, though Hariotii is excluded from the first. The other three groups are in our
opinion unnatural to a greater or less degree. That the hygroscopic character is not the
most fundamental one seems obvious. Both hygroscopic and non-hygroscopic species
are found in the very natural group with sulcate peristomes, as well as in the non-
sulcate group. In related species and even in the same species at times the intensity of
this character is very variable and in individuals may practically disappear. For
example, in arenarius the rays while hygroscopic are usually pliable and thin, while in
the very closely related floriformis they are much more rigid. In asper individuals
appear rarely without any hygroscopic tendency at all, though the species is markedly
hygroscopic.

Within certain limits the structure of the mouth area is the most dependable
character. The sulcate group is a natural one and there are no confusing intermediates
connecting it with the non-sulcate group. So-called intermediates like Morganii are
not troublesome, as their crumpled mouths rarely bear any real resemblance to a truly
sulcate peristome. In the non-sulcate group the distinction between the definite
and indefinite peristome, while adequate enough as a rule, is in some cases obscured by
variation. This is particularly true in floriformis and arenarius. In constructing the
key we have used the characters in the order of their importance as it seems to us, and
the key is therefore as natural a grouping of the species as we can arrange at present.

Literature

Cunningham. Species of the Genus Geaster. Proc. Linn. Soc. New South Wales 51: 72, pis. 2-7.

1926.
Cunningham. The Development of Geaster velutinus. Trans. Brit. Myc. Soc. 12: 12, 9 text figs.

1927.

104 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Destree, Caroline. Revision des Geaster observes dans les Pays-Bas. Neederlandsch Kruidkundig

Archief, 2nd. ser., 6: 488. 1895.
De Toni. Revisio monographica generis Geasteris, Mich. Rev. Myc. 9: 61, 125, pi. 2. 1887.
Lloyd. The Geastrae, figs. 1-80. Cincinnati, 1902.

Lloyd. New Notes on the Geasters. Myc. Notes No. 25: 309, figs. 144-149 and pis. 94-101. 1907.
Morgan. The North American Geasters. Amer. Nat. 18: 963, figs. 1-12. 1884.
Morgan. New American Geasters. Journ.Myc. 1: 7. 1885. Condensed from above paper.
Morgan. Journ. On. Soc. Nat. Hist. 12: 12, pis. 1, 2. 1889.
Morgan. The Genus Geaster. Amer. Nat. 21: 1026, figs. 1-2. 1887.
Petri. Flora Italica Cryptogama (Gasterales),Fasc. 5: 67. 1909.
Rea. British Geasters. Trans. Brit. Myc. Soc. 3: 351, pis. 17-19. 1911.
Rick. Die Gattung Geaster und ihre Arten. Bot. Centralb. 27, Abt. n: 375, figs. 1, 2. 1910.
Scherffel. Bemerkungen iiber Geaster-Aiten. Ber. d. Deutsch. Bot. Ges. 14: 312, pi. 19. 1896.
Smith. Two species of Geaster. Grevillea 2: 35, pis. 13 and 14, fig. 1. 1873.
Smith. British Geasters. Grevillea 2: 76, pis. 13-17 and 19-20. 1873.
Tulasne, L. R. and Ch. Sur les genres Polysaccum et Geaster. Ann. Sci. Nat., 2nd. ser., 18: 129.

1842.
Woodward. An essay towards an history of the British stellated Lycoperdons. Trans. Linn. Soc,

London 2: 32, 323. 1794.

Rey to the Species

Peristome not truly sulcate, i.e., not regularly ridged and grooved throughout the extent of the peris-
tome; irregularly wrinkled and crumpled only in G. Morganii
Mouth area distinct as a silky zone of different texture and often of different color, but not outlined
by a sharp narrow groove (in G. subiculosus the mouth area is scarcely distinct) ; spore sac
without glistening particles and, if stalked, the stalk short and thick; button exposed or,
if submerged, the outer coat not holding earth or trash to any extent and the mycelium
attached only by the center of the base; expanded plant saccate or convex below or im-
perfectly to typically fornicate
Rays not hygroscopic, i. e., not incurved over or around the spore sac when dry or expanding
when wet
Outer surface smooth and glabrous or with the texture of soft leather (rarely scaly);
button with a distinct, narrow point
Outer layer of rays tending to crack into longitudinal strips or less often into irregular
plates, slightly if at all separating from the middle layer; base saccate or plane

or less often concave below; mouth not wrinkled G. triplex (p. 106)

Outer layer of rays stripping off on the central region and over the proximal part of
the rays and remaining convex below, the inner layer arching upward and
elevating the spore sac (pseudofornicate) ; mouth not wrinkled

G. Umbatus (p. 107)

Outer layer and shape of plant as in G. triplex, but outer layer tending to peel off in

irregular flakes; mouth irregularly wrinkled or crumpled (pseudosulcate)

G. Morganii (p. 109)
Outer layer soft, not cracking into strips, but with some tendency at times to peel
from the rays; mouth not wrinkled; plants small (at least in the N. American
form)

Spores 3.4-4.8ju thick G. saccatus (p. 110)

Spores 2.2-3.5/i thick Northern form of G. saccatus (p. Ill)

Outer surface pale buff, nearly or quite glabrous, with the texture of leather; peristome
scarcely distinct; dry button blunt, with a low broad umbo, or the top umbilicate;

spores smooth, 2-3/x G. subiculosus, sense of Lloyd (p. 118)

Outer surface not glabrous, but tomentose, strigose or spongy; button not pointed

Plants medium to large; outer surface minutely felted-tomentose, rather harsh, pale

Expanded plant saccate; spores 2.5-3.3^ G. velutinus (p. 113)

Expanded plant fornicate; spores 3.8-4.8^ G. radicans (p. 115)

LYCOPERDACEAE 105

Hants small, the base pinched to a point; ccspitosc or gregarious in numbers from a
conspicuous, superficial white mycelium which binds together the twigs and
leaves of the substratum ; outer surface strongly tomentose to strigose, at
least when young

Plants not saccate; spore sac and fleshy layer blackish C. Lloydii (p. 123)

Hants saccate; spore sac and fleshy layer not blackish

Spores 3.-M/J C. mirabilis (p. 116)

Spores 3-3.6/1 C. trichifer (p. 1 18)

Rays hygroscopic

Plants of medium size; outer pcridium clean of dirt even in the button

G. mammosus (p. 1 19)

Plants small, covered with sand all over at first G. arcnarius (p. 120)

Mouth area not distinct, at least without a definite boundary (at times the color is paler or darker,
and a depressed zone may be present in G. floriforms); button quite submerged; outer
surface covered with flocculent mycelium holding trash or sand; plants saccate or fornicate
or intermediate
Rays not hygroscopic

Spore sac asperate with stiff hairs G. Hicronymi (p. 123)

Spore sac not asperate

Outer peridium with center elevated to expose the spore sac, which is short-stalked or
sessile

Mycelial (outer) layer adhering to the center and rays G. rufescens (p. 121)

Mycelial layer remaining in the ground and forming a hollow cup with the
strongly reflexed rays attached to its margin (fornicate)

G. fornicates (p. 124)
Outer peridium saccate and enclosing the sessile spore sac like a bowl

G. fimbriates (p. 125)
Rays hygroscopic

Surface of spore sac spongy See Astraeus (p. 185)

Surface of spore sac not spongy

Spores about 3.5-4.5/1 A form of G. arenarius (p. 121)

Spores about 5-7/i G. fioriformis (p. 127)

Mouth area very distinct, silky and outlined by a narrow groove; buttons quite submerged until
dehiscence, and the outer layer covered with earth or trash; expanded plants concave
below and often truly fornicate; spore sac with a slender stalk and covered when fresh
with minute, glistening particles; rays not hygroscopic
Plants of small to medium size, growing on the ground or on humus; spores dark brown

G. coronates (p. 129)
Plants small to very small, growing on mossy trunks of living trees; spores very pale

G. leptospcrmus (p. 131)
Peristome truly sulcate with regularly arranged ridges and grooves composing the peristome; all U. S.
American species (at least) with buttons submerged until dehiscence
Not hygroscopic, the rays bent backward, the base concave below; spore sac stalked or sessile
Peristome not black

Plants of medium size, stalk long, lower part of spore sac often striate radially; spores

about 4.2-5.5/1 G. pectinatus (p. 132)

As above but spore sac not striate and having a flaring collar pendent around its lower

side G. Bryantii (p. 133)

Plants small; stalk short; spores about 3.7-1.4/1 G. Schmidclii (p. 134)

Peristome black or nearly so; spore sac sessile C. Hariotii (p. 135)

Hygroscopic, the rays bent inward (typically) when dry, spreading when wet

Spore sac covered with rather coarse, wart-like particles, short-stalked G. asper (p. 135)

Spore sac minutely furfuraceous, smooth in age, sessile

G. umbilkates, sense of Morgan (p. 136)
Spore sac minutely spiny to pulverulent-granular, sessile G. Drummondii (p. 137)

106 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Geaster triplex Jung.

G. Michelianus W. G. Smith

Plates 63 and 115

Plants large, pointed before opening, the point up to 12 mm. long, the inner perid-
ium subspherical, up to 2.7 cm. thick, quite sessile, gray-brown when fresh, becoming
reddish brown with age; mouth fibrillose and becoming lacerated, surrounded by a more
or less definitely outlined, broadly conical area which may or may not be paler. Outer
peridium splitting into 6-8 segments with long acuminate tips which expand or become
revolute under the convex, flat or less often arched base; fleshy layer thick, in our plants
mostly remaining intact as a moderately thick, reddish brown, cracked crust on the
fibrous layer, the central region in some plants breaking loose from the part on the rays
to form a broad collar or cup around the inner peridium; outer layer usually nearly free
of dirt or trash, but with a few small particles sticking to any part, firm, glabrous, dull
yellowish, mostly adnate, but cracking into radial strips or irregular areas; mycelium
basal and leaving a scar at point of attachment when the plant is lifted ; button supposed
normally to be sunk in the substratum, but its surface does not hold the trash as in G.
fimbriates, G. rufescens, G. fornicatus, and most other subterranean species (for rare
exception see note under Alabama entry below). Columella about 1 cm. long, clavate,
and persistent.

Spores (of No. 5886) deep smoky brown, distinctly warted, spherical, 3.7-4.4/u,
most about 3.9/t thick. Capillitium threads unbranched, about 5,u, up to 6/j thick,
not sinuous, concolorous with the spores.

Geaster lageniformis, as suggested by Morgan (see Lloyd, The Geastrae, p. 38),
is a small form of G. triplex. The cracked surface, definite mouth, pointed button, and
spores are all the same. A typical-looking plant of G. lageniformis from London,
Canada (N. Y. Bot. Gard., labelled G. saccatus), has spores distinctly verrucose,
3.5-4.2yu. Another from the same place and in the same herbarium is Ellis & Ev.,
N. Am. Fungi, No. 2735 (as G. triplex). Our plants are like the G. triplex of Thiimen's
Mycotheca Universalis No. 1410 from Holland, which has spores distinctly papillate,
3.6-4.2,u. Plants from Hungary at the New York Botanical Garden correctly labelled
G. lageniformis are the same, with spores verrucose, 3.4-4/* thick. Similar plants from
Hungary also labelled G. lageniformis are in the Herbarium of the Museum of Paris.
Another example from Charlottenburg in the same herbarium (P. Sydow, coll., as G.
fimbriatus) has spores distinctly verrucose, 3. 7-4.5yit. In American herbaria one often
finds under G. lageniformis plants that we are referring to G. saccatus, northern form.

Illustrations: Clements. Minnesota Mushrooms, fig. 89.

Cunningham. Trans. N. Z. Inst. 57: pi. 11, figs. 35, 36.

Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 2, figs. 3, 4.

Destree. I.e., pi. 9, fig. A.

De Toni. I.e., pi. 63, fig. C.

Hard. Mushrooms, pi. 66.

Hollos. I.e., pi. 11, figs. 1-6 (these figs., if correct, represent a very coarse form); pi. 29, fig. 17.

Lloyd. The Geastrae, figs. 47^9. Also pi. 94, figs. 1-5.

Massee. Ann. Bot. 4: pi. 1, fig. 27 (as G. Michelianus).

Micheli. Nova Plant. Gen., pi. 100, fig. 1.

Morgan. Amer. Nat. 18: fig. 2. 1884.

Illustrations of the form G. lageniformis:

Hollos. I.e., pi. 10, figs. 13 and 14. Figs. 11 and 12 more nearly represent what we are calling

G. triplex.
Lloyd. The Geastrae, figs. 76 and 77.

PLATE 03

Geaster triplex. Xo. 682

LYCOPERDACEAE 107

Smith. C.rrvilloa 2: pi. 20 (as G. saccatus).
Vittadini. Monog. Lycopcrd.. pi. 1. tig. 2.

682. Chapel Hill. December 1908. Old collection with no other data.
5886. On mossy oak stump, October 17, 1922.
7483. Under cedars, Aug. 7, 1924.

Asheville. Beardslee.

South Carolina. Ravenel. (Curtis Herb., as G. fimbriates, No. 3025, and as G. saccatus, No. 1600;
Tath. and Myc. Herb., as G. saccalus, Fung. Car., No. 77.) Another folder of this No. 77
is a smaller plant with softer felted outside and tapering lobes and looks like our G. saccalus
from Asheville.
Florida. Alachua Co. Weber, coll. Spring of 192-1. (U. N. C. Herb., No. 7445). Also collection by
Walker.
Gainesville. Weber, coll. (U. X. C. Herb, and Univ. Florida Herb.). Lageniformis form.
Spores distinctly warted, 4-5. 5m thick, often with hyaline material. Also another collection
which is remarkable in its thin, soft, papery peridium.
Alabama. Spring Hill. Bertolet, coll. (Lloyd Herb.). Some of these have clean surfaces, while in

others the surface is as dirty and trashy as in G. rufescens. Mouth very definite.
Maryland. Smith, coll. (U. S. Nat'l. Herb., as G. saccatus).

New Jersey. Pennsville. Sept. 1891. Ellis and Everhart, N. Am. Fungi, No. 2736. (N. Y. Bot.
Gard. Herb., as G.fimbriatus.) This is good G. triplex. (Path, and Myc. Herb., Washington,
as G. striatus.)
Pennsylvania. New Garden. (Michener Herb., as G. rufescens.)

Ohio. Loveland. James, coll. 1880. (Path, and Myc. Herb., as G. saccatus.) Outer surface mottled
as in Rick, No. 261.
Cincinnati. Spurlock, coll. (Path, and Myc. Herb., as G. lageniformis). Also collection by
James in the same herbarium.
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).
Missouri. St. Louis. Trelease, coll. (Farlow Herb.). Lageniformis form.
Wisconsin. Algonia. Dodge, coll. (Unix-. Wis. Herb, and U. N. C. Herb.).

Madison. Trelease. coll. (Farlow Herb.). A small, papery form.
Washington. Seattle. Morrill, coll. (N. Y. Bot. Gard. Herb., No. 208). With a long single root

2-3 mm. thick, attached to center of base and branched below.
Canada. London. Deamess, coll. (Path, and Myc. Herb., U. N. C. Herb., also Ellis and Everhart,

X. Am. Fungi, No. 2735 in N. Y. Bot. Gard. Herb.).
Mexico. Colima. Murrill. January 1910. Several buttons show a small point with top collapsed
around it; surface hard and glabrous, in some cases with flat, more or less inherent scales like
Scleroderma vulgaris, nearly clean; peristome distinct. Such a form is shown by Lloyd, Myc.
Notes, p. 339, fig. 166.
South America. Paraguay. Balansa, coll. (N. Y. Bot. Gard. Herb., unnamed). Spores 3.7-4.2M.
Brazil. Rick, coll. Fungi Austro-Americani, No. 261. (Path, and Myc. Herb., Washington.)
Exactly like G. triplex except that the outer surface is cracked and mottled to remind one of a
giraffe, a form closely approached by the plant from Ohio cited above. Spores 3.5^1. 1m
thick, surface as in G. triplex. Also 4 plants from Rick. (U. N. C. Herb.)
Venezuela. Fendler, coll. (N. Y. Bot. Gard. Herb., asG. saccatus). Looks exactly like G. triplex.
A large plant with spore sac 2.5 cm. thick; rays long and taper-pointed. Spores distinctly
warted, 3.7— 1.4m thick.
England. Surrey and Ascot. (N, Y. Bot. Gard. Herb, from Massee Herb., as G. Michelianus.)
New Zealand. Cunningham, coll. (U. N. C. Herb.). Spores distinctly warted, 4.2-5.5M thick.

Geaster limbatus Fr. Sense of Bresadola.

Plates 64, 115 and 116

Plants large and resembling G. triplex in most important characters. The follow-
ing description refers to the dried plants: Outer peridial surface firm, glabrous, nearly

108 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

clean of trash except at base, mycelium attached only at a central point below; rays
reflexed, the long tapering tips usually revolute; outer layer not cracked into strips or
flakes, but separating as a rule from the central region and the proximal part of the rays,
and remaining convex below while the inner layer arches upward and elevates the spore
sac (pseudofornicate). Spore sac subglobose with a more or less obvious apophysis,
brown, nearly glabrous and with a short, thick, often flattened stalk; peristome definite,
silky, often paler than the sac, broadly conical, the mouth fimbriate. Fleshy layer
brown, rather thick, mostly adnate and not cracking. Columella subspherical, about
4—5 mm. thick.

Spores (of plant from Wisconsin) spherical, distinctly warted, 3.7-4.8/*, most
about 4fi thick. Capillitium threads straight, not branched, concolorous with the
spores, up to ly. thick.

This plant is evidently nearly related to G. triplex as shown by long-pointed lobes
(indicating a pointed button), absence of flocculence on the outer surface, which is clean
and glabrous in large part, and in the definite mouth and similar spores. It differs
from G. triplex in the splitting away of the outer layer over the central region and the
arching up of the middle layer. The stalk is probably no more obvious than it would be
in G. triplex if it were to assume such a position.

There is much confusion among authors and in herbaria as to the identity of G.
limbatus. This is the only American plant labelled G. limbatus in American herbaria
which cannot be referred to G. triplex or G, rufescens or G. pectinatus. Most of the
collections at the New York Botanical Garden labelled G. limbatus are G. rufescens, e.g.,
from Alabama, Kentucky, Ohio, Missouri, Indiana, Colorado and Canada. All the
material in the Path, and Myc. Herb, at Washington as G. limbatus is G. rufescens, as
Ellis, N. Amer. Fungi, No. 1309, and others from Ohio and Colorado. Morgan figures a
plant as G. limbatus (N. Amer. Geasters, fig. 6) which is probably G. rufescens, as noted
by Lloyd and Hollos. A plant in the Curtis Herbarium from Alabama as G. limbatus
(Peters, No. 934) is G. rufescens with a short stalk. We are adopting the name of G.
limbatus for the present species as it is so determined by Bresadola, as shown by a
European specimen from him in the Lloyd Herbarium. This has exactly the habit of
the American plant described above, the outer surface clean and separated from the
fibrous layer over the center and proximal part of the rays. Another plant from Bresa-
dola so labelled has the outer surface earthy and not separated in part. The mouth is
distinct in both. Our plants described above are also the G. limbatus of Lloyd, at least
in so far as the American specimens are concerned, as shown by a number of plants in
his herbarium. Among Lloyd's illustrations of G. limbatus (The Geastrae, figs. 42^16),
figs. 45 and 46 represent the species as we have it. His three other figures are probably
not the same. Figure 42 of a plant from Rea (England) and fig. 44 of a plant from
Hollos (Hungary) apparently represent the dark species noted below and considered
G. limbatus by English authors and by Hollos.

It is certain that the G. limbatus of Hollos and of Massee and Berkeley and to all
appearances that of Elias Fries* is not our plant, and we have never seen anything like
it in this country. It is a large, usually very dark plant (blackish brown; some forms
are paler and glaucous) with a short thick stalk, a large, definite, fimbriate mouth of

* In the Curtis Herbarium are two plants from E. P. Fries (Upsala) labelled by him G. limbatus, but they are really of
pectinatus. Worthington G. Smith gives a figure (Grevillea 2: pi. 17, fig. 2) as G. limbatus which has all the appearance G.
G. pectinatus except that grooves are not shown on the peristome.

PLATE 64

Geaster limbatus
Burlington, Wisconsin [lower right]; Kansas, Xo. 8252 [other three]

LVC0PERDAC1 \l 109

more brownish tint and with a universal flocculent covering that holds earth tightly as in
G. rufescens. In fact, the appearance is very much that of rufescens except for the black-
ish and more glabrous spore sac and more definite mouth. It is beautifully shown in
Hussey's Illustrations, pi. 2. Good plants from Hollos in the Farlow Herbarium and
from Massee (determined by Berkeley) in the New York Botanical Garden Herbarium
are alike. The former has spores which are distinctly warted, 4-5. 2/i thick, and with
which those of the latter agree (4.2-5.5^). Another plant from Massee (Chiselhurst)
so labelled is the same. Plants from Germany (Berlin; Magnus, coll.) labelled G.
rufescens by the collector are also the same: nearly black, mouth distinct. The outer
surface of this blackish plant of Northern Europe is quite different from that of our
plant, and it does not peel off. Neither Hollos nor any other European author mentions
the separation of the outer layer in limbatus. The spores as given by Hollos are 4-5,
often 6/i thick, which is too large for our plant. Geaster pseudo-limbatus is represented
from Hollos in Lloyd's Herbarium and apparently does not agree with either idea of
limbatus discussed above.

Illustrations: Fries, Th. C. E. Sveriges Gasteromyceter, fig. 29 (spores).
Lloyd. The Geastrae, figs. 44 and 45.

New York. Syracuse. Oct. 1888. Underwood, coll. (N. Y. Bot. Gard. Herb; also a small form as

C. Schae fieri).
Massachusetts. South Hadley. 1888. Hooker, coll. (N. Y. Bot. Gard. Herb.).
Kansas. Bartholomew, coll. (N. Y. Bot. Gard. Herb., Xo. 2215; U. N. C. Herb., No. 8252). Spores

of Xo. 8252 are 3.4-4M thick.
Wisconsin. Burlington. (Univ. Wis. Herb, and U. N. C. Herb.)
Minnesota. Miss Hone, coll. (N. Y. Bot. Gard. Herb, and Lloyd Herb.). Spores dark, distinctly

warted, 3.5— 1.2m, most about 4ju.
Colorado. Brandegee, coll. (N. Y. Bot. Gard. Herb.).
Canada. Macoun, coll. (N. Y. Bot. Gard. Herb.).

London. Dearness, coll. (U. N. C. Herb.). Spores 3.6-4.2m-

Geaster Morganii Lloyd

Plate 115

Button broadly bulb-shaped with a short to rather long point (up to 6 mm.),
yellowish, the base flat; mycelial attachment from a central basal point, the surface
nearly or quite clean, buffy alutaceous, dull, felted to minutely spongy; rays about
7-10, taper-pointed, pliable, the outer layer tending to crack and peel off in irregular
flakes (more separable than in typical triplex or lageniformis) ; fleshy layer dark brown
to blackish, cracking and at times forming a collar as in G. triplex; spore sac sessile, sub-
spherical, usually compressed when dry, minutely granular-felted, wearing glabrous,
brown ; peristome rather vaguely outlined, forming a narrowly conical papilla (typically)
with the sides crumpled toward the top or almost all over, forming a pseudo-sulcate
peristome with lacerated tip.

Spores (of a plant from Ohio; Underwood, coll.) spherical, distinctly warted, 3.4—i.ln
thick. Capillitium threads wavy, up to ly. thick.

The plant seems common in the middle west, less so elsewhere. We know of no
record for the southern states, but there is in the Curtis Herbarium a plant from Ala-
bama (Peters, No. 198) as G. saccatus that we believe to be this species. The mouth
is very long and narrow and not at all like triplex, but is not crumpled. Lloyd mentions
such a mouth as occurring. The nearest relative seems G. triplex, which differs in even

110 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

mouth and smooth outer surface. Lloyd now considers his G. Morganii a synonym of
G. Archeri Berk. (Hooker's Fl. Tasm. 2: 264, pi. 183, fig. 9 bis. 1860) but compared
with American G. Morganii, the lobes of G. Archeri are more slender and tapering, and
the outer surface differs in not being cracked. The fleshy layer looks quite different
and has a hard, cartilaginous appearance with few cracks, instead of the softer, more
leathery appearance of the rimose fleshy layer in G. Morganii. The spores are the same.
The type material of G. Archeri at Kew consists of two separate mounts of one open
plant each, both from Tasmania (Archer, coll.). The plants have long, tapering and
twisted lobes, rather pliable ; fleshy layer thin, continuous ; outer layer with texture of
leather, not cracked in radial lines; spore sac deep brown, glabrous, mouth apparently
as in G. Morganii, but somewhat obscured by pressure. Spores (of No. 28) spherical,
regularly warted, 3.4-4yu thick (pi. 116, fig. 14).

Illustrations: Lloyd. The Geastrae, figs. 31-36.

Morgan. North Amer. Geasters, fig. 3 (as G. striatus).

Delaware. Commons, coll. (N. Y. B. G. Herb.).

New York. Chippewa Bay. Phillips, coll. (Path and Myc. Herb.). Spores minutely warted, 3. i-Aii

thick.
Massachusetts. Peabody. Mackintosh, coll. (Lloyd Herb.). Fine lot with buttons.
Ohio. Underwood, Morgan, Lloyd and James, colls. (All in N. Y. B. G. Herb.).

Columbus. Brewer, coll. (Lloyd Herb.) .

Cincinnati. Lloyd, coll. (Lloyd Herb.).

Dinwood. Lloyd, coll. (Lloyd Herb.).
Indiana. Underwood, coll. (N. Y. B. G. Herb.).
Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).

Geaster saccatus Fr.

Plates 65, 115 and 117

Buttons gregarious, not caespitose or crowded, 7-12 mm. thick, 7-14 mm. long,
including point, when fresh, entirely subterranean or half exposed, abruptly pointed by a
short small mucro about 1.5 mm. long; surface nearly clean, tomentose to spongy-felted,
ochraceous to buff. Rays about 5-7, upright or revolute, the base strongly saccate.
Fleshy layer when fresh pallid, faintly or distinctly pinkish, about 0.8 mm. thick.
Spore sac light or dark drab, glabrous, dull; peristome, broadly conical, definite, sur-
rounded by a paler depression or much paler all over (at times nearly pure white) ;
mycelium attached only as a basal strand, the submerged part of the plant, which is
more often all in the button, not binding trash to it, but coming up nearly clean, the
mycelium forming a powdery-looking, mouldy layer about 1 cm. from the surface.

Spores (of No. 7350) spherical, minutely asperulate, 3. 5-4. 3m-

In Chapel Hill we find this only under cedars, but the Florida collection was under
oaks, though otherwise exactly like the others, the whiter peristomes being matched in
some of our collections. Our plants are very small, only 6-15 mm. wide when open in
the dry state, and they look most like G. mirabilis. That is separated by the strictly
epigeal and crowded habit, absence of point, strigose surface and entirely different
mycelium. The tropical form of saccatus may run much larger than ours, but there are
intermediate sizes. What we are calling the northern form of saccatus runs somewhat
larger than ours and has much smaller spores.

There is still some doubt as to what the true G. saccatus is. As Fries described it,

PI.ATK (o

-.

Geaster mirabilis. No. 7331 [above]
Geaster saccatus. No. 73S0 [below]

l.WOPERDACEAE 111

it is obviously in the G. triplex rather than in the G. fimbriates series, as indicated by
the radicating, basal mycelium and the clean or trashy, adnate, soft, leather colored
outer surface, and long, narrow points of the rays. Authors and collectors have refer-
red to G. saccatus both G. triplex (including the lageniformis form) and G. jimbriatus,
as well as the plant that we are calling G. saccatus. Lloyd confuses the latter two in
America and has also referred the lageniformis form of G. triplex to G. saccatus (see plants
at the New York Botanical Garden, sample box). Morgan distinguishes them properly
by the difference in the mouth, but fails to call attention to the difference in the outer
surface which is at least as important. Rick has distributed from Brazil as G. saccatus
two quite different plants; one (Fungi Aus.-Amer., No. 330) which looks just like G.
fimbriatus, with rlocculent trashy outer surface and indefinite mouth. The spores,
however, are distinctly larger than in G. fimbriatus, faintly warted, 3.8-4.6ju thick.
Another (Fungi Aus.-Amer., No. 261) is just like our G. triplex, except that the outer
surface is cracked in a mottled way rather than in radial lines (spores 3.5-4. \/x). This
mottled cracking may also be noted in U. S. American G. triplex (as plant from Love-
land, Ohio, noted under that species). Rick's figure of G. saccatus (Broteria 5: pi. 2,
figs. 6 and 8) shows a little plant with distinct mouth and clean surface that looks just
like the plants we are calling by that name. What Spegazzini distributed from Argen-
tina as G. saccatus (Dec. Myc. Arg., No. 44) is a hygroscopic plant with an indefinite
mouth and about the size of G. mammosus. It is evidently not the Friesian species.
(This is noted also by Lloyd.)

7173. Under cedars, buttons covered up, The Rocks, Sept. 26, 1923. Peristome very pale. Spores

3.2-i.ln.
7441. Under cedars in Mrs. Kluttz's yard, July 20, 1924. Spores 3.5-4.5/j.
7475. In humus under cedars, Aug. 5, 1924. Spores 3.5-4.2/*.
Also Xos. 7350, 7415, 7479, 7506.

Florida. Columbia Co. On lake bank under oaks, June 28, 1924. Mouth (of all four open plants)

nearly pure white, contrasting strongly with the dark drab sac. Spores minutely spiny,

3.4-l'„.
Alachua Co. Walker, coll. (U. N. C. Herb.). Spores very dark, subspherical, distinctly warted,

3.8-4.8/j thick.
Xew Smyrna. Sams, coll. (Lloyd Herb.).
Gainesville. Among leaves in moist woods, Aug. 16, 1926. Erdman West, coll. (U. N. C.

Herb.). Spores 3.5-4.2/i.
Mexico. Botteri, coll. (Curtis Herb., as saccatus).

Porto Rico. Stevenson, coll. (N. Y. B. G. Herb.). Spores minutely warted, 3.4-3.7//.
Bahamas. Xew Providence, Sept. 1904. Mrs. Britton, coll. (N. Y. B. G. Herb.).
Jamaica. Xew Haven Gap, 5600 ft. elevation. Murrill, coll. Jan. 1909. (N. Y. B. G. Herb.)

Spores 3.7—1.8/1.
Cuba. Wright. ("Curtis Herb., as G. saccatus, Xo. 668 and 870, and as C. fimbriatus, the latter with a

clean, pointed button.)
Bolivia. January 1902. R. S. Williams, coll. (X. Y. B. G. Herb.). Spores spherical, minutely but

distinctly warted, 3.4-4/1 thick.

Geaster saccatus Fr., Northern Form

Plate 115

The following is drawn up entirely from dried plants:

Plants saccate, of medium size or rather small, the expanded rays with a spread of
2.5-4 cm. (see note on a larger form below); buttons subglobose with a point up to 4

112 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

mm. long, or rarely with only a rounded umbo. Mycelium strictly basal and radicating
by flocculent strands from a central basal point ; outer surface felted or nearly smooth,
almost clean or at times with a few bits of rotten wood or leaves, not cracking, with some
tendency for outer layer to separate on the rays which are 6-9, pliable, thin, expanded,
with the tapering rips revolute; fleshy layer reddish brown, thin, adnate, somewhat
cracked. Spore sac 10-13 mm. broad, sessile, surrounded below by the saccate base,
smooth, brownish drab to gray-drab; peristome silky, paler or nearly concolorous,
distinctly or rather vaguely limited. Columella not obvious.

Spores (of plant from Asheville) spherical, dark, minutely but clearly warted,
2.2-3.3/*, distinctly smaller than in the southern form. Capillitium threads about the
color of the spores or paler, 5-7.5/* thick.

Gregarious on leaves of deciduous trees with a tendency to crowding, which is more
or less obvious in most collections. Apparently rare. We have not found it at Chapel
Hill.

Neither this nor the southern form can be distinguished from G. triplex by the
absence of point on the buttons, as there is a point or umbo present both when fresh
and dry. Buttons in the same collection may vary from slightly umbonate to pointed
like our photograph of the Asheville collection. Hollos, Morgan, Trelease, and
Lloyd include our U. S. American plant in G. saccatus. Hollos separates it correctly
from G. lageniformis by its soft outer peridium which is usually free from trash. It
is further separated from that species which has a firm, hard outer surface by the absence
of longitudinal surface cracks that are usually present in the latter, and, in the northern
form, by the smaller and less rough spores. Hollos is wrong in considering G. velutinus
the same as G. saccatus. The former is very easily distinguished from G. saccatus by
its larger size, firmer, harsher, paler, more spongy-looking and more separable surface
layer.

At the New York Botanical Garden is a collection that we take to be this (Gentry,
coll. Two buttons and an open plant; no data). The buttons are larger than in the
others, 2.2 cm. thick and up to 2.6 cm. tall including the point, which in one is about
1 cm. long, in the other about 4 mm. Surface layer soft and felted, very clean except
on the base, separating in places from the middle layer ; mycelium attached to a basal
point. Spore sac sessile, mouth very distinct. Spores spherical, minutely warted,
3-3.6n thick.

In the Lloyd Herbarium there are two collections of G. saccatus from Berlin
(Magnus, coll.). They are like the American plants, with surface nearly or quite clean,
peeling off to some extent from the tips of the rays.

Illustrations: Hollos. 1. c, pi. 10, fig. 19.
Lloyd. The Geastrae, fig. 75b.
Rick. Broteria 5: pi. 2, figs. 6 and 8.

North Carolina. Asheville. Beardslee, coll. 1917. (U. N. C. Herb.)

Maryland. Plummer's Island. P. L. Rickef, coll. 1922. (Path, and Myc. Herb., Washington.)

Ohio. Lloyd. (N. Y. B. G. Herb., 2 colls.) Spores of one minutely rough under high power, 2.3-3.1/*

thick; in the other, 3-3. 5m thick. Also E. & E., North Amer. Fungi, No. 3417. (Path, and

Myc. Herb.)
Cincinnati. Aiken, coll. (Lloyd Herb.).
Kansas. Fort Scott. On leaf mold, July 1902. Garrett, coll. (N. Y. B. G. Herb., no name).

Cespitose, small, soft, surface felted. Exactly like Lloyd's plant from Ohio.
Missouri. St. Louis. Glatfelter, coll. (Lloyd Herb.).
Canada. London. Dearness, coll. (U. N. C. Herb.).

LYCOPERDACEAE 113

Geaster saccatus var. Walkeri

Buttons subspherical with or without B small blunt point, gregarious to subcespi-
tose, 1-1.5 cm. thick, epigean and basally attached to an obvious floccose mycelium,
surface felt-like, not strigose. very dark brown (verona brown of Ridgway), splitting
into 6 or 7 bluntish rays which extend only about one-third to one-half the distance and
confine the spore sac to a deep cup, outer layer not at all separable. Fleshy layer when
drv varying from dark brown to much paler. Spore sac grayish brown, the mouth
without a peristome, fimbriated about as in G.fimbriatus.

Spores minute, faintly rough, 2-3. 2/u.

The plant differs from G. fimbriaius in entirely different habit (basal attachment
of exposed buttons and inseparable spongy surface) and from G. saccatus and its rela-
tives in absence of peristome and very dark color. The plants are larger than any
species of the saccalus group except saccatus itself. The spores are smaller than in
any of the saccatus group except what we have called the small-spored northern form.
Miss Walker writes that "All of the colors had a beautiful rosy tint to them in the fresh
condition."

Nebraska. Lincoln. Fall of 1926. Miss Leva B. Walker, coll. (U. N. C. Herb. No. 4).

Geaster velutinus Morg.

G. Readcri Cooke and Massee
Cydoderma ohiensis Cke.

Plates 31, 66,67 and 115

Unopened plants seated on the substratum (not embedded in it) and therefore
attached to the mycelium only at the base, ovate and bluntly pointed at the top, up to
2.7 cm. broad and' long. Surface dull and finely felted-tomentose, very much like the
surface layer of Lycoperdon leprosum. It is flesh color, becoming creamy yellow or
buff color on drying. Outer peridium splitting into about 7 reflexed or expanded seg-
ments which are a pretty flesh color on the inner (upper) surface when fresh. As these
dry they split more or less into two thin, fibrous, persistent layers, a characteristic habit,
and usually curl backward under the basal part which remains convex or flat below.
Inner peridium smooth, appearing minutely felty under a lens, dark brown or sometimes
light gray (when collected perfectly fresh), subglobose, attached by a broad or rather
narrow base, almost or quite sessile; mouth small, fibrillose and later lacerated, sur-
rounded by a distinct, radially fibrous, broadly conical, silvery gray or light to dark
brown area which is about 5 to 8 mm. in diameter. Columella obvious and clavate in
the button; slender and more obscure in the mature plant.

Spores (of No. 4095) spherical, 2.5-3.3// thick, most about 3yu thick, distinctly as-
perulate and with a halo, deep smoky purplish brown. Capillitium threads rather large,
up to 11m thick, often branched near the tapering tips.

In an unopened button of average size, a microscopic section of the peridia showed
the following: outer peridium composed of four distinct regions, (1) a surface layer
about 35(MO0m thick in which the threads are large, irregular, entangled and ap-
proximately perpendicular to the surface of the plant; (2) a very thin layer 50-65m
thick in which the threads are small, regular and extend approximately parallel to the
surface of the plant; (3) a slightly thicker layer 130-150p thick in which the threads are
small, regular and much entangled; and (4) a very thick (1.5 mm.), fleshy region com-
posed of large parenchymatous cells. Inner peridium composed of a single very thin

114 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

layer made up of delicate threads that run approximately parallel to the surface. The
threads of the four layers of the outer peridium pass into each other, but at maturity
the fleshy layer and the two outer layers (which are an inseparable unit) are easily
removed from the third layer. The columella, inner peridium, and all the layers of the
outer peridium are continuous with each other at the base.

In this species Cunningham finds (1. c.) that the basidia are uninucleate and the
plant appears to be haploid throughout its development. The nucleus divides to form
nuclei for each spore, usually four. Clamp connections were not found in any part of
the plant. All attempts to germinate the spores were unsuccessful.

A rather common, large plant that prefers pine woods. It is easily distinguished
from its relatives of similar habit by the larger size and minutely felted-tomentose,
not hairy, outer peridium which splits into two persistent layers. In many individuals
there is plainly to be seen on the under side a circular scar marking the area of attach-
ment. From a part of the type collection of G. Readeri from Australia (Grevillea 16:
73. 1888), at the New York Botanical Garden, it is shown that it is the same as G.
velutinus, the plants not being distinguishable from the American ones in any way,
the surface pale, velutinate, peeling off in part; peristome distinct, size and shape of
plant the same; spores spherical, very minutely warted, 2.8-3.3^ thick. There are two
other plants mounted in the same herbarium that are also marked "type specimens."
These have some sand on the outer surface, but the spores are the same, 3-3.5/j, thick.
The plant was first named G. australis by Reader, a name previously used by Berkeley.
A plant from New Zealand determined as G. Readeri by Massee (N. Y. Bot. Gard.
Herb.) is not the same. It has the surface more strigose-tomentose and not peeling
off, and the spores are distinctly different, warted, 3.8-5/*. The plant distributed by
Rick in Fungi Austro-Amer., No. 13, as G. triplex is G. velutinus. We have a plant from
Brazil (Murrill, coll.) which seems intermediate between velutinus and subiculosus.
The silky peristome is very distinct, and the button has almost the same surface as G.
velutinus; the mature plant is nearly smooth. The spores are minutely warted, 2.5-3.6/t
thick.

Illustrations: Cunningham. Trans. N. Z. Inst. 57: pi. 10, fig. 33; pi. 11, fig. 34. 1926.
Hard. Mushrooms, fig. 489.
Lloyd. The Geastrae, figs. 62-71. Also pi. 101, figs. 1-2.

846. In pine woods, October 13, 1913.
1260. Under pines in woods, September 24, 1914. Spores 2.7-3m thick, minutely asperulate. Capil-

litium threads 3-5.5^ thick, minutely roughened.
7133. In rotting leaves under shrubs in Arboretum, September 22, 1923. Buttons exposed, subspheri-

cal, not umbonate when fresh.
Also Nos. 999, 1000, 1381, 4095, 7114, 7164, 7219, 7442, 7503.

Statesville. Coker, coll. In pine needles, Aug. 1922. (U. N. C. Herb., No. 5499.)
Asheville. Beardslee, coll.

Florida. Gainesville. Walker, coll. (U.N. C. Herb.).
Alabama. Auburn. Baker, coll. (N.Y. B. G. Herb.).

Cuba. Wright, No. 254. (Curtis Herb., no name.) Button and open plant.

Porto Rico. J. R. Johnson, coll. (N. Y. B. G. Herb.). The plant, though saccate, has slipped entirely
out of the sheath.

•LATE 67

\?

%

^r

L .f$#

Geaster mirabilis. No. 7208 [upper left]

(.1 ISTEB CAESFTTOSUS. Ohio [upper right]

Geaster trichifkk. Morce's Gap, Jamaica [middle left]

Geaster srjBictnosus. No. 184 [middle right]

(,i \>ter velutims (Buttons). No. 7219 [below]

LYCOPERDACEAE 115

Brazil. Rick. (Path, and Myc. Herb., Washington.) Also reported by Sydow (Ann. Myc. 5: 353.
1907).
Mt. Corcovado. Murrill. coll. (X. Y. B. G. Herb, and U. N. C. Herb.).
Africa. Uganda. 4000 ft. elevation. Diimmcr, coll. (Lloyd Herb.). These are buttons which look

exactly like the American ones.
New Zealand. Cunningham, coll. (U. N. C. Herb.). Spores minutely rough, 2.5-3.6^ thick.

Geaster radicans B. & C.

Plates 115 and 116

The co-types of G. radicans (Grevillea 2: 49. 1873) in the Curtis Herbarium
(as G. Curlisii) show it to be closely related to G. vclulinus. In so far as the appearance
goes, it is only a fornicate form of that species, as Lloyd says (Myc. Notes, p. 155;
The Geastrae, p. 31). There are two mounted plants with basal cups gone and one
perfect plant in a box, all Ravenel, No. 953. Ravenel's ms. notes accompany the last
specimen. The velutinate outer surface and the felted spore sac are exactly as in G.
vclulinus. A microscopic examination, however, reveals a difference in the spores,
which are larger than in G. vclulinus, 3.5-4m thick, and more distinctly rough. We have
examined the plants from Florida (Sarasota, on a cedar stump) in Washington, one of
which was photographed by Lloyd (The Geastrae, fig. 57), and find them exactly like
the types, the spores 3.4-4/* thick. The basal cup is a strong, pliable membrane, free
from earth, and in no w^ay like the delicate, submerged cup of G. jornicatus and G.
coronaius. We find at the New York Botanical Garden a specimen of this plant from
Bermuda, which extends its known range. The spores of this plant are spherical,
distinctly warted, 3.8-4.8/u, most about 4.1m thick. The species is reported as new to
Europe by G. Lind in "Danish Fungi as Represented in the Herbarium of E. Rostrup,"
p. 401, 1913 (no description). Rick figures a plant as G. radicans from Brazil (Broteria
5: pi. 5, fig. 9) that is not fornicate and looks exactly like G. triplex with a large collar.
He had earlier reported this as G. triplex, and says that Bresadola referred his plant to
G. radicans. It may be G. vclulinus, but cannot be accepted as G. radicans. Lloyd
thinks that G. Wdvntsckii (from Spain) is the same as the American plant (Myc. Notes,
p. 315, pi. 101, fig. 3).

The Cuban plants reported in the original description as G. radicans are represented
in the Curtis Herbarium as Wright, No. 873. These three plants are all buttons and
are not G. radicans. They were entirely submerged and are covered with humus held
on by a flocculence as in G. rnjescens; a single stout basal strand is attached to buttons
in two cases; not pointed.

The types were sent to Berkeley by Curtis, who received them from Ravenel, who
also sent notes and added that if a new species then "it is G. Curlisii" wishing to honor
Curtis. Berkeley, however, ignored this wish of Ravenel's and named it G. radicans
B. & C. (See Lloyd, Myc. Notes, p. 171.) Ravenel in his notes says "with a sulcate-
plicate mouth," but the mouth is not truly so as we are defining that character. The
best specimen has the mouth pinched into a few crumples, but the others do not show
this.

As the usual form and spores distinguish G. radicans from G. vehttini(s, we think
it well to retain the name, though it would be better to reduce it to a variety.

116 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustration: Lloyd. As cited above.

Patterson and Charles. U. S. Dept. Agric. Bull. 175: pl.36, fig. 1. 1915.

South Carolina. Ravenel, No. 953; Curtis, No. 3041. (Curtis Herb., as G. Curtisii Rav., co-types;

also in Farlow Herb, from Ravenel.)
Florida. Sarasota. On a cedar stump. (Path, and Myc. Herb.)
Bermuda. Millspaugh. (N. Y. Bot. Gard. Herb.)

Geaster mirabilis Mont.

G. caespitosus Lloyd
G. lignicola Berk.

Plates 65, 67, 68, 115 and 116

Densely cespitose to single in considerable numbers, and arising from an obvious
white mycelium which binds together the leaves and twigs. Buttons spherical, not
pointed, but a few with a small umbo when dry, 0.8-1.2 cm. thick, strigose-tomentose,
reddish brown to ochraceous buff, a few blackish; mycelium basal, the buttons quite
exposed or partly covered with loose trash (completely covered in No. 7173). Ex-
panded plants 1.4-2.2 cm. wide; lobes usually 6 or 7, expanded (tending to become
elevated when dry), the basal half bowl-shaped and holding the completely sessile inner
peridium which is 6-9 mm. thick, subspherical, gray, with a delicately felted surface;
peristome distinct, silky, more or less elevated, conical, immediately after exposure pale
from a granular-looking, minute pubescence, with a darker line around the base, soon
darker than the peridium; mouth fimbriated, not sulcate. Fleshy layer when first
exposed about 0.5 mm. thick, smooth, light flesh color, soon darker and drying to a
thin, adnate, continuous, gray-brown membrane.

Spores (of No. 7077) very dark, minutely warted, 3.2-3.8/*, most about 3.6/u.
Capillitium threads sinuous, thick- walled (practically no lumen), much paler than the
spores, about 3.4/* thick, the tapering ends smaller.

From G. saccatus the present species is distinguished by cespitose habit, much
smaller size, hairy surface, less pointed buttons when dry and larger spores. Both have
the habit of growing exposed (epigeal).

The species was originally described from French Guiana and is known also from
Brazil, Africa, and the Orient.

Our plants are the same as plants received through the kindness of M. Patouillard
and collected in Brazil by Rick. The young buttons have the same strigose-tomentose
surface ; the older ones more felted-tomentose ; spores exactly as in our plants, faintly
warted, 3.4-4/i thick, not like the smooth and smaller spores of what we are calling
G. subiculosus. Rick considers G. trichifer the same (see Myc. Notes, p. 804), but
Lloyd thinks it different (see that species). In our No. 7077, some of the plants are
apparently as strigose as G. trichifer (which see), while others show only a matted tomen-
tum, exposure to weather having a considerable effect.

The species is represented in the Kew Herbarium by five cards of material: three
from Cuba (Wright), two from Ceylon (G. H. K. T.), and one from Uganda, Africa,
(Diimmer, No. 1462). All these have the spores of G. mirabilis, and in appearance are
the same as the Chapel Hill and Brazilian specimens except that none shows quite so
strigose a surface. The spores of the plant from Ceylon, No. 184, are 3.2-3.8/* thick,
minutely rough. The African plants connect up directly with G. papyraceus, with
outer peridium more papery-looking than in the others. The spores of this collection
are 3.5-4.2/* thick, with surface as in the others.

ri.\i i 68

[Top row] GEASTER Harioiii. Brazil (left) ; Porto Rico (middle); British Honduras (right)
[Second row] Geaster caespitosus. Type (left)

Geaszes arenarius. Type (light)
[Third row] Geaster florieormis. Grasmcrc, Florida (three tiny plants) ; Denton, Texas

(2 larger plants in middle); North Dakota (one plant on right)
[Bottom row] Geaster asper. Long Pine, Nebraska

1.W0PERDACEAE 117

Caster papyraems B. & C. (Proc. Amer. Acad. Arts and Sciences 4: 124) in the
Curtis Herbarium from Japan and Benin Islands (authentic, but not type) is closely-
related if not the same. The Japanese plants are all open ; peridium extremely thin and
paperv, pliable, pale creamy straw color, outer surface with a concolorous coat of very
thin soft weft in places. Mycelium membranous; fleshy layer almost all gone; inner
peridium broken or worm eaten, peristome not shown. The Bonin Island plants are
strigose-hairy outside; two cespitose buttons, one grown button apparently torn open
at the top, and one open plant; even the last is strigose. These look like G. mirabilis.
Fleshy layer intact and rays much stiffer than in the Japanese plants. Peristome
distinctly silky. A microscopic examination, however, shows the spores to be identical
in the two collections. In the Japanese plants they are spherical, finely but distinctly
warted, 3.5-3.8/i thick, with a very distinct and shiny oil drop. In the Bonin Island
plants they are 3.6-4^ thick, surface and oil drop as in the others. We can confirm
Lloyd's opinion (Myc. Notes, p. 181) that Coilomyces Schweinitzii B. & C. is an un-
opened Geaster related to, if not the same as, G. mirabilis. The plants in the Curtis
Herbarium have spores exactly like our Chapel Hill G. mirabilis, spherical, minutely
warted, 3-An thick (pi. 116, fig. 15).

Geaster caespitosus is apparently the same, with the usual slight variations which
are to be expected. The ample type collection in the Lloyd Herbarium has the exact
appearance of our plants except that the strigose character is not so obvious. The
surface is softly felted, becoming smoother; peristome silky, sharply defined by a ridge;
the plants attached by a pointed base to the abundant mycelium. Spores slightly
smaller than in our collections, very minutely warted, 2.3-3.4^. We find at the New
York Botanical Garden a box of miscellaneous material without names but labelled
Ohio, Lloyd, No. 255, in which, mixed with three other species, is a dense clump of small
plants that are G. caespitosus and just like the types. The type of G. lignicola Berk.
(Journ. Linn. Soc. Bot. 18: 386. 1881) at Kew from Rockingham, Australia (Thozet,
No. 908), is apparently not different from G. mirabilis. There are two crushed buttons
and one open plant on fragments of plant material. Mycelium not very obvious;
surface of both buttons and mature plants nearly smooth. Spore sac dark brown with
fimbriate mouth. Peristome characters not obvious on account of the way the plant is
pressed and glued down. Spores spherical, minutely warted, 3.2-4m thick.

Illustrations: Lloyd. Myc. Works, pi. 100, figs. 2 and 3; also figs. 6-9 (as G. caespitosus).
Rick. Broteria 5: pi. 4, fig. 3. 1906. Freshly opened plants.

7077. On trash and leaves of deciduous trees and a few pine needles, wooded hillside, August 10, 1923.
7208. In rich, rocky soil near cedars, November 11, 1923.
7502. In humus in mixed woods (mostly deciduous) , Aug. 14, 1924.
AlsoXos. 7331,7413, 7414.

Virginia. Warrenton. Upland frondose woods, Sept. 6, 1926. Coker, coll. (U. N. C. Herb.)

Ohio. Lloyd. (X. Y. Bot. Gard. Herb., unnamed.)

Kansas. Cragin, coll. (X. Y. Bot. Gard. Herb., as G. saccatus). Outer surface clean, tomentose;

spread when expanded 1.8 cm. Spores minutely warted, 3.5— 4.2^.
Cuba. Wright, Xo. 696. (Curtis Herb.) Button very small, strigose to tomentose.
Jamaica. Bancroft. Ellis, coll. (Farlow Herb.).
Brazil. Rick, coll. (U. X. C. Herb.).

118 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Geaster trichifer Rick

Plates 67 and 115

Plants gregarious to cespitose; buttons exposed, subspherical to oblong, pinched
below to a point and attached to ropy mycelium, slightly umbonate above, bufly straw
color, covered with more or less fascicled hairs about 1 mm. long; rays about six, splitting
to or below the middle and tending to remain upright or with tips spreading; outer
layer showing a distinct tendency to peel off the rays as in velutinus, but less so; fleshy
layer leather color, thin, adnate. Spore sac subglobose, sessile in the cup-shaped base,
grayish brown, glabrous; peristome very distinct, flat with elevated center, mouth
becoming fimbriate.

Spores very minutely rough, 3-3.6/j. thick.

Murrill's notes on the fresh condition say, "Outside with long, stiff hairs, ferruginous-
fulvous, shaggy-setose . . . fleshy layer dirty white; spore sac flattish acorn-
shaped, avellaneous. . . . Spread 5 cm., center 2 cm." One of the largest plants is
now broadly spread and reaches 4 cm., the smaller ones if spread would be about 3 cm.
The plants are growing on leaves and twigs and have the habit and shape of G.
mirabilis, which differs from the Jamaica collection in being much smaller, less strigose,
with outer layer not peeling, and in the smaller spores.

The original plants were collected by Rick in Brazil. He now considers it a form
of G. mirabilis, while Lloyd thinks it different. The plants shown in Lloyd's photo-
graphs are not larger than G. mirabilis, and in fig. 1251 the surface does not appear more
shaggy than in G. mirabilis.

Illustrations: Lloyd. Myc. Notes, p. 314, figs. 147, 148, and p. 804, fig. 1251.

Jamaica. Morce's Gap. Murrill, coll. No. 739. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.)

Geaster subiculosus Cooke and Massee (sense of Lloyd)

Plates 67, 115 and 116

Plants with the habit of G. mirabilis, crowded and springing from copious, white,
felted mycelium which binds together the leaves, twigs, and sticks on which they grow.
Buttons 9-11 mm. broad, completely exposed, obovoid, pinched below to a point
connecting with the mycelium; apex concave by collapse (when dry), not pointed.
Surface pale buff, wrinkled, glabrous, but with the soft appearance of leather; tardily
opening by 6 or 7 rays, the sinuses extending into about a third of the peridium, leaving
the basal part unchanged as a deep cup enclosing the sessile spore sac. Fleshy layer
drab brown, thin, adherent, not cracked except across the bottom of the rays. Spore
sac 1 cm. broad, subspherical, glabrous, brown. Peristome not distinctly outlined or
distinctly silky, fading into the sac surface about as in G. fimbriatus, sometimes sunken
and then more distinctly outlined; mouth scarcely elevated, the opening fimbriate.

Spores (of Jamaica plant) spherical, smooth, 2-3// thick.

This is clearly distinguished from G. mirabilis Mont, by larger size, absence of
tomentum, obscure peristome, and smooth and much smaller spores. Even very young
buttons are practically glabrous and nearly white. The plant described above is
certainly the same as that illustrated by Lloyd in Lycoperd. Australia, etc., fig. 19, and
Myc. Works, pi. 100, figs. 4 and 5. He refers to it as G. subiculosus, which he considers a
form of G. mirabilis. From the description of G. subiculosus (Grevillea 16: 97. 1887)
one would hardly refer the present plant there, as it is described as furfuraceous and

LYCOPERDACEAE 119

wood colored on the outside, with spores 4;u thick, but the types at Kew from Trinity
Bay, Australia, are essentially like our plants from Jamaica. There are two open
plants and several more or less crushed buttons. The best specimen shows a spore sac
about 7 mm. thick, dark brown; peristome not outlined by a groove or ridge but paler
and fibrous. Surface of buttons minutely spongy, of older plants nearly glabrous.
Largest button 1.1 cm. thick. Subiculum white, membranous. Spores spherical,
practically smooth, some finely dotted, 3-3. S/u.

Lycopcrdon f-usio B. & C. is apparently a Geastcr. The co-types in the Curtis
Herbarium, No. 253 (Cuba), are all buttons, but they look just like another collection
so labelled, also from Cuba (Wright) , in which the plants are open. These last are small
Geasters looking very much like G. subhiilosiis, with pale, smooth surface, and con-
spicuous mycelium. The spores, however, are not those of any member of this group,
but are 3.8-4.Sm thick, with distinct, blunt warts.

Illustrations: Lloyd. As cited above.

Jamaica. Hope. Earle, coll. (N. Y. Eot. Gard. Herb., No. 184).

Geaster mammosus Fr.

G. cor all inns (Batsch) Hollos
G. lugubris Kalch.
G. argentcus Cooke

Plates 115 and 116

Button bulb-shaped with a distinct tapering point about 3^1 mm. long when dry,
covered all over with a finely felted, thin, soft, buffy yellow coat which is usually quite
clean of dirt except at the base and apparently exposed. Mycelium basal; outer scurf
wearing off gradually and exposing the glabrous shining copper brown or darker, fibrous
layer of the rays, which are about 8-12 in number, of unequal width and strongly hy-
groscopic; fleshy layer thick, adnate, smooth, at first gray-brown then darker brown to
blackish. Inner peridium 5-15 mm. thick, subglobose to compressed, very finely
puberulent, sessile, grayish to brown then darker; peristome distinctly outlined by a
depressed border, plane, silky-strigose, the mouth elevated and papillose-fimbriated.
Expanded plants up to 5.5 cm. broad.

Spores (of a Canada plant) spherical, minutely warted, 3-3.7 n-

Easily denned in the hygroscopic section by the clearly outlined silky-fibrous,
plane and non-sulcate peristome. The above description was mostly drawn from a
fine collection from Canada, containing one button and plants of various ages and sizes
(N. Y. Bot. Gard. Herb., no data except Canada).

Plants in the Lloyd Herbarium from Paris (L. Rolland) are like American speci-
mens, with a spongy, yellow, nearly clean superficial layer peeling off in flakes; the mouth
distinct. Plants in the same herbarium from Tucuman, Argentina (Leon Castillon,
coll.), are like the United States and European plants except that the outer layer when
present is covered with sandy earth.

Geaster argentens Cooke (Grevillea 17: 75. 1889) is represented at Kew by three
sets on the same sheet, all from Fort Carlton, Saskatchewan (one lot not named). The
plants of the upper lot with pasted label giving fullest data, we take as the types. The
plants have the appearance of G. mammosus w-ith outer superficial layer weathered off,

120 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

showing the whitish, fibrous layer and blackish fleshy layer; spore sac sessile and largely
enclosed by the strongly hygroscopic lobes. The mouth area is obvious as a silky
zone, the center more or less elevated and lacerate. The spores, which are spherical,
distinctly warted, 3.7-4.5/* (pi. 116, fig. 13) are distinctly those of mammosus and not of
floriformis, which are larger (5-7/z). Hollos considers G. argenteus the same as G.
floriformis, but he apparently had not seen authentic specimens.

Geaster argentatus Cooke and Massee from Brisbane, Australia, is closely related.
A part of the type collection is at the New York Botanical Garden. It has a definite
silky peristome; spread of rays 3 cm., the outer surface scurfy and ochraceous at first,
then smooth and gray to buffy gray; fleshy layer reddish brown; spore sac 12 mm.
thick, finely furfuraceous to sub-granular, gray-brown to reddish brown, sessile. Spores
dark, mostly spherical but some irregular, 4.5-6^ (pi. 116, fig. 17). Other plants from
Kew Herbarium labelled G. argentatus Cooke from Victoria, Australia (not types),
now pasted on the same sheet with G. argenteus, are quite different from the types, and
we take them to be G. Drummondii, which they resemble in all important respects:
mouth sulcate, spore sac granular to subasperate.

Illustrations: Destree. 1. c, pi. 9, fig. B.

De Toni. 1. c, pi. 2, fig. 0 (as G. lugubris).

Hollos. 1. c, pi. 10, figs. 1-3 (asG. corallinus).

Kalchbrenner. Gasteromycetes novi vel minus cogniti, pi. 5, fig. 3 (as G. lugubris). 1884.

Lloyd. The Geastrae, figs. 16, 17. Also pi. 98, figs. 7-11.

Micheli. Nov. Pi. Gen., pi. 100, fig. 3.

Smith. Grevillea2: pi. 19, fig. 1. (Reproduced from Gardener's Chronicle.)

Vittadini. Monog. Lycoperd., pi. 1, fig. 9.

North Carolina. Flat Rock. Memminger, coll. (Lloyd Herb., as G. floriformis).

New Jersey. Ellis, coll. (Lloyd Herb.).

Ohio. Cincinnati. Lloyd, coll. (Lloyd Herb.).

Wyoming. Cody. Davis, coll. (Lloyd Herb.).

Pitchfork. Davis, coll. (U. N. C. Herb.). Spores 3.8-5^.
Iowa. Iowa City. Teeters, coll. (Lloyd Herb.).

Colorado. Clements, coll. (Path and Myc. Herb.); also Demetrio, coll. (N. Y. Bot. Gard. Herb.).

North Dakota. Fargo. Brenckle, coll. (Path, and Myc. Herb., Lloyd Herb., and U. N. C. Herb.).

Note in Lloyd Herb, says "epigean." The plants show pointed buttons with yellow, spongy

surface and some adhering humus, mouth distinct. Spores (of No. 2S4) minutely warted,

3.6-4.4M.

Canada. (New York Bot. Gard., as G. umbilicatus.)

London. Dearness, coll. (Lloyd Herb, and U. N. C. Herb.).
Nova Scotia. Halifax. Crossland, coll. (Lloyd Herb.).

Geaster arenarius Lloyd

Plates 68, 115 and 116

Lloyd describes G. arenarius (The Geastrae, p. 28) as follows:

"Exoperidium subhygroscopic, cut to five to ten segments; drying usually with
segments incurved. Mycelial layer closely adnate with adhering sand. Fleshy layer
closely adnate, light color, not rimose. Inner peridium subglobose, with a very short
but distinct pedicel in some specimens, in others appearing sessile. Mouth even,
conical, acute, definite and usually darker colored than the remainder of inner peridium.
Columella indistinct. Spores globose, rough, 3-4yu."

PLATE 69

Geaster rufescens. Xo. 985

LYCOPERDAC] \r 121

The abundant type collection from Jupiter, Fla. (Culbcrtson, coll.), shows this to
be a remarkable plant intermediate between G. coronatus and G. Jloriformis. In some
the rays are distinctly revolute with the center elevated, in others the rays are closed
in over or entirely under the sac. Though some of the rays are pliable they are dis-
tinctly hygroscopic, expanding quickly in water. Spore sac very faintly felted to
glabrous, not at all granular-dotted as in G. coronatus, about 1 cm. thick, practically
sessile; peristome varying from sharply outlined by a groove (about as in G. coronatus)
to a less emphasized boundary as in G. mammosus; mouth apparently brown in the fresh
condition, washing out pale and becoming lacerate. The outer layer of sand is thin
and the flocculence holding it is very delicate and obscure. Spores minutely and ir-
regularly warted, 3.2-3.8/z thick. Mixed with the types is a good specimen of G.
coronatus, and this is almost certainly the plant referred to by Lloyd in his footnote
as being an exception.

From the appearance of the rays of G. arenarius, which are acute, but not acumi-
nate, the button is probably little if at all pointed. At the New York Botanical Garden
there are two plants from Florida labelled G. fimbriatus (Underwood, coll., No. 1801a)
that are almost certainly this species, though the mouth is only vaguely if at all deter-
minate. The plants are small, hygroscopic, covered with sand on the outside ; peristome
vague, spore sac nearly glabrous; spores minutely warted, 3.7-4.5yu thick, just like those
of Lloyd's plant and smaller than in G. ftoriformis.

The species is very near G. floriformis (see Lloyd, Myc. Notes, p. 143) and this
approach is further emphasized by the vague peristome of the Underwood plants, which
have the same spores as Lloyd's plants. The rays of G. arenarius are distinctly less
rigid than in G. Jloriformis, but the smaller spores of the former are now the principal
character separating the two species.

The extreme variation of Geasters is well shown by what appears to be a giant form
of G. arenarius that is represented in the New York Botanical Garden by two very large
plants from Greencastle, Indiana (as G. rufescens). They are 3.5-4 cm. thick with the
rays closed over the spore sac. The peristome is vague, but the spores are like those of
G. arenarius, minutely warted, 3.5-4^ thick, much smaller than in any other related
species. Another interesting case is that of plants from Lake Titicaca, noted under
G. jloriformis.

Illustrations: Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 5, figs. 24, 25.
Hollos. 1. c, pi. 29, figs. 19, 20.
Lloyd. The Geastrae, figs. 53 and 54.

Florida. Lloyd. (U. N. C. Herb.) Another collection by Underwood. (N. Y. Bot. Card. Herb.,
as C. fimbriatus.)
Lake Helen. Mrs. Xoble, coll. (Lloyd Herb.) Mouth distinct.
Grasmere. Baker, coll. (Lloyd Herb.).
Texas. Denton. Long, coll. (Lloyd Herb.). Mouth distinct. This is variable in its hygroscopic
character.

Geaster rufescens Pers.

Plates 69 and 115

Plants large, the button subspherical, not pointed, entirely buried until dehiscence,
when open 5 to 8 cm. wide; the thin outer mycelial membrane covered with earth or

122 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

trash and tending to split away in patches from the fibrous layer which is flesh colored
when fresh; rays splitting about half way to the base into about 7-9 lobes, or only
shallowly into many more, the lobes when fresh are expanded or recurved, when dry
recurved or the tips incurved, the unsplit basal portion convex above (vaulted beneath)
as in the fornicate group. Fleshy layer of outer peridium very thick when fresh, up to
5 mm.; when dry forming a thinner crust with a felted-looking, brown surface on the
face of the fibrous layer. Spore sac up to 4 cm. thick, depressed-globose, the base
usually constricted into a short thick stalk, not rarely with a low apophysis around the
lower side of the spore sac some distance from the stem; color when fresh varying from
pale flesh color to grayish flesh or deep brownish gray, when dry brown to gray brown,
the surface not glabrous but covered with very minute, inherent scurfy dots (granular-
velvety). Mouth slightly elevated, fibrous and becoming fimbriated, without a definite
peristome (indeterminate). Columella large and subglobose, up to 1.5 cm. thick, the
basal part with a peripheral extension as far as the apophysis.

Spores (of No. 985) globose, deep brown when mature, 3^1.4yu thick, minutely
warted or asperulate, with a halo. Capillitium of simple threads which are about
3.5-4.8n thick.

This fine species, which is not at all rare in America, grows usually around the bases
of oak stumps and occurs in groups. One of the most remarkable characters is the very
pleasant and decided fruity fragrance of the unopened and just opened plants. There
has been considerable confusion regarding this plant and G. limbatus. As understood
in Northern Europe (Fries, Berkeley, Massee), the latter differs in the expanded, not
reversed rays, the darker and smoother inner peridium, the thinner and less spongy
fleshy layer, and the more obvious peristome. For G. limbatus in the sense of Bresadola,
in which sense we are using the name, see under that species. European specimens of
G. rufescens are rare in American herbaria. One from Bresadola in the Lloyd Her-
barium is like our plants, with spores 3.6^1.2/* thick, minutely warted. For comparison
with G. triplex see that species.

Illustrations: Bulliard. Herb. Fr., pi. 471, fig. L.
De Toni. 1. c, pi. 62, fig. 1.
Hollos. 1. c, pi. 11, figs. 7, 8.

Lloyd. The Geastrae, figs. 38-41. Also pi. 97, figs. 6-8.
Morgan. North Amer. Geasters, fig. 11. 1884.
Petri. 1. c, figs. 45, 46.
Rea. 1. c, pi. 19.

985. At base of an old oak stump, November 17, 1913.
2984. On old oak stump, December 4, 1917. Spores 3-iAn thick, apparently pitted in water, but

upon application of KOH the surface appears spiny and with a halo.
7221. On damp clay soil, November 25, 1923. A small form; the spore sac only 1 cm. thick. Spores

3.3-4M.
Also Nos. 683, 1033, 6025, 6053, 6066.

Carolina. (Schw. Herb.; also in Michener Herb, from Schw. Herb.)

Alabama. Auburn. Under a rotting log, August 1897. Baker, coll. (N. Y. Bot. Gard. Herb., as
G. limbatus) . Two folders, each with three buttons, which are up to 3 cm. broad, subspherical
to compressed and completely covered with rotten wood held on by flocculence. Also a col-
lection from Alabama by Peters. (Curtis Herb., as G. limbatus.) This is typical G. rufescens
with a short stalk; outer surface covered with earth held on by flocculence.

New York. Botanical Garden. Williams, coll. (N. Y. Bot. Gard. Herb.).

LYCOPERDACEAE 123

Ohio. Dayton. Foeerstc, coll. Ellis, N. Am. Fungi, No. 130°, as G. limbatus. (N. Y. Bot. Card.
Herb, and Path and Myo. Herb.)

Norwood. Lloyd, coll. l.loyd Herb.).

Chillicothc. Hani, coll. (Lloyd Serb.).

Preston. Morhan, coll. (Lloyd Herb., labelled G. limbatus by Morgan).
Missouri. Saint Louis. I ledgecock, coll. (Lloyd Herb.).
Wisconsin. Milwaukee. Brown, coll. (Lloyd Herb.). Buttons.

Cedarburg. Panly, coll. (Lloyd Herb.).
Ontario. Regina. Willing, coll. (Lloyd Herb.).
Mexico. Ross and Painter, coll. (U. S. Nat'l. Herb.).
Japan. Sapporo. Miyabe, coll. (Lloyd Herb.).

Geaster Hieronymi Hennings

Plate 116

Buttons embedded in the substratum, apparently slightly pointed; outer surface
covered at first with sandy earth or trash which may later strip off in part as a layer.
Rays expanded or the tips incurved, pliable, about 5-8 in number, center concave below;
fleshy layer adnate, thin and brown. Spore sac 1.5-2 cm. broad, subsessile or with a
short stalk, dark brown, harshly asperate with short, setose hairs. Peristome rather
indefinite (not sulcate), silky, mouth fimbriate.

Spores (of plant from Jalapa, Mexico), small and minutely warted, 2.6-3.6^ thick,
often seeming slightly angular.

Except for the asperate spore sac the plant is very like G. rufescens or G. fimbriates
and seems certainly to belong to their group.

\Ye have seen the type collection of G. Hieronymi in the Lloyd Herbarium, and
find that it agrees closely with the Mexican plants. There are also two other lots in
the Lloyd Herbarium labelled G. Hieronymi from Brazil (Rick, coll.). One of these is as
described above and like the type ; the other is something different, with a sulcate mouth.

Illustration: Lloyd. Myc. Works, pi. 229, fig. 2347.

Mexico. Jalapa. Moist forest, alt. 5000 ft., Dec. 1909. Murrill, No. 244. (N. Y. Bot. Gard. Herb.)

Brazil. Rick, coll. (Lloyd Herb.).

Argentine. Hieronymus, coll. (Lloyd Herb.; type).

South Africa. Miss Duthie, coll. (Lloyd Herb. J.

Geaster Lloydii Bresadola

Plate 117

Buttons evidently subterranean or nearly so (submerged in humus), about 1.5 cm.
thick when dry, spherical, not pointed, woolly felted all over or with spots where the
softer wool has collapsed between the stiff er groups of hairs ; largely covered with bits
of leaves and trash, in some places nearly clean; the central point below attached to
obvious mycelial strands. Rays six or more, thin but rather rigid ; center concave below
usually; the thin tomentose outer layer with a strong tendency to peel off except at
tips, as in G. limbatus (sense of Bresadola) ; fleshy layer deep reddish brown to nearly
black, rimose, adherent. Spore sac sessile, thin, collapsing, dark brown with paler
areas at times or rarely pale all over, surface dull, minutely granular velvety, almost
exactly the color and surface of our G.fornicatas; mouth large, fimbriate; peristome not
defined, concolorous. The peeled off outer layer is delicate and pliable, and is usually
torn into strips. As it pulls away from the fibrous layer it leaves a narrow, root-like

124 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

core projecting downward from the central point where the mycelium was attached.
(This core is lacking in No. 8702 from Porto Rico.)

Spores (of No. 7436) sometimes faintly angular as well as minutely warted, 3.5-4.2^
thick. Capillitium threads up to 7.4ju thick, strongly encrusted.

We have not seen the types, but as our plants agree almost perfectly with the
description of G. Lloydii, we refer them to it with confidence (see Myc. Notes, p. 50.
1901). Lloyd, who collected the types in Samoa, later merged the species with velutinus
(Geastrae, p. 35). If our determination is correct, we think this a good species.

The species is in the rufescens-fornicatus group and almost exactly intermediate
between them in habit. The outer layer separates, not as in velutinus, beginning at the
tips, but as in fornicatus, radicans, and limbatus, beginning at the center. It is dis-
tinguished from fornicatus by the imperfectly fornicate habit and by the strigose and
much less trashy surface. The spores are larger than in G. Hieronymi and the spore
sac is not asperate; G. fimbriatus differs in more saccate base, smaller spores, entire
absence of hispid hairs, and in usually lighter color. In an ample collection from
Florida, consisting of 8 buttons and 12 open plants, all but one open plant were very
dark; another collection of a single plant from the same section was pale brown.

This species, as well as G. fimbriatus and Astraeus hygrometricus, shows that in
some, if not all, Geasters with so-called universal mycelium, the true mycelium is really
only basal and the felted "mycelial layer" only an outgrowth from the button. Ex-
panded plants of G. fornicatus often show a hole in the bottom of the sac-like base as if
it had been pulled away from a firmer basal attachment.

Florida. Gainesville. On an old stump. Weber, coll. 1923. (U. N. C. Herb., No. 7436, and Univ.
of Florida Herb.)

Porto Rico. Seaverand Chardon, colls. (N. Y. B. G. Herb, and U. N. C. Herb.). Outer tomentose
layer almost entirely stripped off but fragments present. Spores dark, minutely warted,
3.5-4/i. Capillitium threads paler than the spores, up to 7.5/j thick.

Geaster fornicatus (Huds.) Fr.
G. marchicus Hennings

Plates 70 and 115

Plants of medium to large size, the outer peridium splitting beyond the middle
into four (usually) to five (rarely more) lobes which bend strongly backward and down-
ward, the tips remaining attached to an equal number of shorter lobes on the margin of a
membranous cup, sunken in the earth and trash, which, as the outer layer of the button,
was stripped off from the middle layer when evagination occurred. This cup is always
completely covered with adhering trash and earth due to the flocculent mycelium weft-
ing it. Rays firm, the fleshy layer dark brown and adherent or partly peeled off, in old
plants worn away; the exposed fibrous layer whitish tan. Inner peridium seated on a
distinct, whitish, short stalk about 2-3 mm. long (in our plants), subglobose or urn-
shaped to depressed, constricted near the base so as to form a ring; surface dark brown,
finely velvety; mouth large, slightly elevated, fibrous lacerated, wrinkled, without a
distinct peristome, but the paler brown fibers in older plants have a distinct limit
exactly as in G. limbatus (sense of Hollos), which is very near. Columella not large,
about 3 or 4 mm. high and 2.5 mm. broad (in the Smith's Island plant).

Spores spherical, blackish brown, distinctly warted under high power with rather
regular, blunt warts, 3.7-4.5^j thick. Capillitium threads 3-7. 5/j. thick, about as dark
as the spores; walls roughened.

PLATE 70

Geaster PORNICATDS. Smith's Island
( lEASTEE PMBRIATUS. Xo. 7093 [middle four dry plants];
No. 7097 [lower three fresh plants]

LYCOPERDAI EAE 125

This is a rare plant in the United States and is reported by Lloyd only from Texas
and Catalina Island. In the rest of the world it is supposed to be of wide but erratic
distribution (Lloyd, Lycoperdaceae of Australia, etc., p. 21). The report by Curtis
that it is common in this state (Catl. Plants of North Carolina, p. 110) almost certainly
refers' to what we now call G. coronatus, as all the plants in his herbarium under G.
fornicatus are the latter species. Our single collection from this state consists of three
plants in fine condition that grew under cedar, but the species is reported as growing
under both conifers and frondose plants. It is easily distinguished from G. coronatus
by the absence of an outlined peristome and the entirely different surface of the inner
peridium . The species is really most nearly related to G. rufescens which is very similar
in fundamental characters, as mouth, surface of inner peridium, separating layers of
the outer peridium, and concave base when expanded. The spores, too, are very
similar. The two species, however, are not forms of the same thing, but are easily
distinguished by the fornicate habit, fewer and more rigid lobes, and darker and more
densely velvety inner peridium of G.fornicatus. Geaster marchicus is usually considered
a synonym of G.fornicatus, and plants so determined by Hollos in the U. S. National
Herbarium are certainly this species, as are also those determined by him and repre-
sented by No. 950 of J. Wagner's Kryptogamae exsiccatae. Spores of the latter are
dark, minutely warted, 3.6-4^. For other supposed synonyms, see Hollos, p. 154.

Illustrations: Batsch. Elench. Fung., pi. 29, fig. 168.
Corda. Icon. Fung. 5: pi. 4, fig. 43.
Destree. 1. c, pi. 6, fig. B.
Hollos. 1. c, pi. 8, figs. 9-18; pi. 29, fig. 26.
Lloyd. Myc. Works, pi. 96, figs. 1-3 (fig. 2 was first published as G. fenestrates in Myc. Notes,

p. 70, fig. 33).
Massee. Brit. Fungi and Lichens, pi. 35, fig. 5.
Petri. 1. c, fig. 44, 2.
Rea. 1. c, pi. 17.
Smith. 1. c, pi. IS, fig. 2.
Sowerby. Eng. Fungi 2 : pi. 198.

Smith's Island. In a dense bed of cedar twigs under a cedar in sandy soil, December 29, 1921. Couch
and Grant, colls. (U. N. C. Herb.).

Geaster fimbriatus Fr.
G. tunicalus Vitt.

Plates 70 and 115

Plants small, in our collections up to 3 cm. wide when freshly expanded; inner
peridium about 9-14 mm. thick when fresh, subglabrous to minutely dotted, pale or
dark brown, sessile in and surrounded by the concave lower half of the outer peridium;
mouth on a more or less elevated, concolorous or paler cone, but without a definitely
outlined area around it, the margin fibrous-flocculent and at times lacerate, not sulcate.
Outer peridium splitting about half way down into 5-8 lobes which when freshly ex-
panded are strongly recurved around the margin of the bowl-shaped base, usually
remaining recurved in the dry state, not hygroscopic. Mycelial layer very thin with
wefted surface, pale tan or buff to soaked ochraceous when fresh, tending to peel away
in part from the rays but as a rule not nearly so much so as in G. velulinus.^ Fleshy
layer pallid (nearly white to pale smoky tan; not pinkish) and about 2 mm. thick when
fresh, drying down to an extremely thin, smooth or finely cracked up, brown or ochra-

126 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

ceous membrane or scurf which disappears only after much exposure. Buttons quite
spherical, not pointed, completely covered by humus and trash and retaining trash
over the whole or most of the surface after opening. Columella in the mature plant
represented by a small, subspherical, pale core some distance from the base.

Spores (of No. 7093) dark smoky brown, spherical, very minutely asperulate,
many appearing smooth except under high power, 2.2-3.4//, most about 3/i. Capillitium
threads unbranched, about 5m thick, sometimes up to In, usually lighter than the spores,
but varying in color in plants of the same colony.

This small Geaster grows usually in groups of several, not truly cespitose (rarely
crowded in twos or threes), in partly rotten leaves and trash under cedars or in mixed
woods. As is to be expected, specimens are found (rarely) that are not saccate but have
the outer peridium concave below. It also happens occasionally that the outer layer
may peel off completely. For both of these variations see plants from Lynn, England
(Plowright, coll.), at the New York Botanical Garden.

In American herbaria this is often referred to G. saccatus. Lloyd thinks that G.
fimbriatus does not occur in America (The Geastrae, p. 37), though reported by Morgan
and others. However, as our plants agree clearly with this species in all important
points (indeterminate mouth, splitting of the outer peridium into two layers in part,
etc.) , we see no reason not to refer them to it. In cases where the paleness of the mouth
area is most pronounced as in plants from Buck Hill Falls, Pa. (see below) , some authors
might be inclined to call it definite, but there is no structural boundary and except for
color the mouth is exactly as in other forms. The European plant also has a tendency
to a paler mouth area. As Lloyd says, the mouth is no more fimbriate than in some
other species. The true European G. fimbriatus is represented by plants in Washington
(Path, and Myc. Herb.) from lower Austria collected by Zahlbruckner and determined
by Keissler. They are just like our plants. We have received from Patouillard another
typical European specimen (Jura) with spores minutely warted, 2.5-3.5/i thick. Still a
third (N. Y. Bot. Gard. Herb, from Herb, of Thiimen) has spores a trifle larger (3.4-4/u)
but is otherwise identical. It grew in coniferous woods and still shows fir leaves
attached.

A European collection from Sydow (Flora Marchica, Charlottenburger Schloss-
garten, P. Sydow, coll.) labelled G. fimbriatus and now at the New York Botanical
Garden is not the plant we here describe, but G. lageniformis with much rougher spores
and determinate mouth. Our plants agree with Rea's description of G. fimbriatus
except in size of spores which he gives as 4-5/j. It is very likely that G. triplex (with
G. lageniformis) and G. fimbriatus have been often confused.

Geaster rufescens is the nearest relative of G. fimbriatus, and large plants of the latter
might easily be referred to it. The thinner fleshy layer, more saccate form, and smaller
average size distinguish G. fimbriatus, but there may be connecting forms. Geaster
tunicatus as represented in Saccardo, Mycotheca italica, No. 1625 (Mendola; Bresadola,
coll.), is G. fimbriatus. The mouth is indefinite, the base saccate. The two species are
also treated as synonyms by Hollos.

Illustrations: Berkeley. Outlines Brit. Fung., pi. 20, fig. 4.

Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 6, fig. 36.

Destree. 1. c, pi. 10, fig. A.

De Toni. 1. c, pi. 63, fig. P.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 31.

LYCOPERDACEAE 127

Hollos. 1. c. pi. 10, figs. 17 and 18.

Lloyd. ThcGoastraclig. 74.

Muhaol. l'ulmr f. l'il/.freunde 2: No. 202, figs, a and b. 1918 (2nd ed).

Morgan. North American Gcastcrs, fig. 8.

Petri. 1- c., tig. ■!-■

Rea. 1. c, pi. 19.

Rolland. Champ. France, pi. 1 10, fig. 252.

Schmidel. Icon, riant., pi. 50, figs. 1-4. 1747.

Smith. Grevillea 2: pi. 17, fig. 2.

Soweiby. Engl. Fungi, pi. 80.

1291. In cedar twigs and oak leaves mixed at the base of a cedar tree, September 30, 1914. Spores
minutely asperulate, up to 3.7m thick.

7092. Under cedar tree, August 24, 1923. Spores 2.8-3. 7/i. Capillitium threads thick-walled, nearly

hyaline, up to 6.6m thick.

7093. In rotting frondose leaves, August 24, 1923. Spores dark under the microscope, surface as

usual, 2. 2-3. 4^ thick.
Also Nos. 4096, 7097, 7135, 7439, 7476, 7480.

Statesville. Beardslee, coll. (Lloyd Herb., as G. saccatus).

South Carolina. Society Hill. (Curtis Herb., asG. fimbriates, No. 3833.)

Tennessee. Unaka Springs. Murrill, coll. (N. Y. Bot. Gard. Herb.).

Pennsylvania. New Garden. (Michener Herb., as G. minimus.)

New Hampshire. Shelburne. Farlow, coll. (Farlow Herb.).

Wisconsin. Eagle Heights, near Madison, August 26, 1903. No distinct peristome. Spores very

faintly spiny, 3-3. 7M thick. (U. N. C. Herb.)
Ohio. Miami Valley. Morgan, coll. (N. Y. Bot. Gard. Herb.).

Colorado. Minnehaha. Clements, Crypt. Form. Colo., No. 194, as G. limbatus (in part). (Farlow
Herb.)

Denver. Bethel, coll. (N. Y. Bot. Gard. Herb.).
Ontario. Sarina. Bertolet, coll. (Lloyd Herb., as G. saccatus). Also collection from Canada by
Deamess. (U. N. C. Herb.)

Geaster florifonnis Vitt.
G. delicatus Morg.

Plates 68, 115 and 116

Plants (typically) small, with a spread when expanded of 2-2.5 cm., submerged
until mature ;"rays about 8-10, unequal, hygroscopic and expanding quickly when put
into water, covered at first with sandy earth held on by mycelium, this flaking off by
degrees to leave the surface smooth and pale after exposure; fleshy layer dark brown,
adnate, continuous then rimose; inner peridium about 6-8 mm. thick, subglobose or
depressed, sessile, pale, covered at first with fine granules or furfuraceous particles which
soon wear away in large part; mouth only a puncture or slit with short radiating fissures
with no defined or silky area around it as a rule, after a time the margin becoming
fimbriate.

Spores (of Clements, No. 613) distinctly warted, spherical, 5-7^ thick, or slightly
elongated and about 6.5 x 7 An. Capillitium threads straight, thick-walled, up to 7^
thick.

The American plants are like those of a good collection from Hollos (Hungary)
sent us through the kindness of Patouillard. In these plants the peristome is indefinite,
rays strongly hygroscopic, spores spherical or somewhat irregular, distinctly warted,
5-7 x 6-8/i. Similar plants from Hollos have been seen by us in the Lloyd Herbarium.

128 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Good collections of this species from San Diego, California, in Washington and
New York show an interesting variation in the mouth. In the plants at the New York
Botanical Garden the two plants with mouths visible show a depressed zone around
the mouth but the surface of the zone is not silky as in most Geasters, but of the same
texture as the remainder of the surface. In one of the plants the mouth was an elevated
Dapilla in the center of this zone; in the other it was a plane opening. For a similar
variation see plants from Florida and North Dakota entered below. The plants in
Washington show a plane opening without a depressed zone. In most herbarium
material the spore sac is nearly smooth or faintly powdery. The granules when present
are not so large as in G. asper. The rays when dry are either folded over the top of the
inner peridium or are curved in more abruptly with their tips under it, or they may take
any intermediate position. The species grows in sandy soil and in this country is
apparently confined to the western states, occurring also in Europe and Africa. A
large form of this with the spore sac 2 cm. thick (in the pressed condition) is represented
in the Herbaiium of the Museum of Paris from western Africa. Except in size they
do not differ apparently from the American plants. The spores are distinctly warted,
5-6.5 x 6-7(x. An interesting plant that seems referable to this as a variant is repre-
sented in the Far low Herbarium from Lake Titicaca, Bolivia (Mrs. Shepard, coll.).
The rays are thick, strongly hygroscopic, covered with sand; spore sac densely furfura-
ceous, nearly white, peristome vaguely defined by a low ridge, fimbriate; spores spherical
to irregular, distinctly warted, 4-5/u thick or up to 6// long.

In his treatment of G. delicatus (The Geastrae, p. 11), Lloyd makes the mistake of
referring to G. lageniformis when he should have said G. florijormis (see Hollos, p. 67).

Illustrations: Cunningham. Trans. N. Z. Inst. 57: pi. 11, fig. 37. 1926.
De Toni. 1. c, pi. 1, fig. O.

Hollos. 1. c, pi. 10, figs. 20-25; pi. 29, figs. 29, 30.

Lloyd. The Geastrae, figs. 14, 15 (as G. delicatus) and fig. 78; also pi. 98, figs. 12-18.
Morgan. The Genus Geaster, fig. 2 (as G. delicatus).
Petri. 1. c, fig. 43.
Vittadini. Monog. Lycoperd., pi. 1, fig. 5. 1842.

Florida. Grasmere. Baker, coll. (Lloyd Herb.). Mouth indistinct. Two other boxes of same show

very minute plants, many not over 4-5 mm. in diameter when closed; many with depressed

mouth area.
Alachua Co. Couch, coll. (U. N. C. Herb., No. 7274).
Texas. Denton. Long, coll. (Lloyd Herb.). Columella cylindrical, about 4 mm. long.
California. San Diego. Orcutt, coll. (Path, and Myc. Herb, and N. Y. Bot. Gard. Herb., as G.

delicatus). Spores subspherical, about 5-6.8^ thick.
Catalina Island. Mrs. Trask, coll. (Lloyd Herb.).
South California, 32-36° N. Lat. Parry, coll. (Curtis Herb., as G. rufescens).

Colorado. Yuma. Clements, No. 613. (N. Y. Bot. Gard. Herb.) Spores sacs distinctly granular.
North Dakota. Brenckle, coll. (Path, and Myc. Herb, and U. N. C. Herb., No. 152 of Sydow, Fungi

Exot. Exs.). Spores spherical or somewhat elongated, 4.2-5 x 5-7/x. In Lloyd's Herbarium

plants from N. Dakota (Brenckle, coll.) show a more or less definite, depressed area around the

mouth.
Nebraska. Lincoln. Webber, coll. (N. Y. Bot. Gard. Herb.; also in Path, and Myc. Herb, as Ell.

& Ev., N. Am. Fungi, No. 1941).
Washington. Cheney. Mrs. Tucker, coll. (Lloyd Herb.). Lloyd has a note saying, "Mrs. Tucker

writes me that this species proves quite a pest in flower beds."

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LYCOPERDACEAE 129

Canada, London, nearness, coll. (TJ. N. C. Herb.).

Transvaal. Pretoria. Evans, coll (Lloyd Herb.).

New Zealand. Rangitata. Barker, coll. (Lloyd Herb.).

Geaster coronatus (Schaeff.) Schroet.

G. minimus Schw. (.not Gcastrum minimum Chevallier)
G. marginatus Yitt.
G.quculrithluni Pers.
G. Cesalii Rabenhorst

Plates 71, 72, 115 and 116

Tlants when expanded 1.5-3.5 cm. broad and up to 3.3 cm. tall excluding the basal
cup when present; outer peridium splitting into 4-8 strongly recurved or rarely hori-
zontal lobes, the mycelial layer often being stripped off and remaining as an obscure
weft which holds together a hollow mass of earth or trash out of which the plant was
lifted, the tips of the rays attached to its margin (typical or fornicate form); in other
cases the mycelial layer with the adhering trash is lifted with the plant and forms a
more or less' persistent coat on the outer surface (form minimus) ; fleshy layer brown,
cracking and often seceding in large part from the rays, the central part most persistent
and at times forming a low collar around the base of the stalk. Inner peridium ovoid
to oblong, 4-14/j thick, brown to umber or slate color, the surface, except when weather-
worn, powdered with minute glistening particles; stalk distinct, about 1.5-2.5 mm.
long and expanding above into a ridge which in some specimens may have an acute rim
like a little collar. Mouth delicately fimbriated, not sulcate, low, abruptly or gradu-
ally elevated from a nearly flat, sharply defined silky area which is outlined by a groove.
Columella in mature plant not obvious.

Spores (of No. 3881) dark smoky purplish, distinctly rough, 4-5. 1/x thick, most
about 4.5/1. Capillitium threads up to 6m thick, not sinuous and branching rarely if
at all.

This species has been confused with G. fornicatus both in Europe and America
and it is treated under the latter name by Morgan. We are following Lloyd and Hollos
in using the name G. coronatus for the present species. The true G. fornicatus, as now
understood, is usually larger and is easily distinguished by the absence of a well defined
mouth area, the different surface of the spore sac, and by the smaller and less strongly
warted spores.

The type of G. minimus in the Schweinitz Herbarium is just like the usual G.
minimus of herbaria; the spores distinctly warted, 3.7-4.4ju thick. Another authentic
specimen of G. minimus from Schweinitz, now in the Michener Herbarium in Washing-
ton, is just like the form that we are calling G. minimus. Lloyd has seen the plant in
the Schweinitz Herbarium and says that it is the same as the minimus form. We find
it impossible to distinguish between G. coronatus and G. minimus. If the mycelial
layer with its trash remains attached and is lifted when the rays bend back we have
the minimus form, if not it is the coronatus form, and both forms with intermediates
may occur in the same colony. Smaller size does not distinguish minimus, as we have
seen the fornicate form with the inner peridium less than 5 mm. thick.

In several plants in our collections the ridge around the expanded top of the stem,
which is often obvious in stalked species, is terminated by a distinct, fimbriated frill,
recalling the broader frill of G. Bryantii. The last mentioned is distinguished by its
sulcate and more narrow and elevated mouth, by the distinct columella and by the

130 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

broader frill. Both forms of G. coronatus are widely distributed in the United States
and are not uncommon. In Europe the minimus form seems much rarer. The
European plant is represented in Torrend's Mycotheca Lusitanica No. 79 (in pinetis),
also in Cesati's Herbarium Mycol., No. 1634 (as G. Cesatii. Both in Path, and Myc.
Herb., Washington). In G. Cesatii the spores are distinctly warted, 3.8-4.5^ thick.

Geaster granulosus is an extreme form or variety of G. minimus with distinctly
larger granules on the spore sac and in some cases covering it almost solidly with a
white crust, at least as much so as in G. calceus as shown by Lloyd's fig. 9, pi. 95.
We have studied authentic specimens of G. granulosus from Austria (Fuckel, coll.) and
find the spores to be larger and rougher than in G. minimus, 4.2-5. 6yu thick, often with
an angular appearance from the irregular warts (pi. 116, fig. 12). A plant from Ymala,
Mexico, has the exact superficial appearance of G. granulosus and the spores approach
those of that species, with the same irregularity, but are somewhat smaller, 3.8-5/u.
They have the form of spores of G. minimus shown on pi. 1 15, fig. 24, and are only slightly
larger. Geaster calceus Lloyd (Myc. Notes, p. 311) from South Africa is very near the
last, but with smaller spores (3.5-4.2p), of regular, spherical shape and with equal warts
(pi. 116, fig. 11).

A peculiar plant is represented in the Path, and Myc. Herbarium by three speci-
mens from Waste Land, South Dakota (Wilcox, coll.). It looks like a small, nearly
sessile specimen of G. minimus but the spores are distinctly larger and irregular (4.8-5.5 x
6.4-7.5^), with large, unequal warts (pi. 116, fig. 16). The plant bears the ms. name
of G. lerreus Ellis, n. sp., with description in ms. attached. It may be a new species,
but is more probably an extreme variant of the minimus type. Macbride describes as
a new species, G. juniper inus, a plant from Iowa, very close to G. coronatus. It seems
scarcely more than a form (Mycologia 4 : 84. 1912).

Noack (cited below) reports this fungus, which he calls G. fornicatus, as forming
mycorrhizas on the roots of conifers.

Illustrations: De Toni. 1. c, pi. 63, fig. H and fig. I (as G. marginatus).
Fries, Th. C. E. Sveriges Gasteromyceter, fig. 28 (as G. quadrifidum) .
Hard. Mushrooms, fig. 482 (as G. minimus).

Hollos. 1. c, pi. 7, figs. 11-14 (as G. coronatus); pi. 10, figs. 8-10; pi. 29, fig. 28 (as G. minimus).
Lloyd. The Geastrae, figs. 50-52 (as G. minimus); figs. 58-61 (as G. coronatus; fig. 59 was first
published as G. fornicatus, Myc. Notes, p. 71, fig. 36). Also pi. 95, figs. 1-7 (as G. minimus).
Macbride. Mycologia 4: pi. 62, fig. 2 (as G. marginatus) and fig. 3 (as G. minimus). 1912.
Morgan. North Amer. Geasters, fig. 1 (as G. fornicatus) and fig. 7 (asG. minimus).
Nees von Esenbeck. Syst. Pilze Schw., pi. 12, fig. 128 (as Geastrum quadrifidum). 1817.
Noack. Bot. Zeit. 47: pi. 5, fig. 1 (as G. fornicatus). 1889.
Petri. 1. c, fig. 37, No. 1 (asG. minimus); fig. 44, No. 4 (asG. coronatus).
Schaeffer. Fung. Bavar., pi. 183.

Schmidel. Icon. Plant., pi. 37 figs. 1-3 (as G. coronatus). 1747.
Vittadini. Monog. Lycoperd., pi. 1, fig. 6 (asG. marginatus). 1843.

75. On mossy shaded ground under cedars, November 10, 1911. Fornicate form. Spores 3.7-4.4/1,

most about 3.9m thick. Capillitium threads sinuous, up to 5.5ji thick.
3881. By a decaying oak stump in Strowd's low grounds, December 13, 1919. Fornicate form and

minimus form mixed insame colony.
6065. By a cedar stump in a pasture, February 12, 1923. Fornicate form. Spores distinctly warted,

4-5.3m, most about 4.6p thick. Capillitium threads straight, apparently unbranched, up to

5.1/i thick.

PLATE

Geaster coroxatus. No. 3881
Fresh condition

LYCOPERPACEAE 131

6091. On the ground by a decaying oak stump, May 20, 1923. Fornicate form. Spores distinctly
MOted, 4-5. 1m thick. Capillitium threads straight, apparently unbranched, up to 7/i thick.
Also Nos. o84. 5903, 6074, B077, 7403, 7484.

Raleigh. Wells, c.-ll 0 N (..Herb.).

Smith's Island, Couch and Grant, coll. No. 5965. In sandy soil, December 29, 1921. Minimus
form. Spores minutely warted, 4-5/i.

"Carolina." Pine woods. M ichener Herb, from Schw. Herb., as G.fornicatus and G. strialus.) Also

in Schw. Herb.. No. 2247, as (.'. tiiuuirindiini.
South Carolina. Aiken. Ravenel, coll. (Path, and Mve. Herb., as G. minimus, Fung. Am. Exs.,

No. 472). Also Fung. Car. Exs., No. 74. (Phila. Acad., as(.'. minimus.)
Society Hill, (Curtis Herb., as G. fornicatus, No. 2301.)

Also a collection from South Carolina by Ravenel in lUichener Herb., asC. minimus.
Florida. DeFuniak Springs. Fisher, coll. (Lloyd Herb.). These plants have a distinct apophysis

with a distinct edge as in one or more of our Chapel Hill collections, but is not exactly typical

G. coronoJus as the sac particles arc dull and look more like specks of dust.
New Jersey. In very sandy soil. Ellis, coll. (N. Y. Bot. Card. Herb.). Lobes varying from 4-8;

an imperfect cup attached in some, in others not.
New York. Mohawk. Mrs. Lobcnstcin. (N. Y. Bot. Card. Herb., as G.fornicatus.)
Massachusetts. Boston. Sprague, No. 42. (Curtis Herfa., aa G. mittimus.) There arc also two typi-
cal specimens of minimus from Schweinitz Herb, in the Curtis Herb.
Ohio. Lloyd, coll. (N. Y. Bot. Gard. Herb., as(.\ minimus, and U. N. C. Herb.). Spores distinctly

warted, 3.8-5.5/1 thick. Capillitium threads straight, up to 5.5/i thick.
Michigan. Agricultural College. Wheeler, coll. (Path, and Myc. Herb., as G. minimus).
Wisconsin. I Univ. Wis. Herb., asG. minimus, and U. N. C. Herb.) Spores distinctly warted, 3.7-4.4/1

thick.
Texas. Wright, No. 326. (Curtis Herb., as G.fornicatus.)
California. Pasadena. McClatchie, coll. (N. Y. Bot. Gard. Herb.).
Canada. Macoun, coll. (N. Y. Bot. Gard. Herb., as G. minimus, and U. N. C. Herb.). Minute plants

of both forms. Lobes varying from 4-8, rcvolutc, the earth mass attached in some.
Jamaica. A. E. Wright, coll. (Farlow Herb., as G. limbatus). One is covered with coarse white

particles like Lloyd's G. calceus.
Mexico. Ymala. (Path, and Myc. Herb.) Minimus form with large granules like G. granulosus.

Spores 3.8-5/1.
Ecuador. Millc, coll. (Lloyd Herb.).
Japan. Sapporo. Miyabe, coll. (Lloyd Herb).
New Zealand. Cunningham, coll. (U. N. C. Herb.). Spores strongly warted, 4.2-5.5/1 thick.

GeasterleptospermusAtk. & Coker

Plates 73 and 115

Inner peridium nearly globular, 2.5-5 (rarely 6.5) mm. thick, pale gray to pale tan
in color, minutely dusted with fine whitish particles which also cover the inner surface
of the freshly opened rays; mouth very small, minutely fibrous, surrounded by a conical,
white disc which is distinctly outlined and radially fibrous (not sulcate). Fibrous
layer of the outer peridium splitting into 3-6 rays about half way down or more, the
rays recurving and the base becoming arched up (vaulted beneath), leaving the delicate,
fibrous, mycelial layer as a distinct, membranous, white cup beneath, with its margin
also rayed by slits, the rays attached to the rays of the plant above (typically fornicate).
Inner surface of the rays when freshly opened covered with a succulent, pale buff,
fleshy layer, which when dry forms the thin, tan or light brown, smooth and nearly
complete membrane over the fibrous layer; underside of the rays white and smooth.
Columella not obvious.

132 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Spores (of No. 2028) minute, spherical, 2-3n, usually appearing under high power
as if very minutely punctate or asperulate. Capillitium threads unbranched, about
3m thick. Both spores and capillitium very pale yellow-brown (whitish).

This is a very distinct and well marked little species and is the smallest known
Geaster, the inner peridium of a full-grown plant seldom reaching over 5 mm. in di-
ameter. They have the unique habit of growing in moss on tree trunks, as cedar,
hickory, elm, etc., and usually from about four feet from the ground up, most of our
collection coming from a height of ten to twenty feet. It is not at all uncommon in
Chapel Hill, which is the only known station, and is distinctly a cold weather species,
appearing in mild wet spells in winter and early spring and often associated with Lyco-
perdon acuminatum. The flocculent white mycelium penetrates the old bark and ex-
tends itself abundantly between the planes of cleavage. The species is distinguished
from G. coronatus, which is nearest, by the much smaller average size, smaller and
nearly white spores, and capillitium, and peculiar habitat. For the original descrip-
tion, see Bot. Gaz. 36: 306. 1903.

681, 1085, 1479, 1537, 2028, 2029, 2466, 3222, 3231, 6052, 7586. All on mossy bark of living elm, hickory
or cedar during winter or early spring.

Geaster pectinatus Persoon
G: multifidum var. a Pers.

Plates 74, 75 and 115

Mycelium universal and forming a soft, flocculent coat that holds trash to the
entire outer surface of the plant as in G. rufescens; rays few to numerous, typically
reflexed but the tips often incurved, central region concave below and elevating the
spore sac; mycelial layer with attached earth tending to peel off after expansion as is
also the brown fleshy layer, so that the thin, pale fibrous layer may be left almost
naked in old specimens; spore sac subglobose to urn-shaped, with or without an apophy-
sis, the lower part with radial lines or ridges that in some cases may be quite faint;
stalk slender, typically rather long, up to 5 or 6 mm., often with a collar or ring near
center or base; surface when fresh covered with a fine powder which is whitish against
the spore sac and brown without so as to give the spore sac a whitish bloom where it is
thin and a brown-drab color where it is thicker. Mouth long, narrowly conical, prettily
sulcate-striate, brown.

Spores (of plant from Jamesville, N. Y.) very dark, with large blunt warts, 4.2-5.5/x
thick. Capillitium threads up to 7ju thick.

A rare plant apparently, and Schweinitz's record is so far the only one from North
Carolina. The length of stalk is very variable, e.g., in one good lot from Surrey,
England, at the New York Botanical Garden Herbarium (as G. limbatus), the stalk
is only 2 mm. long. The species runs into two extremes that are most conveniently
treated as species, though intermediates occur. The smallest of these is G. Schmidelii,
which also approaches G. Smithii (G. ambiguus, sense of Hollos). The other is G.
Bryantii which is distinguished by a deflexed collar around the base of the spore sac.
Specimens of G. pectinatus (Upsala) in the Curtis Herbarium are labelled G. limbatus
by E. P. Fries.

A part of the type collection of G. biplicatus (Proc. Am. Acad. Arts and Sciences
4: 124. 1860) in the Curtis Herbarium (Bonin Isles) is so pressed as not to show the
base of the spore sac; peristome truly sulcate; spore sac reddish brown with a grayish

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Geaster pectinatus, North Island, S. C.

LYCOPERDACEAE 133

powder; outer surface covered with humus held by flocculence. Spores distinctly
warted, 3.8-4.8/1, rarely up to 5.2/u thick. Other plants from the type collection in
Washington are similar with inner peridium blackish brown with a grayish powder.
We have a fine specimen from New Caledonia sent us by Patouillard (as G. plicalus =
G. biplicaius) with strongly sulcate base to the spore sac, the stalk about 4 mm. long;
spores spherical to a little irregular, distinctly warted, 4-6/u thick; capillitium straight,
up to 6.5m thick. It is probable that G. plicatus and G. triplicates are the same, as
Patouillard thinks, and both are certainly very near G. pectinatus. The deep and
regular sulcations on the base of the spore sac are a striking character that we have not
seen exactly duplicated in G. pectinatus. We have seen Lloyd's abundant specimens of
G. plicatus from Australia and South Africa, but have not seen the type.

Illustrations: Cunningham. Trans. N. Z. Inst. 57: pi. 9, fig. 27. 1926.
Cunningham. Proc. Linn. Soc. NT. S. \V. 51: pi. 3, figs. 7, 8.
Fries, Th. C. E. Sveriges Gasteromyceter, fig. 30.
Lloyd. The Geastrae, figs. 19-22 (fig. 19 was first published as G. tenuipes in Myc. Notes, p. 72,

fig. 37).
Lloyd. Gen. Gastromycetes, pi. 8, fig. 38.
Petri. I.e., fig. 37, 4-5 and fig. 40.
Rick. Broteria 5: pi. 1, figs. 14-15.

Schmidel. Icon. Plant., pi. 37, figs. 11-14. Edition of 1793. These are referred to by Persoon.
Smith. Grevillea 2: pi. 17, fig. 1 (as G. limbatus). Ridges on mouth obscure.

North Carolina. Salem. (Schw. Herb.) Plant as large as usual in the species; stalk short and rather

thick.
South Carolina. North Island. Under cedars. Coker and party, colls. (U. N. C. Herb.).
New York. Jamesville. Mrs. Goodrich, coll. (N. Y. Bot. Gard. Herb.).

Pompey. (N. Y. Bot. Gard. Herb., as G. Bryantii.)
Ohio. (N. Y. Bot. Gard. Herb., from Lloyd Herb.)
Reported from Florida and Pennsylvania by Lloyd.
Ontario. Hamilton. Dearness, coll. (U. N. C. Herb.). Spores strongly warted, 4.5-6p.

Geaster Bryantii Berk.

Plate 115

Like G. pectinatus except that around the base of the spore sac at some distance
from the stem there hangs a collar-like ring with an acute edge. This character alone
would hardly entitle the plant to more than varietal rank.

Spores of a plant from Massee (England [?], N. Y. Bot. Gard. Herb.) are spherical,
distinctly warted, very dark, 4-5.5/1, most about 4.8/u. In another plant from Europe
(Rabenhorst and Winter, Fungi europaei, No. 2639, N. Y. Bot. Gard. Herb.) the spores
are smaller and less distinctly warted, 3.8-4.8^ thick. The plant is also well represented
in Sydow's Mycotheca germanica, No. 1449 (Path, and Myc. Herb., Washington).

The plant is rare in America and is not reported from the southeastern states.
Lloyd reports it from Maine and Texas, and Morgan from New York.

Illustrations: Berkeley. Grevillea 2: pi. 16, fig. 2.

Bryant. An Historical Account, etc., fig. 19. 1782.

Cunningham. Proc. Linn. Soc. N. S. \V. 51: pi. 3, fig. 10.

DeToni. 1. c, pi. 1, fig. D.

Hollos. 1. c, pi. 9, figs. 1-4.

Lloyd. The Geastrae, figs. 23-26.

Morgan. North Amer. Geasters, fig. 5.

Rea. I.e., pi. 18.

134 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Geaster Schmidelii Vitt.
G. Rabenhorstii Kunze

Plates 75 and 115

Like G. pectinatus, except smaller and with a short stalk and less separable mycelial
layer and smaller spores. A typical lot of this form is No. 5020 of E. Bartholomew's
Fungi Columbiana (Calloway, Nebraska; J. M. Bates, coll.). These have 5-6 reflexed
rays with trashy mycelial layer attached, the thin brown fleshy layer also present;
spore sac suboval, very short-stalked, smooth, dark brown, mouth elevated, strongly
sulcate. Spores medium dark, minutely warted, spherical, 3.7-4.4u, with a visible oil
drop.

Another lot of Bates' plants from Nebraska are in the New York Botanical Garden
(from Lloyd Herb.). The outer surface is covered with sand, the spore sac blackish
with a whitish or brown powder. A less typical lot is from New Jersey (Newfield;
Earle, coll., at N. Y. Bot. Gard. Herb.) . This lot has the rays slightly hygroscopic and
in one of the plants the spore sac is yellowish buff color. Another plant of this lot is
the one shown by Lloyd in his Geastrae, fig. 27 a. The spores of this particular plant
are like those noted above, minutely warted, 3.7-4.5;*. A plant about intermediate
between pectinatus and Schmidelii is at the New York Botanical Garden Herbarium
(Underwood, coll.; Kirkville, N. Y.). The spore sac is buffy and the stalk very short.
European G. Schmidelii as distributed by Torrend (Mycotheca Lusitanica, in Path,
and Myc. Herb., Washington) is like our American plant and like the types of G.
Rabenhorstii. Three plants sent us for examination by Patouillard (Herb. Mus.
Paris; Barla, coll.) as G. Schmidelii are somewhat worn specimens of the minimus form
of G. coronatus. The spores are spherical, warted, 3.7-4.6^.

In freshly opened plants collected by us in Warrenton, Va., the fleshy layer was
nearly white; spore sac drab. On drying the fleshy layer turned brown and cracked
considerably.

. Spores of the type material of G. Rabenhorstii (Fungi selecti exs., No. 10, in Path.
and Myc. Herb., Washington) are spherical, distinctly warted, 3.8-5^, a few up to
5.5^. Geaster elegans Vitt. is very near G. Schmidelii, the only obvious difference being
the entirely sessile spore sac and larger spores of the former. These characters are
shown by plants in Torrend's Mycotheca Lusitanica, No. 78 (Univ. Wis. Herb, and
U. N. C. Herb., as G. elegans), and in his Fungi selecti exs., No. 79 (in the same her-
barium, as G. minimus) ; also in the Herbarium of the University of Paris (from Tu-
lasne Herb.). The spores of the last mentioned are minutely but distinctly rough,
4.2-5.5^ thick (pi. 116, fig. 2). The plant is not known from America.

This would seem to run into a saccate form, apparently quite sessile as shown by a
distribution of 11 good plants by J. Kunze, Fungi selecti exs., as G. striatus DC, from
the same place as his G. Rabenhorstii and also like that in growing in Picea excelsa
needles; mouth, peridium and all else same except that the outer peridium is saccate
or flat and inner peridium with no obvious stalk.

Illustrations: Destree. 1. c, pi. 7, fig. B.
Lloyd. As cited above.
Petri. 1. c, fig. 37, 3.

Trelease. 1. c, pi. 7, fig. 3 (as G. Rabenhorstii).
Vittadini. Monog. Lycoperd., pi. 1, fig. 7.

PLATE 75

^W

» U* *

^*

*-3t

[Top] Geaster PECTIKATUS. Small plant, London, Canada, No. 3023;

larger, Webster's Falls, Canada, Xo. 4235
[Middle] Geaster Schmidei.ii. Newfield, N. J.
[Bottom] Geaster fmbilic iattjs. Newfield, N. J. (left)
Geaster Rabenhoesttj. Type material (right)

LYCOPERDACEAE 135

North Carolina. Salem. Schwcinitz, coll. (Curtis Herb., as C. pectinatus) . Spore sac practically

sessile.
Virginia. Warrenton. In upland frondose woods, Sept. 6, 1926. Coker, coll. (U. N. C. Herb.).

Spores minutely warted, 3.4— Ip.
New Jersey. Newfield. Ellis, coll. (N. Y. Bot. Card. Herb.).
Nebraska. Bates, coll. (N. Y. Bot. Card. Herb., from Lloyd Herb, and Path, and Myc. Herb.,

Washington, as Bartholomew, Fungi Columb., No. 5020).
Texas. Denton. Long, coll. (Path, and Myc. Herb., Washington; Ellis and Everhart, Fungi

Columb., No. 1620, as C. pectinatus). Mixed with these is a plant with strongly granular

spore sac like C. asper except that the outer peridium is not hygroscopic but like the others.
Cuba. Wright, No. 694. (Curtis Herb., as G. tenuipes Berk.). Spore sac apparently sessile, slate

color with a dusting of grayish powder.

Geaster Hariotii Lloyd

Plates 68 and 115

Rays about S, spreading about 3 cm., pliable, not hygroscopic, revolute or in part
expanded, the center concave below and elevating the spore sac; outer surface entirely
covered with earth or trash held on by flocculent mycelium which may in time be
partially shed in flakes. Fleshy layer dark brown, cracking and seceding in places.
Spore sac 1.5 cm. broad, subglobose, sessile, dark brown to black, distinctly pitted and
rough to asperate with small points, but not truly warted or tomentose. Peristome
truly sulcate, mouth elevated and becoming fimbriate.

Spores (of plant from Porto Rico) spherical, dark, faintly rough under very high
power, 3.5-4m thick.

As Lloyd does not mention a rough surface to the peridium (Myc. Notes, p. 311),
we had referred our plant to this species with some doubt until the recent receipt from
Patouillard of a specimen from Brazil that had been seen by Lloyd and which Patouillard
designates as an "original specimen." This is exactly like our plant except that the
surface is somewhat less conspicuously rough; rays not rigid; spores spherical, nearly
smooth, 3-3.6ju thick, often with a visible oil drop. Capillitium threads straight,
encrusted, up to 6.5/u thick. Under G. Archeri Lloyd has two collections, one from
Ceylon (Petch, coll.) and one from Victoria (Winkler, coll.). They are not that
species, but pale forms of G. Hariotii. The mouth in each case is darker than the
granular or spongy peridium; peridium clay-brown to brown; mouth truly sulcate;
outer surface densely flocculent and with trash as in G. Hariolii.

Illustrations: Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 4, fig. 12.
Lloyd. Myc. Works, pi. 99, figs. 7-9.

Porto Rico. March 1923. Seaver, coll. (Seaver Herb., No. 840, and TJ. N. C. Herb.).
British Honduras. M. E. Peck, coll. (N. Y. Bot. Gard. Herb.). Spores smooth, 3-3.7/1 thick.
Brazil. (U. N. C. Herb, from Patouillard.)

Rio. In leaf mold, March 1894. Murrill, coll. (N. Y. Bot. Gard. Herb.).
Dutch Guiana. Samuels, coll. (N. Y. Bot. Gard. Herb.).

Geaster asper (Mich.) Lloyd
G. pseudomammosus Henn.
G. campesiris Morgan

Plates 68 and 115

Plants submerged until mature, the mycelium universal; rays with a spread of
2.5-3.5 cm. (up to 5 cm., Morgan), hygroscopic, rigid, 8-10, rarely more, involute or

136 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

spreading when dry, outer surface covered at first and rather persistently with sandy
earth mixed with fiocculent mycelium, this slowly flaking off or wearing away on
exposure, leaving the surface pale and smooth; fleshy layer adnate, usually continuous,
dark brown to blackish; base concave below and elevating the spore sac which is sub-
globose with a short thickish stalk, the surface gray to brown and covered with wart-
like particles; mouth strongly and prettily sulcate, seated in a depressed zone, con-
colorous or darker, at times nearly black.

Spores (of a plant from Lincoln, Nebraska), spherical, distinctly warted, 4.5-6.5/n.
Capillitium threads straight, up to 5p thick.

A plant of the Mississippi valley and westward, seeming to prefer sandy soil. It
is well marked by the conspicuous granules on the spore sac, which are much coarser
than the fine powdery particles on the sac of G. coronatus and G. leptospermus.

This species varies to a form with lax and pliable outer peridium like that of G.
mammosus and G. Schmidelii. See, for example, one plant among a typical lot of G.
asper from Dakota at Washington (Brenckle, Fungi Dakotensis, No. 35, in Path, and
Myc. Herb.) ; also in the same herbarium a plant from Texas, mixed with G. Schmidelii
(Ell. & Ev., Fungi Columb., No. 1620; Long, coll.).

Illustrations: Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 2, fig. 1; pi. 3, figs. 13, 14; pi. 7, fig. 41
(as G. campestris).
Hollos. 1. c, pi. 9, figs. 12-14, 21, 22; pi. 29, figs. 24, 25.
Lloyd. Myc. Notes No. 7 : fig. 34; Geastrae, figs. 28-30.
Micheli. Nov. Plant. Gen., pi. 100, fig. 2.
Morgan. The Genus Geaster, fig. 1 (as G. campestris).
?Rick. Broteria 5: pi. 1, fig. 13 (as G. asper, but does not look much like ours).

Nebraska. Long Pine. Bates, coll.

Lincoln. Webber, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.).

North Dakota. Kulm, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb., as Sydow, Fungi exot.,
No. 151).

Wyoming. Pitchfork. Davis, coll. (U. N. C. Herb.).

Kansas. Bartholomew, coll. (Path, and Myc. Herb.). In this lot the mouth is nearly black, con-
trasting strongly with the nearly white peridium.

Utah. Fort Douglas. Clements, coll. (Farlow Herb.). Granules on spore sac inconspicuous, but
appearance and spores as in G. asper. Spores spherical, irregularly warted, 4.4-5.5/1, rarely
up to 611 thick.

Texas. Denton. Long, coll. (Ell. & Ev., Fungi Columb., No. 1825 ; Path, and Myc. Herb., asG. striates).

Geaster umbilicatus Fr. Sense of Morgan

G. striatulns Kalch. Sense of Lloyd
? G. Smithii Lloyd

G. ambiguus Mont. Sense of Hollos in part

Plates 75 and 115

Plants small, hygroscopic, the button subterranean and when first expanded
covered completely without by sandy earth; rays about 7-10, unequal in breadth, the
delicate, fiocculent outer layer intimately mixed with earth and gradually wearing away
to leave the rays smooth and glabrous and pale brown to pallid tan, scarcely shining
and not as metallic-looking as in G. mammosus; fleshy layer rather thin, smooth or more
or less rimose, brown to blackish. Spore sac 7-12 mm. thick, sessile, pale tan to dark
brown, minutely furfuraceous until old; peristome prettily and regularly plicate-sulcate,

PLATE 76

AsO^CKlAXjAJUbJ

■ (I ■+-

Suggested Relationships of the Geasters

LYCOPERDACEAE 137

strongly furfuraceous, particularly below, when fresh, the mouth usually elevated and
conical or at times flattened in a depressed zone.

Spores (of plants from New Jersey, Ellis, No. 3472) spherical, faintly warted,
3.5-4m thick. Capillitium threads wavy, up to 6.5m thick.

A plant of sandy soil in Europe and America and evidently rare. It is the plant
distributed by Ellis as No. 110 in his North American Fungi (as G. mammosus), and a
letter to Ellis from Morgan at the New York Botanical Garden states this to be the
same as his umbilicatus. The species is known from New Jersey (Ellis) and from Florida
(Lloyd). Morgan thinks that G. Smilhii Lloyd (Geastrae, p. 21) is the same, but
Cunningham does not agree with this. It should certainly occur on our North Carolina
coast.

The rays in this species while distinctly hygroscopic are more delicate and pliable
than in G. mammosus and much more so than in Astraeus. From G. Drummondii
(G. modulus) and G. asper the plant is easily separated by the delicately and softly
furfuraceous, not warted or asperulate spore sac, and by the smaller spores. From
G. Schmiddii, which is very near, it is separated by its hygroscopic and not revolute
rays and by the sessile spore sac. The spores of the two are very nearly alike. Lloyd
has seen the type of G. ambiguus and thinks it different from G. striahdus, to which he
refers the present plant (Myc. Notes, p. 311).

Geastrum minimum Chev. (Fl. Gen. Env. Paris 1: 360, pi. 10, fig. 3. 1826) does
not look much like this species. It has a distinct, slender stalk, though short, and that
is why Chevallier considered it new. He gives the sessile G. badium Persoon under his
species as a form alpha. Hollos considers G. badium and G. umbilicatus the same as
G. clegans Vitt. (which see under G. Schmiddii). From the description there is no
reason to think that G. striatus var. minimus Wallroth (Fl. Crypt. Germaniae 2: 400
(No. 2274)) is the same as our American plant above described. He refers to Chevallier's
G. minimum as the same.

Illustrations: Hollos. 1. c, pi. 9, figs. 15-17 fas C. ambiguus).

Lloyd. Myc. Notes, p. 71 (as G. striatulus). Also copied on pi. 98, figs. 3, 4.
Morgan. North Amer. Geasters, fig. 4.

Florida. Mrs. Sams, coll. (Lloyd Herb., asG. Smilhii).

New Jersey. New-field. Ellis (N. Am. Fungi, No. 110. Also several other collections by Ellis at the
N. Y. Bot. Card. Herb., as mammosus or umbilicatus) .

Geaster Drummondii Berk.
G. involutus Massee
G. Sckweinfurlhii Hennings

Plates 115 and 116

At the New York Botanical Garden Herbarium are two good plants from Bris-
bane, Australia (Herb. Massee) which appear to truly represent this species. The
plants are :

Hygroscopic, rigid; fleshy layer blackish; outer surface covered with sandy humus;
spore sac small, subglobose, sessile.

One plant has the inner peridium 12 mm. broad, spiny-granular; mouth blackish,
12-ridged. Spores faintly rough, spherical, 5.4-6.5^.

138 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

The other has the inner peridium 9.5 mm. broad, pulverulent-granular; mouth
creamy gray, 19-ridged. Spores faintly rough, 4.5-6.5/u.
Inner peridium of both creamy gray (pale leather color).

Type plants of G. involutus from St. Domingo at the New York Botanical Garden
show that no important difference can be made out to distinguish it from G. Drum-
tnondii. The appearance is identically like that of the plant with spiny peridium,
except that the mouth is not dark. The spores are the same, minutely warted, 4-5m
thick. The inner peridium is asperate and the mouth beautifully sulcate with 12 or 14
ridges. Hollos gives G. avellaneus Kalch. as a synonym of this species, and both as
synonyms of G. ambiguus Mont. The former was published only as a figure, with no
written description. The spore sac is shown as white and not dotted; the outer per-
idium evidently hygroscopic, in one case folded over the spore sac, its outer surface
ochraceous with some earth attached; mouth truly sulcate.

Illustrations: Berkeley. Journal of Bot. for 1845, pi. 1, fig. 4.

Cunningham. Proc. Linn. Soc. N. S. W. 51: pi. 4, figs. 17-18.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 27.

Hennings. Engler's Bot. Jahrb. 14: pi. 6, fig. 7.

Lloyd. Lycoperd. of Australia, etc., fig. 8 (types of G. Drummondii).

St. Domingo. Part of type of G. involutus at N. Y. Bot. Gard. Herb.
Australia. Victoria. (Kew Herb., as G. argentakts).

MYRIOSTOMA Desv.

Outer peridium as in Geaster, coriaceous, pliable, splitting in a stellate way with
segments expanded or reflexed; inner peridium with several short stalks (rarely one by
coalition), membranous, papery, opening by several to many mouths (rarely one).
Columellas several, slender. Capillitium threads free, simple or nearly so.

There is only one species of this genus known, as it has been shown by Patouillard
that Geaster columnatus Lev. is a synonym (see Lloyd, Myc. Notes, p. 156) .

Myriostoma coliformis Corda

Plate 117

Plants subterranean until dehiscence, lobes usually 5-7, acute; fleshy layer thin,
brown, adherent at first but wearing away; outer layer nearly smooth, usually with
adherent sand or trash. Inner peridium subspherical to compressed, 1.5—5 cm. broad,
roughened by minute points and lines, silvery brown. Mouths small, fibrous to
lacerate.

Spores (of plant from Lloyd Herbarium) spherical, strongly warted with irregular
pale warts, 4-6m thick, rarely abnormal ones up to 8/z. Capillitium threads long,
slender, tapering at both ends, entirely free from the peridium and columella, about
2-5/i thick, with a thick wall which often closes the lumen.

This species has a wide but erratic distribution in this country and in Europe.
It is not known from the Carolinas, but has been found in both the northern and
southern states.

Illustrations: Destree. 1. c, pi. 6, fig. A (as Geaster).
Hollos. 1. c, pi. 7, figs. 1-10; pi. 8, fig. 8; pi. 29, fig. 18.
Lloyd. The Geastrae, figs. 1-4.
Petri. Flora Ital. Cryptogama (Gasterales), fasc. 5, figs. 49 and 50.

<

—

a.

bo

LYCOPERDACEAE 139

Florida. Eldorado. Underwood, coll. (X. Y. B. G. Herb.).

Also collection by Mrs. Sams. (N. Y. B. G. Herb, from Lloyd).
Ohio. Sandusky. Schaffner, coll. (N. Y. B. G. Herb.).
Ontario. Point Abino. Cook and Underwood, coll. (N. Y. B. G. Herb.).
Reported from Dakota and Colorado.
Abo from South America in the Xew York Bot. Gard. Herb.

DISCISEDA Czern.

Fruit bodies growing just beneath the surface or partially exposed; outer peridium
thickish and volva-like, composed of two layers, an outer of loosely interwoven white
threads which grow around and enclose many particles of sand and trash and a much
thicker inner layer which is watery and subtranslucent when fresh and composed of
large, thin-walled, parenchyma cells which dry up and collapse at maturity to form a
thin coat on the inside of the outer peridium, the lower part of the outer peridium split-
ing awav at maturity and remaining in the ground, the upper part with the thin, firm,
inner peridium attached, breaking away and becoming free; inner peridium opening by a
small basal mouth to liberate the spores, which are spherical and warted. Gleba
composed of very minute, irregular, anastomosing chambers which are separated by
clusters of basidia and a very little trama; basidia much as in Lycoperdon, with four
slender sterigmata; capillitium threads extending inward from the inner peridium,
sparingly septate and slightly branched, at maturity breaking up into small sections.
Subgleba none.

Up to the present the internal structure of this genus has not been known, so that
its proper position in the system was obscure. It is now apparent, it seems to us, that
the genus is a highly specialized member of the Lycoperdaceae. The best evidence to
this effect are the basidia, which are obviously of the Lycoperdon and Bovista type and
quite different from the Astraeus and Calostoma type. Moreover, while the chambers
are very small and the hymenium poorly defined, there is an obvious resemblance to the
Lycoperdaceae in these characters also.

Three species are known from the southeastern states. Several others are known
from the West. Lloyd extends the genus to include several species without the peculiar
dehiscence and inverted mouth (Myc. Notes, p. 121, pis. 6 and 7; Myc. Notes, p. 1167).

A new genus, Abstoma, has been established by Cunningham (Lycoperdaceae,
etc., p. 206) to cover a single endemic species from New Zealand. It is very near
Disciseda, differing from it essentially only in the absence of a definite mouth, both
peridia wearing away gradually from the attrition of wind-blown sand, etc.

Key to the Species

Spores without a pedicel, or rarely with a short one

Spores 3.6-5,* thick D- Candida

Spores 5.5-9.5,1 thick D. subterranea

Spores with a pedicel up to 12.5m long or rarely longer D. pedicellata

Disciseda Candida (Schw.) Lloyd

Disciseda circumscissa (B. & C.) Hollos

Bovista circumscissa B. & C.

Calastoma circumscissum (B. & C.) Morgan

Plates 31, 78,79, 99 and 118

Plants single or cespitose, compressed globose, with a single rhizomorph-like root
when young and fresh; 0.5-2 cm. thick, 2-3.5 cm. wide before maturation; 0.8-1.5 x

140 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

1.5-2.5 cm. when dry; growing in the soil and partly exposed when fully grown; sur-
rounded until maturity by a thickish cortex (outer peridium) the upper and thicker part
of which remains attached to the inner peridium by a layer of spongy fibrous material,
the lower part tearing irregularly from the upper and separating more or less completely
from the inner peridium and remaining in the ground as broken and inconspicuous
fragments. The upper part with the inner peridium attached is now free and is easily
knocked out of the cup-like lower part by rain, etc., turning over so that the exposed
part of the inner peridium is now above. A small torn hole now appears at the place of
attachment of the stalk, in the center of this upturned base, and the spores begin to
escape. The cortex is a sand case held together by the woven white mycelium. The
inner peridium is rather firm and rigid, pale brown then silvery gray to slate gray in
age, minutely granular or scurfy on the exposed (lower) part, densely scurfy above
under the cortex. This thick spongy tissue above and thin granular layer below are the
remains of a watery translucent layer, 0.6-1 mm. thick which lay between the more
fibrous layers in youth. The inner peridium is very indistinct in sections when fresh,
becoming conspicuous on maturing and drying. The whole plant shrinks to scarcely
more than half its original size upon drying. Glebal chambers extremely minute,
5-25/1 wide, but often longer; no definite tramal plates are formed, as in Lycoperdon and
Scleroderma, but the basidia arise from irregularly anastomosing strings of threads and
the chambers are very irregular. Gleba white when fresh, changing as the plant
matures through yellowish olive to brown, at times faintly purplish. There is no
sterile base.

Spores (of No. 5967) brown, at times with a tint of purple, globose, 3.6-4.4/i thick,
warted, and with a short pedicel (mucro). Capillitium threads 3-5/i thick, irregular,
not rarely branched, extending inwards from the walls of the peridium; after maturity
breaking up into short pieces. Basidia (of No. 6092) short-pyriform, 6-7.4 x 9-14/1,
usually with 4 slender apical sterigmata of equal length.

This little plant is easily recognized by its peculiar habit of dehiscence and reversal.
It is not at all rare in pastures, the fresh, nearly mature plants appearing in mid-summer,
the liberated plants obvious in fall and winter. The earthy color makes the plant in-
conspicuous and it has seldom been reported.

The type of Bovista Candida Schw. in the Schweinitz Herbarium is this species, as
Lloyd says (Myc. Notes, p. 93), but it is so pasted on its sheet as to obscure its char-
acters, and the spores, while of the same size, are less rough than in our plants and in the
co-type of B. circiimscissa B. & C. The spores are 3.7-4.4/1, nearly smooth. The plant
may have been collected before fully mature. The capillitium threads are unmistak-
ably those of this genus, made up of irregular fragments, 3.3-4.6/1 thick, rather rarely
branched. We have some Chapel Hill specimens (as No. 6092) with spores less warted
and approaching those of the type of Bovista Candida. The co-type of Bovista cir-
ciimscissa in the Curtis Herbarium (Blake, No. 164) is like our No. 7115. The spores
are warted 3.9-4.6/1 thick.

The form we have described above is the Chapel Hill plant which is typical as
described by Hollos for the European plant, though running smaller than Morgan's
drawings. We have a collection from Asheville (Beardslee) which are 2.5 cm. broad,
counting the open cup. They are a brighter brown than our plants, about snuff-brown,
but the spores and other characters are the same.

Illustrations: Clements. Minnesota Mushrooms, fig. 88 (as Catastoma).
Hollos. 1. c, pi. 22, figs. 25-27; pi. 29, fig. 37.
Lloyd. Genera of Gastromycetes, fig. 33.

PLATE 7*

I t

t

fc

%

S

(

c

i>

c t

DlSCISEDA CANDIDA. No. 596V

LYCOPERDACEAE 141

Lloyd. Myc. Works, pi. 6, figs. 1-7.

Masscc. Journ. Bot. 26: pi. 282, fig. 6 (as Bovista). 1888.

Morgan. Journ. Cin. Soc. Nat. Hist. 14: pi. 5, figs. 4-9. 1892.

596". On mossy bank of branch, November 6, 1922.

6092. In pasture below Cobb Terrace, partly submerged, June 14, 1923. In fresh, nearly grown con-
dition; basidia in various stages of development.
7117. On ground in the drug garden, Sept. 10, 1923. Spores warted,4-4.8M thick.
7246. On ground in Arboretum, March 18, 1924. Very small specimen, only 5 mm. wide.
Also Nos. 5969, 5974, 5980, 6007, 7109, 7115, 7507, 8151.

Asheville. Beardslee, coll.

"Mountains." Buckley, coll. (Curtis Herb.).

South Carolina. Greenville. Sept. 1926. Coker, coll. (U. N. C. Herb.). Spores 3.8-4.2M.
Virginia. White Post. Dodge, coll. (N. Y. B. G. Herb.).

New Jersey. Newfield. Ellis, No. 3600. (N. Y. B. G. Herb.) Spores minutely warted, 3.7-5/i.
Ohio. Norwood. Lloyd, coll. (N. Y. B. G. Herb.). Spores 3.8-4.4m, sometimes with a short pedicel.
Nebraska. Atkinson, coll. (N. Y. B. G. Herb.). Spores 3.6-4.5/1.

Disciseda subterranea (Peck)

Plates 80 and 118

We have not seen this in the immature condition and have collected it only once.
As it appears in herbaria, it is usually of larger size than D. circumscissa, but individuals
may be smaller than the average in that species. The size of the plant is therefore not
a constant distinguishing feature, and the larger spores (5.5-9.5/z) must be finally relied
on for determination. The present species was described from Dakota and seems to be
common in the middle and far west (see Lloyd, Myc. Notes, p. 122; Morgan, 1. c, p.
143). We have a good specimen from Denver, Colorado (Bethel, coll.), and have also
seen a specimen from Canada kindly sent us by Dearness. These plants are about
1.8-2.8 cm. broad, subspherical or depressed; in the Canadian plant the outer peridium
covering about one-half of the inner; in the Denver plants forming only a disk opposite
the mouth.

Our collection from North Island, South Carolina, is apparently the first collection
from the southeastern states. As this station invades the territory of D. pedicellata,
which was plausibly separated from this species by the different regional distribution
and pedicellate spores, confidence in a real specific difference is distinctly reduced.
We can now make out no difference between the last two except the presence of a
conspicuous and persistent pedicel on the spores of D. pedicellata. The North Island
plants are smaller than the Denver and Canadian specimens and, like the former, are
not at all or scarcely depressed and retain only a small, sandy, basal pad representing
the outer peridium. Both the Denver and South Carolina specimens, as well as Rav-
enel's Xo. 15 (D. pedicellata) at Philadelphia, show several small openings in the inner
peridium in addition to the apical one. These may be due to insects but appear
natural.

As Petri gives the spores as only 4.5-5.3^, it is doubtful if he had this species.
Disciseda verrucosa Cun. (Lycoperdaceae, etc., p. 205) should be compared with this
species. From the description it is very close if not the same.

142 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustrations: Hollos. 1. c, pi. 22, figs. 20-24; pi. 29, fig. 36 (all as D. Debreceniensis).
Lloyd. Myc. Notes, p. 263, fig. 98; pi. 7, figs. 1-3 (as Colostoma).
PPetri. Flora Italica Cryptogama (Gasterales), fasc. 5, figs. 51, 52.

South Carolina. North Island. In sand dunes under cedars. Coker and party, colls. (U. N. C.

Herb.). Plants only 1.2-1.4 cm. thick. Spores strongly and irregularly warted, 7-9.5/j

thick.
Colorado. Denver. Bethel, coll. (U. N. C. Herb.). Spores spherical, strongly warted, 6-8.5m

thick, with traces of a halo.
Yuma. Clements, coll. Crypt. Form. Coloradensium. (N. Y. B. G. Herb., as Colostoma Candida.)
Spores minutely rough, 6-8m thick, often with a short pedicel.
Wyoming. Pitchfork. Davis, coll. (U. N. C. Herb.). Spores 5.5-7m, sometimes with a short pedicel.
Canada. Dorchester. Dearness, No. 2841. (Dearness Herb.) Spores strongly warted, 6-7.5/j

thick. Capillitium threads broken up into bits, 3.2-4.5m thick, wavy and irregular, with thick

walls.

Disciseda pedicellata (Morg.) Hollos
Catastoma pedicellatum Morgan

Plates 80 and 118

Plants somewhat flattened, about 2-3 cm. broad; outer peridium not thick, usually-
reduced to a small basal pad when collected; inner peridium purplish brown or pallid
grayish brown, soft and pHable. Spore mass ashy brown at full maturity.

Spores (of Hartsville plant) spherical, strongly warted, 6.5-8/u thick with a pedicel
about 2-9;u long [the short ones may have been broken]. Capillitium threads wavy,
fragmented, about 4m thick.

We have collected this plant at Hartsville, S. C, and have compared it carefully
with Ravenel's Bovista nigrescens, No. 15, as represented in the Herbarium of the
Academy of Natural Sciences of Philadelphia (ex Herb. Geo. Martin). Morgan es-
tablished the species on a South Carolina specimen of this series from Ravenel. The
Ravenel plant has spores irregularly warted, spherical, 7-9/x thick with pedicels up to
6/i long, in all respects like those mentioned above; capillitium threads fragmented,
sinuous, up to 5ju thick, with thick walls.

A plant received from Lloyd without locality data differs from South Carolina
plants in somewhat larger spores (7.4-9^) with more cylindrical warts and distinctly
longer pedicels (12-20/x). Otherwise the differences are inconspicuous and probably
represent a regional variety. For comparison with D. subterranea see that species.
For references see Lloyd, Myc. Notes, p. 121; also Letter 42 and Letter 62, Note 433;
Morgan, Journ. Cin. Soc. Nat. Hist. 14: 143.

Illustrations: Lloyd. Myc. Works, pi. 7, figs. 4-7.

South Carolina. Hartsville. In bare soil in a garden. Coker and party, colls. (U. N. C. Herb.).
Alabama. Auburn. Underwood, coll. (N. Y. B. G. Herb, and U. N. C. Herb.). Spores spherical,
strongly warted, 7-9m thick, with a pedicel up to 12. 5^ long.

PLATE 79

^ }

■ .

-. ; /

Discisf.da Candida. Xo. 6092. Fresh condition

PLATE 80

0 f>

Disciseda slbterraxea. Denver, Colo, [upper left]
Disciseda SUBTEKRANEA. No. 2841. Ontario [upper right]
Disciseda pedicellata. Hartsville, S. C. [center]
Diplocystis Wrichtii. Bahama Islands [Below]

1 \ i OPERDACEAE 143

Diplocystis Wrightii B. & C.

Plates 80 and 117

Fruit bodies small, crowded together in large numbers on a common, firmly leathery
stroma about 1-2 mm. thick which superficially covers the substratum. Individual
fruit bodies touching each other or here and there as much as 4 mm. apart; about 6-10
mm. thick, flattened above; at maturity the flattened top becoming pale and cracking
into pieces which fall away, leaving the firm, deep brown, leathery sides to form a cup
only slightly lower than the puffball within. Inner peridium paler brown, thin, pliable,
opening by a small apical pore, collapsing gradually as the spores escape and finally
settling into the bottom of the cup.

Spores yellowish brown under the microscope, spherical, minutely warted, 4-5. 5m,
sometimes with a small mucro visible. Capillitium very scarce, pale, threads long or
fragmented, branching, thick-walled with lumen almost closed, usually surrounded by
remnants of collapsed threads, about 4.8-1 1/u thick, with a rough wall.

The type is from Cuba and it is also known from the Bahamas (Inagua, Andros,
Long Island) and Guadaloupe.

As Lloyd mentions, the mouth is not protruding as shown in Fischer's figure. Our
specimens, while ample, do not show stages young enough to determine the internal
structure in youth, but the arrangement of the capillitium in a mount suggests that there
were chambers. We therefore place the genus provisionally in the Lycoperdaceae.

Illustrations: Fischer. Englerand Prantl, Pflanzenfamilien ll: 323, fig. 167 D.
Lloyd. Myc. Works, pi. 15.

Bahamas. On bare earth on 'hillside, Clarence Harbor, Long Island. Coker, coll. (U. N. C. Herb.).

Broomeia. The North American (Albany) record for this little known genus, as
given in Saccardo, is a mistake, as noted by Lloyd, the plant really being found in the
district of Albany at the Cape of Good Hope (Leveille; Ann. Sci. Nat., 3rd ser., 9:
129. 1848). This leaves the type species, B. congregala, known only from South
Africa. Of the other two described species, B. gnadalupensis, is almost certainly
Diplocystis Wrightii. For discussion and reference to figures of Broomeia see Lloyd,
Myc. Notes, p. 917. The genus Broomeia is distinguished from Diplocystis by a uni-
versal volva covering a crowded group of small peridia which are seated on a more or
less elevated, common base.

ARACHNIACEAE

Fruit bodies exposed; peridium single, very thin, crumbling at maturity. Gleba
with distinct chambers lined by the hymenium, these chambers forming at maturity a
mass of minute, separate, hollow peridioles which crumble and escape like grains of
sand. Capillitium and sterile base none.

This family is most nearly related to the Lycoperdaceae, from which it differs in
the simple peridium, absence of capillitium, and presence of peridioles. The peridioles
in Pisolithus are of a very different nature, being solid masses at all stages and without a
hymenium. We are including in the family only the genus Arachnion, as Arachniopsis
and Eolocotylon are not yet well enough known to be placed. If the former genus is
good and Long's description is accurate, it is very different from Arachnion, and if
Lloyd is right in thinking it the same as Arachnion (Myc. Notes, p. 1134) it need not be
considered. Eolocotylon may be related to Arachnion, and if so the family description
will have to be modified by the omission of peridioles.

ARACHNION Schw.

This genus is characterized by the peculiar nature of the contents of the peridium
at maturity. It is filled not with the typical gleba consisting of capillitium and spores
that are found in the Lycoperdons, but with a mass of granular particles like grains of
sand. They are grayish or ash-colored, and look like a mass of sandy earth. These
granules are peridioles or little sacks containing the spores. Each particle consists of a
woven web of hyphae enclosing a minute chamber around the walls of which are borne
the basidia pointing inwards, thus showing a close relationship to Lycoperdon. In
youth the structure is like that of Lycoperdon, the empty chambers being separated by
tramal plates of delicately woven hyphae. The peridium is smooth, not double, very
thin and fragile, and breaks easily into small particles at maturity, exposing the granular
contents. There is no sterile base. Originally there was only one species recognized,
A. album, described by Schweinitz from this state in 1822, but other tropical and austral
species have since been added. Long has described a genus Arachniopsis from Texas as
closely related to Arachnion (Mycologia 9: 272. 1917), but Lloyd, who saw the types,
thinks they are not different. Eolocotylon from Mexico and southern Texas is a
genus apparently related (Lloyd, Myc. Notes No. 21 : 254, pi. 73, figs. 5-8. 1906.
Also No. 22: 271. 1906).

Literature

Lloyd. The Genus Arachnion. Myc. Notes No. 21: 252. 1906; No. 46: 643. 1917. Also Myc.

Notes No. 66: 1133. 1922.
Lloyd. The Lycoperdaceae of Australia, etc., p. 39. 1905.

Long. Notes on New or Rare Species of Gasteromycetes. Mycologia 9: 271. 1917.
Saccardo. Syll. Fung. 7: 150. 1888.

For other literature see p. 194.

144

PLATE 81

6

Arachnion album. No. 730. Enlarged one-fourth

ARACIIXIACEAE 145

Arachnion album Schw.

Plates 81 and 118

Plants with the fruiting body above ground, irregularly spherical with a more or
less pointed base which extends into a small toughish root, lower surface more or less
grooved or pitted, surface smooth, dull white until maturity then buffy when dry, or
yellowish if water-soaked. Peridium thin, 0.5 mm. thick or less, the same thickness
all around, toughish. Gleba pure white then gray or brownish olive at maturity,
composed of a large number of minute empty chambers lined by a hymenial layer,
apparently made up only of basidia, the context between the chambers consisting of a
loose weft of delicate, interwoven hyphae. At maturity this loose tissue breaks down
and leaves the chambers as distinct granular particles which contain the spores and fall
apart like sand at maturity or decay into a sordid mass in wet weather. This structure
indicates a close affinity to Lycoperdon. The peridium has no definite opening but
falls to pieces slowly by decay or usually by the attacks of grubs (the larvae of the
fire-fly playing a large part). Odor none until ripe, then decided and nitrous, resem-
bling somewhat that of Bovistella radicata and some Lycoperdons. There is no capil-
litium or columella or sterile base.

Spores (of No. 730) brown under the microscope, smooth, short-elliptic, with a
decided mucro or short stalk about 1/i long, though sometimes twice the length of the
spore, 3.2-1.3 x 4.3-5.5/i- Basidia clavate, 3.7 x 11/*, apparently bearing only one
mature spore at a time though two or four sterigmata may be seen in the young state;
sterigmata very peculiar, a long, very slender part remaining attached to the basidium
and a shorter, stouter part falling off with the spores as its stalk.

It was from this state that Schweinitz first described this interesting plant. It is
plentiful in Chapel Hill in summer and seems to prefer cedars, but it also grows in
deciduous groves and in open places and we have found it in abundance in flower beds.
We have also found it at Hartsville, S. C. It is widely distributed in the eastern United
States and, according to Lloyd, in other parts of the world. It has been found only
once in Europe (Italy).

Illustrations: Lloyd. 1. c, pi. 16, pi. 73, figs. 1-3 and Myc. Notes No. 46: figs. 917, 918.
Schweinitz. Syn. Fung. Car. Sup., pi. 1, fig. 2.

730. In grass under hickory, September 9, 1913. Spores 3-4.8 x 3.7-5.5/1.

2364. Under cedars at The Rocks and on Rosemary Street, July 5, 1916.

3115. On damp soil in Strowd's lowgrounds, June 29, 1918.

5209. On sandy soil, June 22, 1922. Basidia 3.7 x llAi,'2-spored.

5224. On ground in Arboretum, June 23, 1922.

Salem. (Schw. Herb., No. 2221, type).
Hillsboro. Curtis, coll. (Curtis Herb.).

South Carolina. Ravenel. Fungi Car. Exs. No. 75. (Phil. Acad. Herb.).

Society Hill. Curtis, coll. (Curtis Herb.).

Hartsville. Coker, coll. In sandy soil in an apple orchard, June 1913.
Alabama. Earle, coll. (N. Y. B. G. Herb.).
Ohio. Berlin. Morgan, coll. (N. Y. B. G. Herb.).

SPHAEROBOLACEAE

Characters of the Genus Sphaerobolus

SPHAEROBOLUS Tode

Minute plants, growing on rotting wood (common form), on earth (Alabama)
and on dung of herbivorous animals (Mississippi, Africa). Fruit bodies subspherical
when on wood, breaking through without an exposed mycelium or less often seated on
the wood and surrounded and covered when young by the white mycelium. Peridium
of several layers, an outer, thick layer of woven hyphae, much like the mycelium; next a
pseudoparenchymatous, gelatinous layer (lacking in 5. iowensis), a thin filamentous
layer, and, finally a thick highly specialized inner layer of long, densely packed, pris-
matic cells radially arranged. This last is sometimes called the receptaculum. The
spore-bearing gleba is an inner ball of fertile tissue which at maturity is indistinctly
divided into areas by thin sterile plates. In youth these areas may be hollow and lined
with a distinct hymenium or stuffed and without a distinct hymenium, depending on the
species. However, even in species with stuffed chambers, the basidia are not indis-
criminately scattered, but are arranged in definite groups. This with the obvious
chambers of S. iowensis clearly indicates a relationship with the Lycoperdaceae, rather
than the Sclerodermataceae, as pointed out by Miss Walker. The dark, shining and
viscid glebal ball is shot out with force at full maturity by the sudden evagination
of the receptaculum. Spores smooth, sessile on the oblong basidia. Gemmae capable
of sprouting are also found in the gleba.

This is the only genus of the family. About eight species have been described but
the validity of several of them is doubtful (see Saccardo, Syll. 7, p. 46, and 17, p. 216.
For 5. impaticus Boud. see Hollos, p. 185).

Key to the Species

Gleba without chambers or well defined hymenium; gelatinous layer present in peridium

5. stellalus and var. giganteus
Gleba before maturity with chambers lined with hymenium; gelatinous layer lacking in peridium

S. iowensis

Literature

Corda. Icon. Fung. 5: 66, pi. 6, fig. 48. 1842.

Cunningham. Sphaerobolus stellalus Tode. N. Z. Journ. Sci. & Tech. 6: 16, figs. 1-6. 1923.

Fischer. In Engler and Prantl's Pflanzenfamilien l1: 346, fig. 182. 1900.

Fries. Systema Mycologicum 2: 309. 1823.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 43.

Gillet. Champ. Fr. (Gasteromycetes), pi. 2.

Hollos. Die Gasteromyceten Ungarns, p. 139, pi. 28, figs. 27-30.

Lloyd. Myc. Notes, p. 431, figs. 24S-247; also pi. 1 1 1.

Massee. Ann. Bot. 4: 60, pi. 4, figs. 55-55c (mostly from Fischer, Bot Zeit. 42: 1884). 1889.

Micheli. Nova Plantarum Genera, p. 221, pi. 101. 1729.

Nees von Esenbeck. Syst. Pilze Schw., pi. 11, fig. 122.

Pillay. Zur Entwicklungsgeschichte von Sphaerobolus stellalus Tode. Jahrb. der Phil. Fakultat

II der Universitat Bern 3: 197-219. 1923.
Pitra. Zur Kenntnis des Sphaerobolus stellatus. Bot. Zeit. 28: 681, 697, 713. 1870.

146

SPHAEROBOLACEAE 147

IVtri. Flora Italicacryptogama (Gasterales, fasc. 5: 135, fig. 83. 1909.

QttClet Champ. Juract Vosg., pt. 2, pi. 3, fig. 2. 1873.

Kabinowitsch. Bcitr. zur Entwickclungsgeschichte der FluchlkArper einigcr Gastromycetcn. Flora

79: 385-418. 1894.
Schroctor. In Cohn's Krypt-FL Schlesien 3, pt. 1: 688. 1889.
Sowerby. Engl. Fungi 1: pi. 22 (as LyCOperdon carpobolus).

Walker ami Andersen. Relation of Glycogen to Spore-ejection. Mycologia 17: 154, pi. 18. 1925.
Walker. Development and Mechanism of Discharge in Spliacrobolus idttiensis n. sp. and S. stdlatus

Tode. Journ. E. M. Sci. Soc. 42: 151, pis. 16-25. 1927.

Sphaerobolus stellatus Tode
5. carpobolus (L.) Schroeter
Carpobolus stdlatus (Mich.) Desm.

Plates 82 and 117

Plants arising from within the soft wood or bark, and pressing through to expose
its upper surface or the greater part; subspherical, about 1.5-2 mm. thick, at first dull
ochraceous from the surface color, then whitish from the cracking of this superficial
layer into flakes; soon after exposure the little sphere opens at the top in most cases
by four to seven stellate lobes and the outer, translucent-white layer (receptaculum)
of the gleba suddenly reverses itself outward and throws the little central flattish ball
containing the spores to a distance of several feet (Miss Walker reports a height throw of
over 14 feet). The reversed and evaginated receptaculum now appears as a watery
white sphere sitting on the lobes of the outer peridium. The ejected ball is slippery,
smooth and very dark chestnut brown, looking like a miniature horse chestnut seed.
Within it is divided into sections by delicate walls and in these areas are borne the
oblong basidia, 4-4.8/* thick, with four to six, rarely eight (up to nine in Pillay's form),
sessile spores. According to Miss Walker, these areas are never cavernous in their
development, and there is no definite hymenium. She gives the following cytological
facts for S. stellatus. All cells of the fruit body are binucleate; the two nuclei fuse in the
young basidium; the fusion nucleus divides to form 4-8; the nuclear walls disappear ;
the chromatin center of a nucleus enters each spore, and where there are less than 8
spores, some of these granules are left behind; the young spores are uninucleate, the
older ones may be binucleate.

Spores smooth, unevenly oblong, 3.7-4.8 x 7.4-10^. Mixed with the spores are
certain irregular densely filled cells (gemmae) formed by the hyphae which are also
capable of sprouting to form new threads. The entire ball of the gleba is said (Fischer)
to sprout without breaking up, sending out many hyphae from the gemmae and spores
within. For references to illustrations and literature see under the genus.

Misses Walker and Andersen give evidence that the discharge of the bolus in
Sphaerobolus is due to the rapid transformation of glycogen into sugars, followed by a
great increase in osmotic pressure in the cells of the peridium. Miss Walker has grown
four forms of Sphaerobolus in pure culture and obtained abundant fruiting. Her last
paper gives further details of structure and behavior and proposes a new variety of 5.
stellatus and a new species. We are including her descriptions of these below.

5112. On decaying oak log, May 16, 1922.
5204. On rotten deciduous wood, June 22, 1922.
7530. On decorticated pine board, Sept. 23, 1924.

Asheville. Beardslee.

"Carolina." (Schw. Herb.).
Pennsylvania. Bethlehem. iSchw. Herb.).

148 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Sphaerobolus stellatus var. giganteus Walker

"Basidiocarps having the color, structure, and appearance of 5. stellatus but being
much larger — up to 4 mm. in diameter (usually about 3 mm.) of unopened basidiocarps
and 5-6 mm. from tip to tip of peridium in opened basidiocarps. Spores globular with a
slight apiculus, 5-7 x 6-8ju, mostly 6 x 7/t."

Type locality: Starkville, Miss., on dung (horse?).

Sphaerobolus iowensis Walker

"Unopened basidiocarps 1-1.5 mm. in diameter. Peridium breaking stellately at
the apex into 3-8 parts (usually 4-5), the tips becoming only slightly recurved so that
the peridium of the opened fruit-body is somewhat cup-shaped. Interior of peridium
cadmium yellow when first opened, glebal mass raw umber when first exposed, soon
becoming almost black. Peridium separating into two regions, an outer and an inner.
Outer peridium during its development and at maturity composed of two layers, an
outer filamentous and an inner pseudoparenchymatic. Inner region of a layer of
tangential hyphae and a palisade layer. Gleba containing during its development many
definite chambers lined with basidia. Wall of glebal mass firm, composed of cells
6-15m in diameter. Contents of glebal mass soft and gluey, never drying hard. Spores
oval, 5-6 x 6-IOai (usually 5.6 x 8/u)."

It is to be noted that Cunningham (1. c, p. 16), speaking of Australian plants that
he refers to S. stellatus, described the gleba as having "several" cavities separated by
thin walls and says that these cavities are lined with hymenium. Whether by several
he means many is not indicated by the figures. However, as his plant has the gelati-
nous layer in the peridium, it does not agree fully with 5. iowensis.

Type locality: Hunters, Iowa, on old coniferous boards.

PLATE 82

J-

Sphaerobolus stellatus. No. 7530 [above]
Scleroderma ceaster. Stafford Co., Va.

From Doctor Howard A. Kelly. Reduced

TYLOSTOMATACEAE

Plants formed underground, emerging at maturity and consisting of a tough,
fibrous, stalk bearing the globular spore sac, the outer wall of which (in Tylostoma)
consists of a thickish, sandy layer which either breaks up and falls away soon in flakes
or persistently adheres and slowly wears away. The inner layer is a thin, tough, rather
firm, persistent membrane which contains the unchambered homogeneous gleba
without tramal plates; the spores escaping by a definite apical mouth (at times several)
or by an irregular or circumscissile dehiscence. Capillitium present, of varied char-
acter, arising from the entire surface of the inner peridium. Basidia club-shaped with 4,
irregularly scattered, nearly sessile spores.

The outer peridium in Queletia has not been described in detail. Lloyd (Myc.
Notes, p. 135) say's that it is "apparently a thin white coat that breaks up into granular
particles and mostly disappears." Six genera are included in the family by Miss White,
five of which are from America. Only two of these, Tylostoma and Queletia, are known
east of the Mississippi. Chlamydopus has been found in New Mexico and Washington,
Dictyocephalos in Colorado, Battarrea in several western states, and Sphaericeps is
known only from Angola in western Africa. See Miss White for descriptions of these.
For convenience we include in the key all the genera of the family. With some changes
it is copied from Miss White.

Key to the Genera

Peridium opening by an apical mouth

No free volva present; stem not flaring above Tylostoma (p. 149)

Free volva present; stem flaring above Chlamydopus

Peridium circumscissile

Peridium nearly plane below, dehiscing around the basal margin Battarrea

Peridium spherical, dehiscing at the equator Sphaericeps

Peridium opening irregularly

Peridium easily separating from the stem; capillitium not embedded in a tissue . . Queletia (p. 158)
Peridium firmly attached to the stem; capillitium embedded in a tissue Dictyocephalos

TYLOSTOMA Pers.

Mycelium subterranean, composed of delicate, white, much branched strands
which bear here and there white, sclerotium-like balls of various sizes, these developing
(apparently slowly) into the full sized buttons. Peridium composed of an outer sandy
case formed of delicate flocculence holding the earth,— in which in the button stage is
also included the unextended stem,— and an inner, very thin layer formed of delicate,
interwoven hyphae which, without apparent change, turn inward and continue as the
elements of the gleba. Stalk cylindrical, tough and firm, with a central column of
delicate soft, white, cottony tissue; surface usually brown and more or less marked with
scales and fibers. Capillitium abundant, arising from the entire inner surface of the
peridium. including the base, pale to nearly hyaline, frequently septate, much branched,
not rarely with cross bar connections as if anastomosing had occurred (but probably
only a method of branching) ; with walls irregular in thickness in the same thread, in
places closing the lumen; tapering tips lacking, the ends appearing abrupt as if broken.

149

150 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Basidia short-clavate, usually 4-spored, peculiar in having the spores scattered over the
surface. There is no trace of a hymenium or of tramal plates, the basidia being dis-
tributed equally on the interwoven threads of the homogeneous gleba. Spores spherical
or subspherical, rust colored to cinnamon, warted or smooth.

The genus is distinguished from others of the family by the smaller size and (except
in Chlamydopus) by the definite, circular, smooth or fimbriated apical mouth (or
mouths) by which the inner peridium opens to liberate the spores. Miss White reports
seventeen species from North America, eight of which are described as new. Tulostoma
is the spelling used by some authors, and Tulasnodea Fr. is a synonym.

In the unexpanded button the stem is a short plug enclosed in the outer peridium.
When the expanded plant dries the stem shrinks considerably in thickness and draws
away from the outer peridium which surrounds it above. This leaves a space between
the two, and the descending margin of the outer peridium which is often acute is the
so-called collar around the stem apex. If moistened after drying the stem swells back
to its original size and fills the collar again. The spores are often somewhat collapsed
when dry so that even those that were smooth when fresh may appear angular or
nodulated. In such cases they should be plumped out by an application of KOH or
lactic acid.

Literature

Bessey. Growth of Tulostoma mammosum. Amer. Nat. 21: 665. 1887.

Lloyd. Myc. Notes, p. 133, pis. 10 and 11 and text figs. 66,67. 1903.

Lloyd. The Tylostomeae, figs. 1-6 and pis. 20 and 74-85. 1906.

Morgan. Journ. Cin. Soc. Nat. Hist. 12: 163, pi. 16, figs. 1-5. 1890.

Petri. Sul valore diagnostico del capillizio nel genere "Tylostoma" Pers. Ann. Myc. 2: 412, pi. 6.

1904.
Petri. Flora Italica Cryptogama (Gasteralesl, Fasc. 5: 115. 1909.
Schroeter. Ueber die Entwickelung und die systematische Stellung von Tulostoma Pers. Cohn's

Beitrage sur Biologie der Pfianzen 2: 65, 1 text fig. 1877.
White. The Tylostomaceae of North America. Bull. Torr. Bot. Club 28: 421, pis. 31-40. 1901.

For other literature see p. 194.

Key to the Species

Mouth projecting as a circular glabrous tube with a sharp edge

Spores nearly smooth T. mammosum (p. 151)

Spores distinctly warted

\T. simulans (p. 151)
'\T.floi

Spore sac not warted \j. fioridonwn (p. 153)

Spore sac warted T. verrucosum (p. 153)

Mouth less projecting, sometimes not at all so, edge not composed of a fibrous roll; often elliptical or

linear; more than one mouth often present

Spores smooth, minute, 2-3/i T. Finkii (p. 154)

Spores smooth, larger, 4.2-5.5 x 5-7.4/1 T. vohulatum (p. 155)

Mouth appearing in the center of a slightly elevated fibrous mat, its edge eroded and fibrous unless

much worn
Spores smooth

Spore sac pale T. poculalum (p. 155)

Spore sac reddish brown, stem very long T. Lloydii (p. 156)

Spores minutely warted T. campestre (p. 156)

Spores distinctly warted T. Berkeleyii (p. 157)

TYI.OSTOMATACEAE 151

Tylostoma mammosum l'r.

T. peJunculdtiun (L.) Schroeter
T. brumalc Pers.

Plates 83 and 119

Spore sac 0.6-1 cm. thick, tan to buff or slightly darker; outer peridium thin,
wearing away gradually, leaving the inner peridium quite smooth or with persistent
particles except around the base where an adherent ring remains. Mouth cylindrical,
elevated, smooth, with a sharp edge, concolorous or often distinctly darker. Stem
1-2 cm. long, slender, brown, superficially spongy to slightly scaly, wearing away slowly
to expose the paler layer beneath; basal mat small.

Spores (of plant from North Dakota) spherical, very faintly warted, 3.8-4.5 x
4-5. 5m- Capillitium threads somewhat irregular, occasionally branched and septate,
up to Sju thick.

Lloyd thinks that the typical European form of this species does not occur in
America, but we find plants with dark mouths and exactly the microscopic characters
of the European plant. They differ from T. simulans and T. floridanum in the darker
mouth area and much smoother spores. We have examined a typical specimen of
T. mammosum from Europe with a dark mouth area (A. Vill, Fungi bavarici, No. 926)
and find that it agrees almost perfectly with the American plant: the spores are almost
smooth, 4-5. 5ji thick, and the capillitium threads agree. Another plant like it
(Schlesien; Dresler, coll., as T. brumale) at the New York Botanical Garden has similar
spores.

Illustrations: Batsch. Elench. Fung. 2: pi. 29, fig. 167 (as Lye. pedunculalum).
Gillet. Champ. Fr. (Gasteromycetes) pi. 7.
Greville. Scott. Crypt. Fl. 6: pi. 340 (as T. brumale). 1828.
Hollos. 1. c, pi. 11, figs. 18-20; pi. 12, fig. 1.
Lloyd. Myc. Works, pi. 78, figs. 5-8.
Morgan. Joum. Cin. Soc. Xat. Hist. 12: pi. 16, fig. 1.
Xees von Esenbeck. Syst. Pilze Schw., pi. 12, fig. 130.
Sorokine. Rev. Myc. 1890, pi. 98, fig. 346; pi. 103, fig. 366.
Sowerby. Engl. Fungi, pi. 406 (as Lye. pedunculatum).

North Dakota. Brenckle, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.). Mouth dark and spore

sac quite smooth and clean over most of its surface.
Wisconsin. Eagle Heights. (Univ. Wis. Herb, and U. N. C. Herb.) Spores spherical to oval,

minutely warted and ridged, 4-5.5 x 4.5-6^.
Kansas. Rooks Co. Bartholomew, coll. (N. Y. B. G. Herb.).

Tylostoma simulans Lloyd

Plates 84, 85 and 119

Spore case subglobose, 0.7-1.8 cm. thick, reddish brown when fresh, but so covered
with sand and earth as to obscure the color ; over this sandy coat is a delicate flocculent
coat of clean white wool which persists only in places as the plant emerges from the
ground and is soon shed. The sandy cortex is persistent for a long time and slowly
wears away above and even more slowly below to expose the brown or whitish inner
peridium which opens by a single, small, apical mouth with a smooth edge, at the top
of a little tubular elevation which in a few of our plants is surrounded by a slightly
darker area.

152 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Stem 1.5-3 cm. long, 3^1 mm. thick when fresh, 1.5-2.5 mm. thick when dry, the
base slightly enlarged by the soft, white, cottony mycelium which brings up a little
earth; surface often nearly covered with reddish brown lacerate scales which wear away
gradually on exposure and in age are represented only by the inherent parts; in some
plants of the same colony the reddish brown cortex instead of being lacerated may be
more even and adhesive and form a cracked and mottled outer layer of the stem which
may persist for a long time; color under the scales buffy brown, becoming dull dark
brown with age; body of the stem longitudinally striate when dry, deeply inserted in
the outer peridium and on drying shrinking away from the peridium and leaving the
usual collar which may or may not have a thin, free margin. Context of stem white,
fibrous, brittle when dry, containing a central cylinder that is partly filled with delicate,
silky fibers.

Spores (of No. 87) globose to subglobose, yellowish cinnamon in color, closely
set with thick, coarse, pale warts, 4.2-6 x 4.5-6.8/i thick counting the warts, which are
up to a fraction over lju long; in KOH solution appearing spinulose with a hyaline
material between the spines. Capillitium threads buff, moderately branched, septate,
somewhat swollen at the joints, 4.5-6.5/* thick.

This species is reported by Lloyd from Ohio and Texas. It may be considered the
southern form of T. mammosum, from which it is separated by the more persistent outer
peridium and by the strongly warted spores. There is also much less tendency to a
dark mouth area, though in our No. 87 there is a somewhat darker diffuse area around
the mouth. For a comparison with T. floridanum see that species. The present
species is peculiar in the paleness of the warts on the spores and their vagueness in
surface view. This seems to be due to the abundance of the nearly colorless amorphous
material which makes up the greater part of the warts and in places fills the space
between them.

Examination of T. mammosum in Ellis and Everhart's North American Fungi,
No. 2734, shows it to be exactly like the Chapel Hill plants. The spores are 4-6/u
thick, distinctly rough, subspherical; capillitium threads 3-6yu thick, thin- to thick-
walled, swollen at the septa. Their T. pedunculatum (Fungi Columbiani, No. 1888) is
not different. Spores 4-6/* thick, subspherical, distinctly rough. Capillitium threads
3-7/x thick, thin- to thick-walled, swollen at the septa. The T. mammosum of Curtis'
list is probably this species.

Illustrations: Lloyd. Myc. Works, pi. 79, figs. 2 and 3

87. On a dry, pebbly hillside among cedars, April 1903, and April 1904.
5930. Same location as No. 87, Dec. 9, 1922. Spores yellowish cinnamon, subspherical, distinctly

warted, 5.5-6.5/j thick.
7134. In grass under cedars, Sept. 22, 1923. Capillitium threads swollen at the joints. Spores

4.4-5. 1m thick, distinctly warted.
Also Nos. 5913, 7504, 7554.

Salem. Schweinitz, coll. (Curtis Herb., as T. bramale).
Asheville. Beardslee.

South Carolina. North Island. Coker and party, colls. (U. N. C. Herb.) .

Florida. Gainesville. Weber, coll. (U. N. C. Herb.). Spores rather pale, coarsely warted, 4.2-6ji

thick.
Alabama. Peters, coll. (Curtis Herb., as T. mammosum).

PLATE 83

n U

Tylostoma mammosum. North Dakota [upper left] Wisconsin [upper right]
Tylostoma floridanum. Type [center leftl

Tylostoma Berkeleyii. Folly Is., S. C, and Wilmington, N. C. [center right]
Tylostoma campestre. Colorado, Canada and Wisconsin [left to right below]

TYLOSTOMATACEAE 153

Iowa. Iowa tity. Macbride, coll. Ell. ft Ev., X. Amer. Fungi, No. 2734, as T. mammosum. (Path.
\ Myc Hub.)

Kansas. Stockton. Ell. & Ev., Fungi Columbiana, No. 18S8, as 7". pendunadaium. (,1'ath. & Myc.
Herb.)

Tylostoma floridanum Lloyd

Plates 83 and 119

We have examined specimens of the type collection of this species from the Lloyd
Herbarium. The plants are small and slender; spore sac about 5-7 mm. thick. We
find the spores to be spherical to oval or irregular, 4-6 x 4.8-6. 8n, distinctly warted
with elongated, irregular warts. Capillitium threads irregular, septate, branched,
up to 8.5/i thick with thickened walls.

Lloyd's description follows (The Tylostomeae, p. 18):

"Peridium dark castaneous color, with a small, tubular, circular, protruding mouth.
Cortex separating imperfectly, particles adhering to the peridium, thickened and sub-
persistent below. Stem slender, dark reddish brown, subsmooth, substriate with no
trace of scales, white within, hollow with central fibrils. Capillitium subhyaline, some-
times wavy [written waxy, in error?] with septa both swollen and even. Spores 5/i,
strongly asperate."

This plant is nearest T. simulans, from forms of which it can hardly be distin-
guished. The spores, while warted in about the same way as in T. simulans, show the
warts much more clearly in surface view. Other characters mentioned by Lloyd, as
slender, dark stem and colored peridium can be nearly or quite matched in simulans
or mammosum. Lloyd has described another species, T. pygmaeum, from Florida
(The Tylostomaceae, p. 16, pi. 78, figs. 3, 4) which, from the description, differs from
T. floridanum only in the unimportant character of color.

Illustrations: Lloyd. Myc. Works, pi. 80, figs. 1 and 2.

Florida. Lloyd. (Lloyd Herb., type, and U. N. C. Herb.)

Tylostoma verrucosum Morg.

Plate 119

This rare species has been found but a few times. Morgan described it from Ohio,
and Lloyd reports it from Texas (Long, coll.). We have seen authentic specimens
collected by Lloyd and determined by Morgan, and find the spores to be spherical to
irregular, distinctly warted, 4-6^ thick. Capillitium threads up to 8^ thick, irregular,
swollen at septa, walls irregularly thickened.

The original description by Morgan is as follows (Journ Cin. Soc. Nat. Hist. 12:
164):

"Peridium depressed-globose, thickish, becoming firm and rigid, with a dense
brown cortex of minute persistent scales and warts ; the mouth small, circular, prominent,
entire. Stipe long, slender, with a surface of brown lacerate scales, internally white,
with a central pith of long loose fibers; the mycelial bulb large, irregularly depressed-
globose. Threads of the capillitium long, slender, about as thick as the spores, hyaline,
branched; spores irregularly globose, minutely warted, pale brown, 5-6/u in diameter.

154 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

"Growing on the ground in rich soil in woods."

Lloyd says that the species is very close to T. squammosum of Europe, from which
it differs in verrucose cortex and more robust habit.

Illustrations: Lloyd. Myc. Works, pi. 76, figs. 3-5.

Morgan. Journ. Cin. Soc. Nat. Hist. 12: pi. 16, fig. 2.
Petri. Ann. Myc. 2: p. 424 (text fig.). 1904.

Ohio. Lloyd, coll. (N. Y. Bot. Gard. Herb.).

Tylostoma Finkii Lloyd

Plates 85 and 119

Spore case depressed-globose, 0.7-1 cm. high by 0.9-1.4 cm. wide, color when fresh
ochraceous to tan where not covered with sand, fading when dry to a mouse-gray with a
tint of salmon (under the sand) ; outer peridium consisting of a rough sandy cortex which
slowly wears away above and is even more persistent and somewhat thicker below where
it may appear with an obscure margin like a cup after the wearing away of most of the
upper part; mouth or mouths (1-4 in number) circular or elongated and irregular,
elevated (up to 0.8 mm. high) or scarcely so, concolorous with the peridium or slightly
lighter, the margin eroded or nearly smooth, and not surrounded by a fibrillose pad.
Stem 1.8-2.7 cm. long by 3-3.6 mm. thick when fresh, 1.5-3.2 mm. thick when dry and
then with longitudinal furrows, of equal diameter throughout or slightly tapering toward
the base which ends in a small myceloid swelling; surface minutely and evenly lacerate
scaly (the scales concolorous with the body of the stem and running horizontally around
it and often with remnants of the gray, myceloid, outer peridium attached; context
solid except for a small central strand of loose fibers; collar irregular, about 1 mm.
distant from the dry stem, in contact with it when fresh, concolorous with the top of
the stem.

Spores very minute, 2-3/i thick, smooth, spherical or slightly angled with a minute
mucro, spores cinnamon color (Ridgway) in mass, pale under the microscope. Capil-
litium threads lighter colored than the spores, irregular, wavy, sparingly septate, 3.8;u
thick, considerably swollen at the joints and at the ends which may be up to 12p thick _

In one of our plants there are four scarcely elevated mouths in a row and all elliptic
to elongated, in another there are two circular mouths and in two others there is a single
circular mouth.

As T. Finkii (Lloyd; Myc. Notes, p. 1169, pi. 225, fig. 2307) agrees with our plant
in its very characteristic spores, distinctly smaller than in any other species except in
T. Longii, described by Lloyd as a form of T. albicans, which is white and has large
rough spores, and as there are no essential disagreements we refer our plants to it. The
multiple and somewhat elongated mouths in some of our specimens point to group 6
of Lloyd's Tylostomeae as its proper location. Lloyd's specimens having only single
tubular mouths, he refers them to group 1. Tylostoma kansense and T. americanum
(Lloyd thinks these the same) and the two new species proposed by Miss White (T.
gracile and T. minutum) that if good might fall in the same group, are all easily sep-
arated by their much larger spores. In T. Lloydii Bres. (in Petri, Ann. Myc. 2: 423.
1904) the spores are smooth and small but distinctly larger than in the present species,
and the cortex is said to separate perfectly above, the mouth to be fibrillose until
weather-worn. We have looked through the species represented in the Schweinitz
and Curtis Herbaria, as well as in Washington and New York, and find nothing that
agrees with our plant.

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TYLOSTOMATACEAE 155

Illustrations: Lloyd. As cited above.

6060. By white oak tree in pasture, upland woods, Feb. 9, 1923.

Tylostoma volvulatum Borsch, in Sorokine. Slender form

Plate 119

Peridium up to 1 cm. thick, covered until age with a rather thin, brown, sandy coat
which wears away slowly and about equally over the surface, exposing the nearly white
inner peridium. Mouth plane, not regularly circular, margin clean and smooth, single
in all nine plants of our collection. There is no cortical cup at the base of the peridium.
Collar short, fimbriate. Stem 2-3 cm. long, 1.5-3 mm. thick, with a brown, more or
less scaly but not conspicuously lacerate surface, cespitose at times, in which cases the
peridia may be fused.

Spores subspherical, irregular, somewhat angled, practically smooth (even under
oil immersion), 4.2-5.5 x 5-7.4/*. Capillitium threads very irregular, nearly hyaline,
paler than the spores, up to 5n thick, with walls of varying thickness.

This agrees very well with T. volvulatum as treated by Lloyd (Tylostomeae, p.
19, pi. 81) except for more slender aspect. The mouth is almost exactly like that of our
Xo. 6060 (T. Finkii), as well as of the T. volvulatum-cespitosum-americanum group.

Illustrations. Lloyd. As cited above.

Sorokine. Revue Myc, 1890, pi. 98, figs. 347, 348; pi. 101, fig. 353a.

Porto Rico. Seaver and Chardon, colls. No. 142. (N. Y. B. G. Herb, and U. N. C. Herb.)

Tylostoma poculatum White

Plate 119

Miss White reports this species from Alabama (1. c, p. 431). We have examined
what appears to be a true example of the species from Calloway, Nebraska (Bates,
coll.; E. & E., Fungi Columbiani, No. 1889). The spores are quite smooth, irregularly
subglobose, 4— 5/x thick, with the contents precipitated or fragmented into irregular
particles which at first sight give somewhat the appearance of wrinkles on the surface.
We find no actual wrinkles as figured by Miss White. Tylostoma obesum Cke. & Ell.
is so close that we hardly see how the two can be separated. The spores of the Ellis
plant at N. Y. Botanical Garden, mentioned by both Miss White and Lloyd, have been
examined by us and found to be exactly like those of T. poculatum in size, shape, and
surface, the contents differing somewhat in being less coagulated (pi. 119, fig. 12).
Tylostoma granulosum of Europe is also very close. As represented by D. Saccardo
(Mycotheca italica, No. 424) the spores are smooth and just like those of the American
plants mentioned above, spherical to slightly irregular, 3.7— 1.4/j thick (pi. 119, fig. 13).
Lloyd gives the spores of T. granulosum as granular and Hollos gives them as warted.
Miss White's description of T. poculatum follows (Bull. Torr. Bot. Club 28: 431.
1901):

"Peridium globose, somewhat depressed, 1-1.5 cm. high, 1-2 cm. in diameter,
smooth, fawn-colored, membranaceous; outer peridium scaly, but more persistent than
in most species, remaining in the shape of a cup-like involucre round the base of the
peridium, mouth slightly raised, fimbriate, mostly large: collar entire, inconspicuous:

156 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

stem 1-3 cm. long, 3-6 mm. in diameter, cylindrical, firm, slightly bulbose, hollow or
stuffed, often with considerable remnants of the outer peridium attached: capillitium
lightish yellow, sparingly branched, septate, swollen at joints, 4-7/i wide, rather thick-
walled: spores ferruginous, subglobose, smooth, or irregularly ridged in the older
specimens, owing to the shrinking of the inner substance, short pediceled, 4-5/i in
diameter."

Illustrations: Lloyd. Myc. Works, pi. 83, figs. 1-2.
White. 1. c, pi. 34, figs. 4-6.

Tylostoma Lloydii Bres.

This species, known only from Cincinnati, Ohio, is near T. poculatum, from which
it seems to be distinguished only by its dark color and long stem. We have not seen
it and give below the original description by Bresadola (in Petri, Ann. Myc. 2: 423.
1904; see also Lloyd, The Tylostomeae, p. 22) :

"Outer peridium membranaceous, brown, soon falling away in pieces and leaving a
concolorous zone at the base; inner peridium subglobose, papery, smooth, pale, sub-
cinereous, flattened at the base and deeply umbilicate, about 1 cm. broad, 8-9 mm.
high, mouth white, oblong, 2 x 1§ mm., slightly protruding, like a subfimbriate, tomen-
tose circle, about 0.5 mm. broad, defined; stem hollow, fibrous-subwoody, brown,
covered all over with rather thickly set, at length deciduous scales, apex tapering and
sunken in the cup of the peridium and free from it on the sides, considerably thickened
below and furnished with a pale, subvolviform, fimbriate membrane at the base, 6-7.5
cm. long, 2.5-3 mm. thick at top, 4-5 mm. at base, white within; gleba subochraceous;
spores smooth, often inequilateral, 3.5-4 x 3.5/u; threads of the capillitium thick-walled,
not easily separating into joints, hyaline, 3-9/i thick."

Illustrations: Bresadola in Petri. Ann. Myc. 2: pi. 6, fig. 4, and one text nguie. 1904.
Lloyd. Myc. Works, pi. 82, figs. 5-8. ■

Tylostoma campestre Morg.

Plates 83 and 119

Spore case subglobose, about 1-1.8 cm. thick; outer peridium a sandy coat which
wears away except for a basal disk exposing about two-thirds or three-fourths of the
pale grayish to pale brownish, smooth or somewhat pitted, inner peridium. Mouth
area a distinct, spongy, somewhat elevated disk which opens in the center by a circular
or lacerate mouth to liberate the spores. Stalk 1.7-5.5 cm. long and 3^.5 mm. thick,
usually longitudinally sulcate when dry, and shrinking away from the outer peridium
so as to leave a distinct collar around its apex; surface layer dark brown, lacerate-scaly
over most of the area, white within and with a central cottony cylinder as is usual in
the genus; base expanding into a sandy bulb of moderate size.

Spores (of No. 861 Ip) pale, spherical or slightly oval, minutely warted, 4.5-6.8/*
or if elongated rarely up to 7.2^. Capillitium threads septate and slightly swollen at
the joints, irregular and constricted at places with the lumen closed by the thickening
of the walls, 3-6. 5/* thick.

This species is a northern and western plant, and we have not seen specimens of
it from east of the Mississippi. However, as it occurs on the shores of Lake Erie in
Canada we include it as eastern American. As Lloyd says, this is the American
representative of T. granulosum Lev. of Europe, of which T . fimbriatuni Fr. is a probable
synonym. We have carefully examined the spores of two typical examples of T.

PLATE 85

r1

*

V

S**

iW

Tylostoma simulans. No. 7504 [above]
Tylostoma Imnkii. No. 6060 [below]
Second plant from left moistened and swollen; others dry

TYLOSTOMATACEAE 157

granulosum represented in D. Saccardo's Mycotheca italica, No. 424, and find them
to be smaller and smoother than those of the American plant. In one plant (Path.
& Myc. Herb.) the spores are spherical to slightly irregular, smooth or nearly so,
3.7 4.4/u thick; in the specimen at the New York Botanical Garden the spores are
spherical to oval. 3.7-4.2p thick or if oval up to 5.5/i long.

The southern representative of T. campestre has been named T. Berkeleyii by Lloyd,
and while the differences are not great we are retaining the name (see that species for
comparison). Lloyd is right in considering T. punctatitm Pk. only a form of T. cam-
pesire. A part of the type collection at the New York Botanical Garden, noted below,
is similar and with identical spores, minutely roughened, 5-6.5/1 thick.

Illustrations: Morgan. Joum. Cin. Soc. Nat Hist. 12: pi. 16, fig. 4.

Petri. Ann. Myc. 2: p. 433 and p. 434 (text figs., the latter as T. punctatum). 1904.
Petri. Flora Italica Cryptogama (Gasterales), fasc. 5, fig. 76.

District of Columbia. Braendle, coll. (U. N. C. Herb, and N. Y. B. G. Herb.). Spores warted but

not reticulated, 5-6.8 x 5.5-7.2/1.
Wisconsin. (Univ. Wis. Herb, and U. N. C. Herb.) Spores 4.8-6 x 5.5-8.2M.
South Dakota. Chamberlain, coll. (Univ. Wis. Herb., as T. granulosum, and U. N. C. Herb.).
Nebraska. Long Pine. Bates, coll. Ell. & Ev., N. Amer. Fungi, No. 3514. (Path. & Myc. Herb.)
Also Bates, No. 499. (N. Y. Bot. Gard. Herb.) Spores spherical or elongated, minutely rough,

6-7 Aii thick.
Colorado. Harkness, coll. (N. Y. Bot. Gard. Herb., as T. fimbrialum) . Spores nearly spherical,

minutely warted with elongated warts, 5— 7.2/x thick. Capillitium threads septate, swollen

at the joints, usually up to about 6/x thick, up to 8m or 9ji at the joints: walls irregularly

thickened, closing the lumen in places.
Canada. Ontario. Deamess, coll. No. 2394 Pe. In sand by shore of Lake Erie, Oct. 5, 1922. No.

861. Port Frank on Lake Huron. Spores minutely but distinctly warted, 4.5-6.5^ thick.

No. 861 Ip. On shore of Lake Huron. No. 2394/861. London, Sept. 27, 1895.

Tylostoma Berkeleyii Lloyd

Plates 83 and 119

Spore case subglobose, 1—1.7 cm. thick; outer peridium as in T. campestre, wearing
away slowly but with traces nearly always remaining as small, inherent flecks on the
grayish to ochraceous tan inner peridium; the basal portion persistent as a closely
applied, sandy cup. Mouth as in T. campestre. Stem about 2.5-3 cm. long, rather
slender, with a deep brown spongy or flaky superficial layer, not so scaly as in T. cam-
pestre, apparently smooth in No. 8707 from Porto Rico, sulcate when dry, base with a
small bulb.

Spores (of plant from Folly Island, S. C.) spherical to oval, distinctly warted and
partially reticulated, 4.5-6/n thick or if oval up to 7.2^ long. Capillitium sparingly
septate, swollen at the joints, branched, irregular, 4-9/u thick, larger at the swollen
joints.

This is the southern representative of T. campestre. The spores of the present
species differ only in having slightly more prominent warts which are distinctly more
elongated so as to form a partial reticulum. The stem is less scaly (in plants seen);
there is more color in the denuded inner peridium, and the size averages smaller.

Illustrations: Lloyd. Myc. Works, pi. 84, figs. 8 and 9.

158 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

North Carolina. Chapel Hill. No. 5908. On bank of branch in a pasture, Nov. 20, 1922. Spores
irregularly spherical, 4.4-6^ thick, warted and often distinctly reticulated.

Wilmington. Dr. F. F. Wood, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.). Spores
spherical to oval or irregular, 5-6.5/1 or if elongated up to 7.4/i long, warted and often
partially reticulated.

Salem. Schweinitz, coll. (Schw. Herb., No. 2267, as T. squammosum) . Spores distinctly warted
and often with a partial reticulum, 5-6.5/1 or rarely 6.5 x 7.8/i.
South Carolina. Folly Island. Small and Bragg, colls. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.).

Hartsville. In sandy loam in a vegetable garden, Dec. 24, 1925. Coker, coll. (U. N. C. Herb.).
Spores distinctly warted and often with a partial reticulum, 4.5-6.5 x 5.4-7.4/1. These are
the largest plants we have seen, the case varying from about 1.2-1.7 cm. thick and with a
base with a very large sand bulb.

Also from South Carolina in Ravenel's Fungi Car. Exs. No. 80, as Titlasnodea fimbriata. (N.
Y. Bot. Gard. Herb., Phil. Acad., and U. N. C. Herb.) The spores of the plant in New
York are distinctly warted and with partial reticulum, 5-6.5 x 6-7.4m; capillitium threads fre-
quently septate, 3-7.4^ or up to 1 1m thick at the swollen joints.
Florida. Gainesville. Ravenel. Fungi Amer. Exs. No. 137 as T. mammosum. (Path, and Myc.
Herb, and U. N. C. Herb.) Mouth opening surrounded by a thick, felted, chestnut colored
mat; upper part of inner peridium quite clean. Spores strongly reticulated, 4-6.6m thick.
Alabama. Auburn. Earle, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.).

QUELETIA Fries

We give below a translation of the original description (Ofvers. K. Vetesnk.-Akad.
Forh. 28: 171. 1871):

"Peridium simple, membranous, smooth, fragile, furnished at the basal margin
with a membranous collar and discrete from the stem, at length dehiscing irregularly.
The apex of the stem extending into the peridium supports a rudimentary columella.
Capillitium flocculent, scarce, attached to the peridium on all sides, continuous, con-
torted, branched, hyaline. Spores very plentiful, in masses, short-pediceled, globose,
verrucose-echinulate. Stem strong, fleshy-fibrous, loosely broken up externally in a
peculiar way into fibrous shavings. Only one species known: Queletia mirabilis."

The genus differs from Tylostoma in larger size and absence of a definite mouth or
mouths, the inner peridium cracking and breaking up irregularly. There is no volva,
but the structure and behavior of the outer peridium are not well known. In speaking
of the capillitium as scarce, Fries seems to have been in error, as Miss White says it is
abundant (see below). For references to Lloyd's observations see under the species.

Queletia mirabilis Fr.

Plate 119

We take the following description from Miss White, 1. c, p. 441:

"Peridium globose, 2.5-3.5 cm. high, 3-4 cm. in diameter, fragile, easily separating
from the stem, rupturing irregularly, of a reddish brown color; collar irregular, of the
same substance as the peridium; stem 6.5-8 cm. long, 1.5 cm. wide at the top, 2 cm. at
the base, fascicular, reddish brown, within and without, like the peridium, solid, lacerate,
fibrillose, particularly at the base; capillitium very abundant and interwoven, forming
with the spores a felt-like mass, reddish brown, single threads whitish yellow, thick-
walled, hollow as shown by the truncated ends, septa rare or wanting, 5-9/x wide,
branches rather short, free ends rounded and recurved; spores subglobose, coarsely
warted, 4-6/x in diameter, some short-pediceled, inner portion breaking up and issuing
from the thin-warted coating which is then hyaline and shrivelled."

TYI.OSTOMATACEAE 159

A rare and interesting plant of curious distribution and habits. Lloyd in Myc
Notes, p. 1S5, refers to the four places that the plant had been found up to that time.
He later refers to another find in France (Myc. Notes, p. 337) and there gives structural
details of young plants. There is only one American record, on tanbark at Trexlertown,
Pennsylvania, where it was collected by Dr. Herbst, who found it three years in
succession.

We have examined the capillitium and spores of one of Dr. Herbst's plants from the
Pennsylvania station (now in the New York Botanical Garden) and do not find the
spores behaving in the way described by Miss White. They are strongly and ir-
regularly warted, 6-9/i thick, often with a short pedicel. The capillitium is peculiar in
fragmenting up at the joints into fairly long contorted sections with closed ends. They
may of course be broken at other places to allow the entrance of air or water, but the
completely closed sections are characteristic.

For other references by Lloyd, see Myc. Notes, pages 135, 217, 323.

Illustrations: Fries. Ofversigt Kongl. Vetens. Akad. Forhandlingar, Stockholm, pi. 4. 1871. Copied
by Fischer in Pflanzenfamilien l1: 343, fig. 180.
Lloyd. Genera Gasteromycetes, pi. 3, fig. 23.
Lloyd. Myc. Works, pi. 10, figs. 5-7; pi. 122.
Quelet. Champ. Jura, etc., part 2, pi. 3. fig. 8.
White. Bull. Torr.Bot. Club 28: pi. 38. 1901.

Pennsylvania. Trexlertown. On tanbark. Dr. Herbst, coll. (N. Y. B. G. Herb, and U. N. C.
Herb.).

SCLERODERMATACEAE

Plants more or less globular, usually partly or wholly exposed above ground at
maturity (but at times entirely embedded) ;* peridium thick or thin, tough or fragile
at maturity, rupturing irregularly and tardily, or splitting at maturity into stellate
lobes, or, in Pisolithus, flaking off in fragments from above downward; gleba divided
irregularly by thin or thick sterile tramal plates into areas which from the first are
stuffed full of the fertile tissue containing the irregularly arranged basidia. At maturity
both plates and fertile tissue break down into a powder at times by slow degrees. Plants
attached below by a thick mass of mycelial plates and fibers and sometimes stalked.
Basidia swollen, pyriform to subspherical; spores spherical, echinulate and sometimes
also reticulate, brown; capillitium none or rudimentary.

A closely related family is Podaxaceae, a long-stalked group of large plants not
represented in eastern United States. Podaxon and Phellorina occur in the western
states.

Key to the Genera

Walls of the gleba chambers very thin and not forming distinctly outlined or complete chambers, dis-
organizing throughout at maturity, together with the fertile tissue, to form a flocculent or
powdery mass containing the spores Scleroderma

Walls of the gleba chambers thick, forming distinct peridioles which slowly pulverize from above down-
ward after maturity and exposure Pisolithus

SCLERODERMA Pers.

Characters of the family and distinguished from Pisolithus by the absence of
distinct and persistent peridioles in the gleba. The plants are sessile except for a root-
like attachment of plates and fibers mixed with earth. The brownish or purplish or
olive-brown spores are echinulate or in several species obviously reticulated. In some
cases there is visible regularly or at times (before much exposure) a peculiar veil or halo
connecting the tips of the spines or reticulum. This effect is probably the result of a
hyaline material which covers the spore between the spines. The spores of this genus
show a remarkable peculiarity in their development. In 1889 it was shown by Beck
(1. c.) that in Phlyctospora fusca Corda (Scleroderma Juscum (Corda) E. Fischer) the
spores when somewhat more than half grown become surrounded by a sheath of cells
which grow in from surrounding hyphae or much less often from the basidia. This
sheath persists until the maturity of the spores and as the basidia have disappeared
when the sheath is formed, the remaining growth and maturation of the spores is made
possible by nourishment derived from the sheath. The sheath in most species is not
so prominent as in S. Juscum, but in essentials the development of the spores is the same
in all species investigated. We have studied S. lycoperdoides through these stages and
find that the basidia disappear when the spores are only half grown and quite colorless.
See details under that species.

* Zeller has recently described a peculiar Scleroderma from Oregon that is normally quite subterranean and that is still
more remarkable in having a deliquescent gleba. (Mycologia 14: 193. 1922.) Scleroderma pteridis is also subterranean
(see p. 162).

160

PLATE 86

Scleroderma geaster. . No. 657

SCLERODERMATACEAE 161

The plants are firm or even hard until mature, and while probably not harmful so
far as known are not of much value as food. Mcllvaine says (American Fungi, p. 615)
that all the Sclerodermas are edible when young and never poisonous. Marshall says
(Mushroom Book, p. 134) that S. aiiranlium (S. vitlgarc) has been eaten without harm,
but is pronounced very unattractive. According to Bulliard it is claimed that S.
vcrrucosum is deadly if eaten, but it is pronounced good by Mcllvaine (at least the plant
he discusses as such). The spiny spores are distinctly irritating to the eyes.

Literature

Bambeke. Bull. Soc. Roy. Belgique 43: 104, figs. 1-4. 1906.

Beck, t'ber die Sporenbildung der Gattung Plilyctospora Cda. Ber. Deutsch. Bot. Ges. 7: 212, one

figure. 1889.
Lloyd. The Genus Melanogaster. Myc. Notes, p. 1064. 1921.
Petri. Flora Italica Cryptogama (Gasterales), Fasc. 5: 93. 1909.
Rabinowitsch. Bcitrage zur Fntwickelungsgeschichte der Fruchtkbrper einiger Gastromyceten.

Flora 79: 385, pis. 10 and 11. 1894.
Sorokine. FieVeloppement du Scleroderma icrrucosum. Ann. Sci. Nat., 6th ser., 3: 30, pis. 5 and 6.

1876.
Tulasne, L. R. and Ch. De la Fructification des Scleroderma comparce a. celle des Lycoperdon et des

Bovisla. Ann. Sci. Nat., 2nd ser., 17: 5, pis. 1 and 2. 1842.

Key to the Species

Plant large (4-10 cm. thick), the peridium thick and scaly and splitting at maturity into several more

or less stellate and recurved lobes 5. geasler (p. 161)

Plant smaller (2-5.5 cm. thick), the peridium thinner (from very thin up to 1 mm. when dry) and less
stellate in dehiscence
Peridium straw-colored before opening, smooth except for coarse cracks; spores not reticulated

S. flavidum (p. 162)
Peridium yellowish, nearly smooth or finely areolated; spores strongly reticulated

5. bovisla (p. 163)

Peridium ochraceous or brownish, thick when fresh, thin when dry, the surface cracked into distinct,

inherent scales about 1-3 mm. wide, which are with or without a stout central wart;

spores strongly reticulated S. auraniium (p. 165)

Peridium leather or bay color, thick when fresh, thin when dry, the surface only delicately cracked

into areas and in large part smooth; spores not reticulated S. cepa (p. 167)

Peridium light brown or yellow-brown, dotted over with minute, dark brown, inherent scales; thin
when fresh and very thin when dry

Spores spiny, not reticulated S. lycoperdoides (p. 168)

Spores reticulated S. lycoperdoides var. reticulatum (p. 170)

Scleroderma geaster Fr.

Sderangium polyrhizon (Gmel.) Lev.
Stella amerkana Mass.

Plates 82, 86 and 120

Peridium about 4-13.5 cm. thick when closed and up to 15 cm. broad when open,
subglobose, depressed and often irregular and lobed; sessile and entirely embedded
until nearly mature, often remaining more than half buried; connected beneath with
diffused plates and strands of floccose mycelium; peridium hard and rigid, up to 5 mm.
thick when fresh, about 1-2 mm. thick when dry, nearly white, then yellow or dull
yellowish clay or straw color, rough and more or less cracked into areolations and scales;
splitting up irregularly at maturity into a varying number of lobes which curl back
as they dry and expose the dark spore mass which is deep brown when fully mature.

162 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Spores deep brown with a slight tint of purple, globose, asperulate, not rarely with
an imperfect reticulum, 5-10,u (most about 6.5-8yu) thick, including the spines which
are up to about 0.5— 1/x long.

This species gets its name from the way in which it cracks up, which is more or less
stellate like the dehiscence of the outer peridium in the genus Geaster. It is a common
plant in Chapel Hill, and is found from September through the winter in hard ground,
preferring clay. The plants open in the fall and persist a long time, becoming black-
ened and empty with age. They often appear on the clean, newly eroded pure clay
sides of gullies, a most remarkable choice for a home. Scleroderma pteridis Shear
(Bull. T. B. C. 29: 451. 1902) does not seem to differ essentially from S. geaster. The
type was collected 2-A feet below the surface of the ground and attached to dead rhi-
zomes of Pteris aquilina. The spores are given as 6-8/* and the peridium as 3-8 mm.
thick, and the author states that the species differs from S. geaster in the thicker perid-
ium and smaller spores. In our numerous collections of the latter the great majority
of the spores come within these limits, and the peridium thickness is almost exactly
3-8 mm. in moist, unopened specimens. A plant from the type collection kindly sent
us by Dr. Shear is unopened, as were all his plants, and most of the spores are im-
mature. We find, however, a few that are like those of S. geaster, about 7.7-10/i
thick, with surface spines. Dr. Shear writes (Nov. 27, 1923) that he is "now inclined
to think that this may be merely a form or variety of 5. geaster."

Illustrations: Boudier. Icon. Myc, pi. 186.
Hard. Mushrooms, pi. 65.

Lloyd. Myc. Notes No. 9: fig. 47. 1902; also pi. 30, figs. 1-3. 1905.
Massee. 1. c, pi. 2, figs. 35-35a.
Murrill. Mycologia2: pi. 17, fig. 9. 1910.
Vittadini. Monog. Lycoperd., pi. 2, fig. 1 1 ; pi. 3, fig. 14. 1842.

674. In woods, top of Lone Pine Hill, and along roadside in fields, Sept. 20, 1908.
973. In hillside pasture, Nov. 11, 1913. Spores 6-8.3/u thick.
1014. In a clay gully, Oct. 22, 1913. Spores 5-8.6/x, including spines, not rarely with an imperfect

reticulum.
2938. Unearthed in digging trenches (depth not known), Nov. 1, 1917. Spores asperulate, 6.7-9^

thick, omitting spines.
7571. Under cedars, Oct. 29, 1924.
Also Nos. 657, 660, 920, 978, 1029, 1761, 7106.

Asheville. Beardslee.

South Carolina. Hartsville. Coker, coll. (U. N. C. Herb., No. 6014). In Bermuda grass lawn,
Dec. 26, 1922. Spores warted and with a faint sign of a reticulum, 7.2-11^ thick.
Ravenel. Fungi Car. Exs. No. 76. (Phil. Acad. Herb.)

North Island. Coker and party, colls. (U. N. C. Herb.). Spores scurfy-warted, 7-10/j thick.
Florida. Gainesville. Weber, coll. (U. N. C. Herb., No. 7576). Stalked form. Spores very dark,
strongly warted and partly reticulated, 7.4—11.2^1.

Scleroderma flavidum E. & E.

Plates 87, 88 and 120

Plants gregarious, at times crowded, 2-4 cm. broad, subglobose, compressed,
sessile, plicate below around the thick basal attachment which holds the earth firmly;

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surface rather light straw-yellow, more or less cracked above and on the sides into small
to large, inherent , Bat areas like dried mud or in a more one-sided way to give a shingled
appearance (in some specimens the greater part not cracked); peridium tough, thin,
only 0.5 1 mm. thick (dry), splitting above into several persistent lobes that open up
irregularly (stellate, incurved or erect) to expose the spore mass, which varies remark-
ably in color in our collections, as fuscous then mummy brown (Ridgway) at full
maturity in No. 5S57. while in No. 675 it is Saccardo's umber.

Spores (of No, 675) spherical, 7.4-12.9/u, including the stout spines which reach a
length of 1.5/i and not rarely show faint traces of a halo around their tips. They are
mixed with numerous threads of the persistent trama which under the lens are nearly
hyaline, branched, and 2.5-4.5^ thick.

This is a good species and easily separated from 5. geaster, its nearest relative, by
the smaller size, much smoother and thinner peridium and the larger spores with longer
spines. Before opening the plants are partly or almost completely buried in the earth.
The spore color of No. 675 is nearly the same as our S. bovista (sense of Bresadola),
but the spores of the latter are entirely different, much larger and with a conspicuous
and perfect reticulum. Smooth forms of the present species approach S. cepa in ap-
pearance, but that can be separated from this by the thinner, more flexible, and gen-
erally darker peridium.

We have studied an authentic specimen of this species (E. & E., N. Amer. Fungi,
No. 169S, 2nd ser.) and find the spores to be spherical or nearly so, spiny-warted, 8-12/x
thick.

660. On ditch bank at side of road west of Carrboro, Nov. 3, 1912. Spores 7.8-14/1 counting spines,

which are rarely up to 3/i long, usually shorter.
3109. Pasture by New Hope Creek, June 23, 1918. Spores spherical, 7.8-14.8/1 thick counting spines,

which are rarely up to 3m long.
7562. In poor soil on campus, Oct. 14, 1924. Spores spiny, 8-1 lju.

Raleigh. L. R. Detjen, No. 1889. Near an old stump in a yard, Oct. 9, 1915. Spores S-ll/i,

excluding the spines which are up to 2y. long.
Asheville. Beardslee. Sept. 18, 1918. Spores 7.5-13p thick, counting spines, which are not rarely

up to iy, long. This plant agrees perfectly with our Chapel Hill plants.
Mecklenburg Co. On sandy bank, July 21, 1924. Couch, coll. (U. N. C. Herb., No. 7444).
Bladen Co. In sandy loam, Sept. 6, 1924. Alma Holland, coll. (U. N. C. Herb., No. 7509).
Spores very dark, strongly spiny-warted, 9.7-14.8^, with a halo.
South Carolina. Hartsville. Coker, No. 675. Sandy soil on Coker College campus, Nov. 1908.
No. 1962. In Bermuda grass lawn, Oct. 23, 1915. Spores 9.3-11^ thick, distinctly spiny.
No. 5857. Again on Coker College campus, Aug. 30, 1922. Spores 7-10.5^, excluding the
spines which are up to X.ly. long; hyaline threads mixed with the spores. No. 6022. On
Coker College campus, Dec. 26, 1922.
Florida. Mell, coll. (N. Y. B. G. Herb.).

Alabama. Reported as common. (Bull. Ala. Agric. Exp. Sta. 80: 268. 1897).
Mississippi. Biloxi. Tracy, coll. (N. Y. B. G. Herb.).
Wisconsin. Palmyra. (Univ. Wis. Herb, and U. N. C. Herb.)

Scleroderma bovista Fr. Sense of Hollos

Plates 89 and 120

Plants roughly spherical, usually flattened horizontally, at times lobed, about
1.5-4 cm. broad, narrowly or rather broadly attached to earth by a thick or slender root
which diffuses at once into mycelial plates and fibers; surface when young smooth, dull,

164 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

remaining smooth or less often becoming cracked up toward maturity over part of the
top into rather small, flattish areas about 1-3 mm. broad. Color buffy yellow to buffy
when young, then leather color or sordid brown, the top tending to become blackish
after maturity, and with black spots often present on paler plants. Peridium when
fresh only 0.7-1 mm. thick, pure white in section and not changing when cut or rubbed,
hardly more than 0.5 mm. thick when dry, at maturity splitting irregularly at the top;
the spore mass soon becoming very dark blackish brown with a tint of chocolate.
Tramal plates obviously yellow and until full maturity or after very distinct and perfect,
the threads of the plates peculiar in that they swell and regain their form in water and
can be distinctly seen to have clamp connections.

Spores (of No. 725) about blackish brown of Ridgway, spherical, in most cases
reticulated, but only imperfectly so in some, with a halo about the reticulum, 9-15.5m
thick, including the reticulum which is up to 3yu high.

There are in herbaria two plants passing for S. bovista, the one here described (our
No. 725, etc.) and the following (our No. 5920). The first is in almost full agreement
with Hollos's description and figures and we are considering it the species in his sense.
Both plants are found in Bresadola's herbarium under the name S. bovista. The one
here described is represented there by plants from Cincinnati, Ohio, sent by Lloyd, and
illustrated by him as S. Texense (cited below). Lloyd's plant is the exact form as ours
in every detail. The other form is represented by a European plant collected by
Bresadola which is exactly like our No. 5920. It is larger than No. 725, has a longer
and stouter rooting mass, a more yellow color, much thicker peridium, distinctly
paler and larger spores, and tramal plate cells which do not regain their form when
put in water. The English plant figured by Lloyd in Myc. Notes No. 8, fig. 43 is like
our No. 5920 in appearance, and he says he has the same thing from Falmouth, Mass.,
growing in clear beach sand, exactly the kind of habitat of our No. 5920.

Which of these two plants is the S. bovista of Fries we do not know, but think it
likely that the species is truly represented by our No. 725 and Lloyd's plants from
Cincinnati; and that our No. 5920 and the plant from Bresadola (and probably Lloyd's
plant from Massachusetts) represent another species that has not been correctly
understood. Petri's treatment of 5. bovista Fries apparently covers this plant.

Illustrations: Hollos. 1. c, pi. 23, figs. 16-20.

Lloyd. Lycoperdaceae of Australia, pi. 31, figs. 2-5. 1905.

Massee. 1. c, pi. 2, fig. 36 (the spore drawn here does not show the reticulum).

Petri. 1. c, fig. 54, No. 2.

600a. On cinder track with clay underneath, October 13, 1911.

725. On damp ground in Arboretum, September 8, 1913.
5880 and 5885. On damp soil on bank of New Hope Creek, October 8, 1922.

7113. On ground in Arboretum, September 10, 1923. Spores 9.7-14.8^ thick, usually reticulated.
8150. Grassy pasture on Glen Burnie farm, Oct. 11, 1926. Spores strongly reticulated, 9.5-13.5/:

thick.
Ohio. Cincinnati. Lloyd, coll. (Bresadola Herb.).

Scleroderma bovista Fries. Sense of Bresadola

Plates 90 and 120

Fruit body irregularly globose; depressed above, deeply and widely plicate beneath,
3-5 cm. thick; sordid yellow, the peridium rigid and brittle when dry and about 1 mm.
thick, very slightly to considerably squamulose, the very small inherent scales being

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formed by U irregular cracking of the outer surface of the pcridium; rooted by a dense
mass of entangled mycelioid cords, and opening when mature by irregular cracks and
lobes. Glebadark yellowish brown (sepia of Ridgw.); tramal plates yellow and quite
persistent.

Spores (of plant from Smith's Island) spherical, 11-18. 5m (including the reticulum),
the majority between 12.5m and 14m thick, with a coarse reticulum, surrounded by a
semi-transparent, pale yellow substance; treated with KOH solution the spore becomes
lighter, the reticulum clearer, and the gelatinous material swells considerably.

The distinguishing characters of the plant are its smooth to minutely areolated
yellow surface, thick peridium (even when dry) which is white in section, long thick
root, olive-brown spore mass mixed with yellow trama, and the large and coarsely re-
ticulated spores. For a comparison with S. bovista, sense of Hollos, see that species.
The species can readily be distinguished from 5. lycoperdoides by the larger size, more
yellow color, paler and far less conspicuous areolations, and by the strongly reticulated
spores. Scleroderma cepa, which is about the same size and color, differs (when dry)
in the thinner and less brittle peridium, more reddish spore mass and non-reticulated
spores; S.flavidum differs in the much paler straw color, coarsely cracked surface, and
non-reticulated spores; 5. aurantium has a much rougher surface, and when dry a
thinner and less brittle peridium.

We have compared our plant with a good example in Bresadola's herbarium col-
lected by him in Europe, and find them just alike in appearance and spores, which are
in the latter 9.3-14m thick, not counting the reticulum which is up to 3fi high.

North Carolina. Smith's Island. No. 5920. In very sandy soil near the sea, December 28, 1921.

(U. N. C. Herb.)
Canada. London. Dearness, No. 13SB. (U. N. C. Herb.)

Pelee. Dearness, No. 4826. (U. N. C. Herb.) Spores 12-25^, most about 19m, reticulated.

Scleroderma aurantium (Vaill.) Pers.
5. vulgar e Horn.

Plates 91 and 120

Fruit body about 2-5 cm. thick, subspherical and usually compressed horizontally,
plicate below and rarely lobed, sessile and attached by a thick mass of fibers; peridium
yellowish ochraceous or brownish, thick (about 2 mm.) when fresh and turning pink
when cut (when dry less than 1 mm. thick), the surface cracked into distinct areolations
which may or may not have a central wart, the warts and areolations often arranged in a
beautifully embossed pattern; after maturity slowly cracking into irregular lobes which
do not open in a stellate way. Gleba gray then nearly black, the tramal plates white.

Spores (of No. 5622) blackish brown, strongly reticulated, 8-1 1m thick, not counting
the reticulum w^hich is 1.5-2m high.

This is recognized by its verrucose and usually distinctly warted surface, thickish
peridium which turns pink when cut in the fresh state, white tramal plates and re-
ticulated spores. It is nothern in its range and in the south we have found it only in
the mountains. It grows in either deciduous or coniferous woods, but seems to prefer
mossy humus or very rotten wood under hemlock, spruce or pine. This is the species
that is occasionally parasitized by a Boletus (B. parasiticus) and we have found a fine
example of this condition at Blowing Rock (see Journ. E. M. Sci. Soc. 42: pi. 48.
1927).

166 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

A plant from Bresadola's herbarium determined by him as 5. vulgare is the same
as 5. aurantium, with spores 7-11.8/i thick.

Maire's cytological work on this species we quote below (Comp. Rend. 131 (2):
1247. 1900). It will be noted that he states the spore number to be four. Tulasne
says they are "usually four, sometimes two, three or five," and he also states that their
respective positions are very variable.

"The study of the basidium is especially interesting in Scleroderma vulgare. At the
beginning of the prophase of the first division, the cytoplasm contains a certain number
of granules which stain black in iron haematoxylin, around which radiate the micro-
somes. It seems clear that two of these granules become the centrosomes, for a little
later only two from among them show as center of radiation; they are then placed on
each side of the nucleus whose nucleolus and membrane soon disappear, while the
chromatic reticulum is transformed into two irregular knotty clubs which represent two
chromosomes and which extend almost from one centrosome to another.

"At the same time there is organized between the centrosomes a spindle; the two
chromosomes contract while increasing in thickness, then divide into two longitudinally,
while shaping themselves into a "V" until they are end to end parallel to the axis of the
spindle. They are then directed toward the poles and there reunite into a chromatic
mass which covers the centrosomes and from which seem still to radiate the astral rays.
The spindle soon disappears and the second division begins almost immediately : there
is formed around each nucleus a spindle with centrosomes and asters. All the granules
of the cytoplasm are massed on the radiations of the asters, except some which remain
in a parietal layer at the base of the basidium, in such a way that, except for this residue,
everything that is visible in the cytoplasm is oriented with regard to the four centro-
somes. The second division is like the first, since the events take place as if the four
centrosomes determined the formation of the four spores; they disappear moreover on
the arrival of the nuclei at the latter, to reappear only at the mitosis, which takes place
in the spore some time after its formation."

Maire also makes the general statement in regard to this species, as well as all
others studied [Geaster kygrometrkus, Lycoperdon caelatum, excipuliforme, gemmatum;
Nidularia globosa, Cyathus hirsutus] : "We have observed the fusion of only two nuclei
in the young basidia. The subhymenial cells always contain two associated nuclei
with conjugate mitoses. Many cells of the other tissues when they are old contain a
more or less large number of nuclei, but these nuclei come from an amitotic fragmenta-
tion of the two primitive nuclei. The number of chromosomes is two in all species
studied."

Illustrations: Bambeke. I. c, fig. 2.

Boudier. Icon. Myc, pi. 145. This shows the condition parasitized by Boletus parasiticus.

Bulliard. Champ. Fr., pi. 270.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 37.

Gillet. Champ. Fr. (GasteromycetesJ, pi. 15.

Hard. Mushrooms, pi. 64 and fig. 475.

Hollos. 1. c, pi. 23, figs. 8-15.

Hussey. Ills. Brit. Myc, pi. 17.

Marshall. Mushroom Book, p. 134.

Massee. Brit. Fungi, pi. 35, fig. 6.

PLATE 91

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S( l.ERODERMATACEAE 167

Michael. Fiihrerf. Pil/.freunde 1: \o. 82. 1918 (2nd ed.).
Petri. I.e.. figs, S3 , 54, No. 4 (both as 5. vulgarc) .
Tulasne. Ann. Sd, Nat., 2nd. ser. 17: pi. 1. 1842.

North Carolina. Blowing Rock. Coker and party, August 1922. No. 5506. Under balsam and
spruce on Grandfather Mountain. No. 5528. By a chestnut oak stump. No. 5622. On
rotting wood and deep woods mold by Glen Burnie Falls. Xo. 5656. At foot of a chestnut.
Spores S-13ji thick not counting the spines which are up to 1.8m long. No. 5681. In de-
ciduous woods by roadside. Spores 8-14m thick, with a distinct reticulum. No. 5686. In
mossy humus under hemlock. Parasitized by Boletus parasiticus.

Pennsylvania.' Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G.).

New York. Cold Spring Harbor, L. I. Couch, coll. (U. N. C. Herb.).

Illinois. Urbana. McDougall, coll. (U. N. C. Herb.).

Wisconsin. Devil's Lake. (Univ. Wis. Herb, and U. N. C. Herb.) Spores reticulated, 9.5-12^.

Canada. London. Dearness, Xo. 578F. (U. N. C. Herb.)
Avon. Dearness, No. 578 Av. (U. N. C. Herb.)

Scleroderma cepa (Vaill.) Pers.

Plates 92, 93 and 120

Plants subglobose or lobed, often flattened, about 1.5-6 cm. broad, sessile or nearly
so and attached by a thick mass of fibrous mycelium ; surface nearly pure white when
young, soon straw colored or yellowish ochraceous to dull leather brown, turning quickly
deep vinaceous when rubbed, finely areolated, or covered over most of the surface with
very small, inherent, concolorous scales, or in large part smooth; peridium up to 1.5
mm. thick when fresh, less than 0.5 mm. thick when dry, firm, not very brittle, much
thinner and less brittle than in S.flavidum, white in section at first then turning vinace-
ous on exposure; spore mass watery white until at least half grown, then rapidly becom-
ing nearly black with a purple tint and finally less dark, varying from Saccardo's umber
to sepia (Ridgway) depending on the time collected; odorless.

Spores (of No. 916) globose, strongly spinulose, 7-11.4/1 thick, including the sharp
spines which not rarely are up to 2/i long; not at all reticulate and with or without a
halo connecting the spine tips. Tramal plates only partly reswelling in water, heavily
encrusted with yellow crystals.

Smooth forms of this plant resemble S. bovista (in sense of Hollos) in surface ap-
pearance, but can be distinguished from the latter by the less rigid and less brittle
peridium, less dark spore-mass (if mature) and the smaller, spiny spores which are
without a reticulum. Young plants are separated by the paler color and purplish color
change of the peridium. Rough forms approach 5. flavidum in appearance, but that is
easily separated by the thicker, more brittle and paler peridium.

Illustrations: Bambeke. 1. c, fig. 4 (spores).

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 38.
Hollow. 1. c, pi. 23, figs. 3-7.
Lloyd. Myc. Works, pi. 31. 1905.
Petri. 1. c, fig. 54, Xo. 1 and fig. 55.

916. In grass on campus, Oct. 15, 1913.
7112. In sandy soil by campus path, Sept. 10, 1923. Fresh plants distinctly tinted lavender when

bruised.
7201. In swamp of Bowlin's Creek, Oct. 26, 1923. Spores 10-12.5*1 thick, counting the spines which

are long, sharp and triangular; no reticulum.

168 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

7515. Under a hawthorn bush in the Arboretum, Sept. 2, 1924. Spores dark, spiny-warted, 8-12ji
with a halo. In this collection the peridium had apparently grown more after the maturity
of the gleba, leaving the latter as a loose ball inside.

Also Nos. 5394, 5998, 6093, 7547, 7561.

Wisconsin. Burlington. (Univ. Wis. Herb., as 5. verrucosum, and U. N. C. Herb.) Spores warted,
7-10/t.

Scleroderma lycoperdoides Schw.

Plates 94 and 120

Gregarious or cespitose; body 0.8-5 cm. (most about 1.5-2.5 cm.) broad, usually-
broader than thick; surface light brown or yellowish brown and dotted all over with
minute darker brown or reddish brown, separated, inherent scales; becoming blackish
on long exposure; smooth or minutely roughened; flat below and abruptly rooted by a
stout embedded stalk of varying length which is soon dissipated into a few strong,
stout strands and plates or into many smaller fibers. Peridium thin, when fresh and
immature 0.5-0.7 mm. thick, becoming thinner and on the inside obscurely defined, the
inner layer becoming fibrous like the trama and continuous with it and wearing away
so as to leave in age only a very thin outer layer of a more leathery texture. Glebal
chambers very small, about 0.2-0.5 mm., the plates very thin and composed of delicate
rather straight hyphae about 2.5-3.5/u thick. Spore mass when very young watery
cream color, then deep brown and faintly purplish, changing to paler grayish brown
with or without a faint tint of olive after maturity; the delicate threads and plates of the
trama are not quickly disintegrated but remain as a decided matrix for the exposed
spores as they are slowly dissipated. There is no distinct dehiscence, but after much
delay there appears an irregular worn opening in the tip and there are often other pores
made by grubs. In age the stalk and all the basal part of the body are left as an empty
saucer.

Spores deep brown, spherical, asperulate, not reticulated, very variable in size of
body and length of spines in the same plant: in No. 5201 they are 7. 5-18. 5^ (most about
11-14/i) thick excluding the spines which vary from a length of 2\x (rarely) to scarcely
more than pointed warts (all from one fully mature plant). In No. 3488 a halo was
noticed connecting the spine tips over a part of the spore surface in many cases. Basidia
(of No. 5253) 7-8.5ju thick with 4-8 sessile spores.

While still half grown and before the gleba has a tint of brown the spores are formed
and the fertile tissue stuffing the chambers, including the basidia, has undergone a
disintegration into a translucent almost structureless mass surrounding and embedding
the spores, which at this time are hyaline, smooth and only about half the size they
attain later. It seems certain that the spores are nourished by the surrounding matrix
and undergo growth and maturation after separation from the basidia, as in 5. fuscum
(see under the genus).

This is our commonest Scleroderma in Chapel Hill. It prefers damp, shaded places
on bare ground or in thin grass or moss, but is often found on rotten logs in swamps.
It appears every year in the same places. The correct name has been in much doubt.
In American herbaria and literature it appears as S. verrucosum (Bull.) Pers., or S.
tenerum B. & C. (Cuban Fungi, No. 512). It is certainly near S. verrucosum, as that
species is now understood in Europe, and larger individuals pass for that species in
America. In fact one meets rarely with small European specimens which cannot be
distinguished from the typical American form. Such a plant is represented in Rathay's
Flora Exs. Austro-Hungarica, No. 1559 (U. S. National Herbarium). We have always
felt with Lloyd that the European name could hardly be applied to the present small
species. A European plant from Bresadola's herbarium (as S. verrucosum) has spores

PLATE 93

*SSk>

Scleroderma cepa. No. 7515 [above] ; No. 7547 [below]

SCLERODERMATACEAE 169

that can be matched by our plants, asperulate, 6.6-18/i, most about ll-14/i (pi. 120,
fig. 12), but the surface, while cracked into areas, is not dotted with small separated
scales of a different color. Lloyd uses S. tcnerum for our U. S. American plant, and the
Cuban plant certainly represents ours in the tropics. The types at Kew (kindly loaned
us by the director) consist of two good plants. One has the surface of our plants;
the other has more pronounced warts (pi. 95). The spores of both are the same and are
like the American form except for the average smaller size, 8-11/x, most about 8.5-10/x
(pi. 120, fig. S). They are almost exactly the size of the spores of S. cepa. We have
fortunately been able to find a good name for our form of the plant as proved by an
authentic specimen of 5. lycoperdoides in Michener's herbarium from the Schweinitz
Herbarium. The specimen is in good condition and shows all the characters of our
plants, the spores 10-15/i thick, covered with short or long, sharp spines which become
clear only after a treatment of K.OH has dissolved away the gelatinous, granular
material between them. Lloyd did not find the species represented in the Schweinitz
Herbarium in Philadelphia and so hesitated to use the name. There is, however, a
fragment of this in the Schweinitz Herbarium which has the same spores as the plant
above mentioned (10.5— 13ju)- There are also fragments from the Schweinitz Herbarium
in the Curtis Herbarium labelled S. lycoperdoides which seem to be the same. The
spore structure of S\ lycoperdoides is as in 5. cepa (No. 3109) except that the spores in the
present species are considerably larger and more variable. For a supposed variety of
this species see Peck, Rept. N. Y. St. Mus. 53: 848, pi. B, figs. 8-12. 1899.

Illustrations: Hard. Mushrooms, fig. 476 (as S. tenerum).

Lloyd. Myc. Notes No. 8: fig. 38. 1901 (as 5. verrucosum).

Trelease. Trans. Wis. Acad. Sci. Arts and Letters 7: pi. 9, fig. 7. 1888 (as S. verrucosum).
For the European S. verrucosum see: Bigeard. Flore Champ. 1: pi. 51, fig. 1.

Bulliard. Herb. Fr., pi. 24.

Gillet. Champ. Fr. (Gasteromycetes), pi. 15.

Hollos. I.e., pi. 23, figs. 21-29.

Hussey. Illustrations, pi. 17, fig. 1.

Leuba. Champ. Comest., etc., pi. 52, figs. 7-11. 1887-90.

523, 726, 2396, 5253. All from damp soil in the Arboretum, June to October.

5395. On rotten logs in a swamp, July 24, 1922.

7000. On earth by Meeting of Waters branch, July 30, 1923. Some of these plants were mere shells,

the spores having all been washed out.
Also Nos. 1013, 1785a, 5201.

Asheville. Beardslee.

Blowing Rock. Coker and party, August 1922. (U. N. C. Herb.)

Also collection by Coker, Sept. 3, 1924. Spores spiny-warted, 10-14.8m thick.
Winston-Salem. Vogler. In moss and on ground, July 30, 1919. (U. N. C. Herb., No. 3488).

This is also probably the type locality.

Florida. Marion Co. Couch, coll. (U. N. C. Herb., Nos. 7278 and 7279). Spores (of No. 7278)
irregularly spiny and granular with hyaline material, 8-12*i thick.

Gainesville. Walker, coll. (U. N. C. Herb.).
Alabama. Peters, coll. (Curtis Herb., as 5. vulgare).
Virginia. Falls Church. Murrill, coll. (N. Y. B. G. Herb.).

Warrenton. Coker, coll. (U. N. C. Herb.).
Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.).
New York. Represented from various places in N. Y. B. G. Herb.

170 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Connecticut. Redding. Underwood, coll. (N. Y. B. G. Herb.).

Massachusetts. Sprague, No. 737. (Curtis Herb., as Lycopcrdon gemmatum.)

Wisconsin. Nelson's Woods near Madison. (Univ. Wis. Herb, and U. N. C. Herb.) Spores spiny-

warted, 8-11*1.
Canada. Ontario. Dearness, coll. (U. N. C. Herb., several collections). Spores (of No. 3841b)

spiny, 7.5-11/i thick.

Scleroderma lycoperdoides var. reticulatum n. var.

Size, habit and surface appearance about as in the species, from which it differs
in distinctly olive colored spore mass at maturity and in the strongly reticulated instead
of merely spiny spores and the more delicate, less massive roots. The color may run a
little darker and the spots somewhat less conspicuous than is usual in the species, but
these characters are too variable to be emphasized. The gleba color and spore differ-
ence are very obvious, the latter being easily seen under moderate power. When the
dark olivaceous spores are shaken out, the remaining matrix is bright olivaceous yellow.

Spores (of No. 8076) spherical, spiny but also strongly reticulated, 10-13ju.

North Carolina. In mixed woods by Pisgah Creek, Haywood Co., N. C, Aug. 7, 1926. Totten, coll.

OJ.N.C. Herb., No. 8076).
New York. Under shrubs in New York Botanical Garden, July 27, 1919. Coker, coll. (U. N. C.

Herb.). Spores strongly reticulated as in No. 8076.

PISOLITHUS Alb. & Schw.

Distinguished from Scleroderma by the distinct peridioles or irregular bodies that
with their separating walls make up the body of the gleba and contain the spores. A
number of species have been described, but all those of the northern temperate zone are
probably only forms of the one that occurs here. There is certainly nothing in the
descriptions to separate them. Differences in form, as presence or absence of a con-
spicuous, simple or complicated stalk, and surface color of peridium are all certainly
worthless. The plant is a common one in Chapel Hill and all forms occur. The fresh
young plants if unweathered are very obviously yellow, but this soon changes to shades
of brown or blackish on exposure.

This has been widely treated as Polysaccum, a genus now known to be antedated
by Pisolithus.

Literature

Bambeke. Bull. Soc. Roy. Bot. Belgique 42: 178. 1906.
Massee. Revision of Polysaccum. Grevillea 16: 26, 76. 1887-88.
Petri. Flora Italica Cryptogama (Gasterales), Fasc. 5: 104. 1909.

Tulasne, L. R. and Ch. Sur les genres Polysaccum et Ceaster. Ann. Sci. Nat., 2nd ser., 18: 129
pi. 5, figs. 1-7. 1842.

Pisolithus tinctorius (Pers.)
P. arenarius A. & S.
Polysaccum pisocarpium Fr.
Polysaccum crassipes DC.

Plates 96, 97 and 120

Mycelium olivaceous yellow. Fruiting body irregularly globose or pear-shaped
and up to 18 cm. high and 10.5 cm. broad, narrowing below to a stout, stem-like, ir-

PLATE 94

•■

Scleroderma lycoperdoides. No. 523

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SCLEROPERMATACEAE 171

regular base of very variable length and thickness, at times sessile and attached as in
Scleroderma; solid and firm, at first olivaceous yellow but usually dull ochraceous when
found, then brownish to blackish, the surface nearly glabrous; peridium very thin and
at maturity becoming brittle and soon crackinginto flakes above until the whole contents
are exposed. Peridioles in section whitish or yellow, then watery vinaceous, then
through darker vinaceous to nearly black, subspherical, irregularly angular, usually
compressed, about 1-2 mm. thick and up to 4 mm. long, smaller toward the base;
texture that of a firm elastic and translucent jelly until the desiccation of maturity.
Peridioles separated by a nearly black jelly enclosing a densely woven mass of delicate
threads with clamp connections.

Spore powder cinnamon brown; spores globose, spiny but with the spines covered
with a gelatinous material that makes them look blunt when first put into water,
7— 11.5m thick, counting the spines which are up to ly. long.

This peculiar plant is common in fall in well drained gravelly and sandy soil, as
pastures, old fields, thin places in lawns, etc. The dry spore powder is scattered by the
breaking down of the peridioles after the peridium cracks off. This disorganization
proceeds by degrees and the lower peridioles are intact for some time while the upper
part is being dissipated. The old, woody, discolored base remains intact for months.

The blackish jelly between the peridioles and the peridium also of this plant gets
its color from a bright olivaceous yellow pigment which quickly stains cloth or paper
and is hard to remove. It is said to be used for dye for cloth in France (Lloyd, Lyco-
perdaceae of Aus., p. 12). When put under the microscope this jelly itself is seen to
contain most of the pigment, the threads being pale. The vinaceous purple color of the
jelly in the peridioles is due to the maturing spores, the threads and jelly around them
being colorless. If a ripening peridiole be put into water the purplish pigment in the
spores will diffuse into the water and color it. The plant thus contains two distinct
pigments, a yellow and a purple one. The purple tint of the spores is lost at full
maturity, the dry spore powder being deep brown without purple. The very surface
of the peridioles is covered with a thin weft of yellow fibers that are not gelatinized and
contrast strongly with the jell}' on each side. When fresh the very thin weft of fibers
that make up the peridium is lined beneath by about one millimeter of the same black
jelly that surrounds the peridioles. This is mottled with yellow plates as if the outer
peridioles had been aborted.

The spores of P. crassipes as illustrated by Tulasne show thick, rounded warts
rather than spines, but we do not take this to be evidence that the plants are different.
When first put into water many of the spores have thick, rounded warts but these
become sharper after standing awhile or on appb'cation of KOH.

As in the case of Scleroderma and the Nidulariaceae, it is obvious that the spores
grow considerably after the basidia disappear by gelatinization. At the stage shown in
fig. 17 the basidia have lost all their contents and are almost invisible, yet the spores
are only about two-thirds grown. Figure 18 shows the mature spores at the same
magnification.

As Persoon's Synopsis Fungorum is now generally accepted as the starting point
for the nomenclature of the Gasteromycetes, there is no reason why his specific name
(based on Micheli's plate) should not be applied to this plant. Though Lloyd suggests
this, we cannot find that the combination we use has ever been published before. For
other synonyms, see Hollos, p. 179.

172 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Illustrations: Albertini and Schweinitz. Consp. Fung., pi. 1, fig. 3.
Boudier. Icon. Myc, pi. 185.

Bruns. Beitrag zur Kenntniss der Gattung Polysaccum. Flora 78: 67. 1894.
Hollos. 1. c, pi. 27 and pi. 28, figs. 1-6.
Lloyd. Lycoperdaceae of Australia, pi. 29. 1905.
Michael. Fiihrer f. Pilzfreunde 3: No. 345. 1919 (2nd ed.).
Micheli. Nova Plant. Gen., pi. 98, fig. 1.
Nees von Esenbeck. Syst. Pilze Schw., pi. 13, fig. 131 B.
Sowerby. Engl. Fungi, pi. 425 (as Lycoperdon capsuliferum) .
Vittadini. Monog. Lycoperd., pi. 2, fig. 8 and pi. 3, fig. 13.

663. By path northeast of athletic field, November 26, 1912.
974. Among scattered pines on hillside pasture, Nov. 11, 1913.
4903. On ground in a yard, October 17, 1921. Spores spherical, densely but rather bluntly spiny,

7-9.5/1 thick, rarely ll/i.
7090. In clay soil by road west of President's house, August 23, 1923. Plant 18 cm. high and 10.5

cm. thick when gathered.
Also Nos. 686, 687, 723, 902, 2219, 5402, 7125, 8170.

South Carolina. Marion Co. Sandy soil near Little Pee Dee River. Coker, coll. (U. N. C. Herb.

No. 8136) . This seems to be an immature or conditional form. The spores are bluntly warted

instead of spiny.
Florida. Gainesville. In sandy pine woods. Weber, coll. (U. N. C. Herb.). Spores spiny- warted,

8.5-11.5^.
Alabama. Tuskegee. Beaumont, coll. (Curtis Herb.).

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PLATE 97

PlSOLITHUS tinxtorius. No. 5402

NIDULARIACEAE

Plants small, growing on the ground, on dead vegetable matter or on dung; cup-
shaped or goblet-shaped or globular; the mouth in two genera closed until maturity
by a membrane (the epiphram); gleba in the form of separate, lens-shaped bodies
(the periodioles) of easily visible size, which contain a number of large, smooth spores
and are covered by a dark, hard, protective coat which is with or without an obvious
superficial, white, thinner layer (the tunic). Basidia clavate, with 2-8 smooth, sessile
or stipitate, apical or scattered spores, forming in Crucibulum a definite hymenial
layer near the center, but in Cyathus occupying irregularly, together with sterile threads,
a large central area of the peridiole. Spores smooth, usually flattened, binucleate in
the species studied. Capillitium lacking.

The peridioles do not open of their own accord, but liberate the spores only when
eaten by insects (or when decayed?). The tunic is thick and removable in Crucibulum,
very thin and obscure in Cyathus (Lloyd thinks it entirely absent in C. stercoreus).
The hard wall of the peridioles is only superficially black or brown, and composed almost
entirely of very irregular, branching and knobbed, very thick-walled sclerotic cells,
which, when a peridium is crushed, are found scattered abundantly among the spores.
In nature they retain their position and are not mixed with the spores. These cells
are only remotely, if at all, homologous to the capillitium of other genera. Next within
this hard layer comes the spore-bearing tissue which may or may not form a definite
hymenial layer. See under the different genera for details on this point. The spores
have been made to germinate in water and in culture media and have produced fruit
bodies in pure culture (Walker, Hesse, Eidam) . On account of the peculiar resemblance
to little nests with eggs, these plants are popularly known as "bird's-nest fungi." So
far three of the four genera have been found in the eastern states. In two of these the
peridioles are connected on their under side to the wall of the peridium by a cord (the
funiculus), which is elastic when wet and brittle when dry. In Crucibulum it is usually
very obscure after maturity and often disappears from gelatinization. In Cyathus,
the funiculus, while much more conspicuous, may be rarely entirely lacking from some
of the upper peridioles (Miss White, Lloyd). In the other they are free from the wall,
but embedded in a mucus. The spores are hyaline, and are elliptic or subglobose.
They vary a great deal in size from the comparatively small ones of Crucibulum vitlgare
to the immense ones of Cyathus melanospermus. They also vary considerably in the
same collection and at times in the same peridiole, and they are much more abundant
in the small-spored species than in those having large spores. In 1902 Miss White
(1. c.) brought together in convenient form the species known from North America and
established a new genus, NMula, with two species from the western states and Canada.
Lloyd (1. c.) adds a third species (the Cyathus emodensis of Berkeley, Kew Journ.
Bot. , p. 204. 1854) and extends the geographical range of the genus. For the structure
of the peridioles, see Tulasne, Walker, Lloyd, or Fischer, as cited below. Fischer sug-
gests that the family may have descended from the Hymenogastraceae, mentioning
Octaviania, but the origin of this group is certainly very obscure. Lohwag (cited on

173

174 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

p. 194) expresses the surprising opinion that these plants are related to Tylostoma
and represent a compound group of Tylostoma fruit bodies enclosed in a common sheath.

Literature

Andrews. Notes on a Species of Cyalhus common in Lawns at Middleburg, Vermont. Rhodora 2:

99, pi. 17. 1900.
Brefeld. Botanische Untersuchungen iiber Schimmelpilze 3: 176. 1877.
Cunningham. A Revision of the New Zealand Nidulariales, or "Birds-nest Fungi." Trans. New

Zealand Institute 55: 59, pis. 3 and 4. 1924.
Eidam. Die Keimung der Sporen und die Eutstehung der Fruchtkorper bei den Nidularieen. Cohn's

Beitrage der Pflanzen 2: 221. 1877.
Fischer. 1. c, in Pflanzenfamilien, p. 324, which see for other literature.
Fries, R. E. Om utvecklingen af fruktkroppen och peridiolerna hos Nidularia. Svensk bot. Tidskr.

4: 126, one plate. 1910.
Fries, R. E. Ueber die cytologischen Verhaltnisse bei der sporenbildung von Nidularia. Zeitschr.

Bot. 3: 145, pis. 1 and 2. 1911.
Hesse. Keimung der Sporen von Cyathus strialus Willd., einer Gastromycetenspecies. Jahrb. Wiss.

Bot. 10: 199. 1876.
Lloyd. The Nidulariaceae, figs. 1-20 and pis. 102-111. Cincinnati, 1906
Maire. Recherches cytologiques et taxonomiques sur les basidiomycetes. Bull. Soc. Myc. France 18:

178 (of supplement), pi. 6. 1902.
Martin. Basidia and Spores of the Nidulariaceae. Mycologia 19: 239, pis. 22, 23. 1927.
Mollaird. Le cycle de deveioppement du Crucibuhim mdgare Tul. et de quelques Champignons supen-

eurs obtenu en cultures pures. Bull. Soc. Bot. France 56: 91. 1909.
Sachs. Morphologie des Crucibuhim vulgare Tulasne. Bot. Zeit. 13: 833, pis. 13, 14. 1855.
Tulasne, L. R. and Ch. Recherches sur l'organisation et le mode de fructification des Champignons

de la tribu des Nidulariees. Arm. Sci. Nat., 3rd ser., 1: 41. 1844.
Walker. Development of Cyathus fascicularis, C. striatus, and Crucibuhim vulgare. Bot. Gaz. 70:

1, pis. 1-6 and 3 figs. 1920.
White. The Nidulariaceae of North America. Bull. Torr. Bot. Club 29: 251, pis. 14-18. 1902.

Key to the Genera

Peridioles (or most of them) attached by cords to the cups

Peridium cup-shaped with a thick wall composed of a single layer; peridioles whitish from the thick

white tunica Crucibuhim (p. 181)

Peridium bell-shaped or goblet-shaped, with walls of three obvious layers; peridioles gray or nearly

black; tunica thin or wanting Cyathus (p. 174)

Peridioles not attached by cords, but embedded when fresh in a mucus

Peridium thin, spherical, falling into fragments Nidularia (p. 183)

Peridium thick, cup-shaped, opening at the top Nidula

CYATHUS Haller

Plants distinctly bell-shaped or goblet-shaped; the tough peridium composed of
three distinct layers, in expanding towards maturity exposing a thin, pale, usually
almost smooth epiphram which is stretched across the top and which disappears with
age. Peridioles attached by a cord, black or dark brown, in some cases covered with a
very thin tunica, which, however, does not completely hide their dark color. Basidia
not forming a distinct or homogeneous hymenium, but scattered at irregular heights
throughout a large central area and intermingled with numerous delicate threads which
at maturity have their cell walls greatly thickened and gelatinized to form a solid horny
matrix, throughout which the spores are unevenly scattered. At the time the spores
are growing, this matrix is not horny but gelatinous, becoming horny at maturity.

MDULARIACKAF. 175

Under ordinary powers of the microscope the almost closed lumens of these thick-walled
threads have the appearance of delicate filaments running among the spores. Minute,
amorphous granules are also found in abundance. The hyaline, sclerotic layer, com-
posing the greater part of the wall, is composed of crumpled and distorted cell units
with very thick walls. In Cyutlius these units do not separate when crushed but break
up into irregular masses and bits of cells; in Crucibulum they separate in great part when
crushed and the individual cells can be easily seen and drawn. For other differences
in structure see under Crucibulum vulgare.

The spores of this genus show the remarkable peculiarity of carrying on a large
part of their growth after the basidia that produced them have entirely disappeared.
When the spores are about half grown or less, the basidia become empty and gelatinize,
leaving the spores embedded in the gelatinous matrix mentioned above. As shown in
several figures on plates 122 and 123, the threads producing the matrix can be seen to
wrap about the spores in large numbers. It is obvious that these threads furnish the
nutrition necessary for the post-basidial growth of the spores. This remarkable con-
dition is found also to a somewhat less degree in Pisolithus and Scleroderma, which see.

At the Washington meeting of the Botanical Society of America (Dec. 1926).
Prof. George W. Martin gave a paper on the basidia of the Nidulariaceae in which he
brought out this fact. Our work on the subject had been completed two years earlier.

In addition to the species listed below, Miss White includes from our territory C.
intermedins, referring to it a plant from Delaware. Other collections of this species are
from the tropics, and from Miss White's description the Delaware plants could readily
be referred to C. striates. For C. rufipes see discussion under C. stercoreus.

Key to the Species

Cups distinctly striate inside but not outside; spores 8.3-10.4 x 15-20.5^ C. striatus (p. 175)

Cups distinctly striate both inside and outside, at least after removal of the hairy surface; spores 11-20.3

x 22-40/n C. Poeppigii (p. 177)

Cups even inside

Outer surface covered with matted, woolly hairs
Mouth hardly if at all fimbriate

Spores 11.4-14.8 x 15-18.5^ (8-10 x 10-14/i, Miss White); outer surface of cup dark

chestnut brown C. rugispermus (p. 178)

Spores 18-29 x 20-31^; outer surface of cup soiled drab to paler; cup very slender

C. Lesueurii (p. 179)
Spores larger, 18-30 x 22-35/*; outer surface of cup tawny to gray brown or paler

C. stercoreus (p. 177)

Mouth strongly fimbriate; spores very large, 25-35 x 27-48/x C. melanospermus (p. 179)

Outer surface comparatively smooth, with fine appressed hairs; spores 5-7.4 x 8.6-12m

C. vernicosus (p. 180)

Cyathus striatus Willd.

Cyathia hirsuta (Schaeff.) White

Plates 98, 99 and 121

Plants long goblet-shaped, with a short stem, 8-12 mm. high and 6-8 mm. broad,
growing on twigs, branches, dead leaves, etc., with the stem sometimes attached to the
substratum by a dark cinnamon-brown mycelial pad, as in several other species; outer
surface tawny when quite fresh, becoming more brown, the darker specimens dark
brown (raw sienna), covered with coarse shaggy hairs, often obscurely or when the

176 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

hairs are worn off distinctly striate above; inner surface variable in color, pale to nearly
black, glabrous, shining, distinctly striate; mouth closed by a thin, pale, smooth epi-
phragm that breaks and withers away at maturity. Peridioles confined to lower half
of the cup, 1.5-2 mm. in diameter, usually attached to the cup by a strong, very elastic
cord. Tunica very thin (ll-30/i), often absent in places, but usually obvious, so that
the peridioles appear gray or silvery brown; composed of light brown, collapsed threads;
black layer thin (about 20/*), dense; sclerotic layer 110-365/j thick, its cells vaguely
outlined and not separable as units on crushing.

Spores (of No. 2432) scattered as usual in the central horny tissue, elliptic, hyaline,
rather thick-walled, 8.3-12 x 15-22/1. When young the wall is very thick, becoming
thinner as the spores mature. Basidia long-clavate, 6.8-7.8/* thick, with 4 sessile
spores.

Not rare on dead wood and other vegetable matter. The young plants have
rounded hairy tips, the peridium expanding above and the hairs seceding so as to expose
the thin diaphragm only a little while before full maturity.

For cytological information by Maire covering this and other species, see under
Scleroderma aurantium.

Illustrations: Bolton. Hist. Fungi Halifax 3: pi. 102, fig. 2 (as Peziza strialus).
Dufour. Atlas des Champ. Comest. et Ven., pi. 71, No. 160. 1891.
Eidam. 1. c, pi. 10, figs. 1-22.
Fries, Th. C. E. Sveriges Gasteromyceter, fig. 34.
Gillet. Champ. Fr. (Gasteromycetes), pi. 1.
Hard. Mushrooms, fig. 441.
Hesse. 1. c, pi. 13.
Hollos. 1. c, pi. 28, figs. 7-10.

Michael. Ftthrer f. Pilzfreunde 2: No. 206. 1918 (Second edition).
Micheli. Nova Plant Gen., pi. 102, fig. 2.
Murrill. Mycologia 3: pi. 40, fig. 11. 1911.
Nees von Esenbeck. Syst. Pilze. Schw., pi. 13, fig. 132 A-B.
Walker. 1. c, pi. 4, figs. 34-48.
White. Bull. Torr. Bot. Club 29: pis. 14 and 18. 1902.

68. In ravine northeast of Tenney's on twigs, October 4, 1911.
2432. On top of a rock wall by President's house on twigs and leaves, July 26, 1916.
5280. Deciduous wood in Arboretum, July 5, 1922. Spores 9.3-10.4 x 15-20.5/1.
Also Nos. 2987, 5367, 5911.

Asheville. Beardslee.

Blowing Rock. On log of deciduous tree, Aug. 1922. Coker, coll. (TJ. N. C. Herb., No. 5590).
Buncombe Co. Standley and Bollman, coll. (U. S. Nat'l. Herb., as Cyathia hirsuta).
Haywood Co. On rotting leaves, Aug. 2, 1926. Totten, coll. (U. N. C. Herb.). Spores 8-9.5 i
12.5-17/*.

Alabama. Mobile. Mohr, coll. (U. S. Nat'l. Herb.).
Tennessee. Elizabethtown. Murrill, coll. (N. Y. Bot. Gard. Herb.).
Virginia. Peaks of Otter. Murrill, coll. (N. Y. Bot. Gard. Herb., no name).
Pennsylvania. Trexlertown. Lloyd, coll. (N. Y. Bot. Gard. Herb.).

Ohiopyle. Murrill, coll. (N. Y. Bot. Gard. Herb.). Spores elliptic, 7-9.3 x 11.5-20/1.
New York. Green Co. On twigs. Wilson, coll. (N. Y. Bot. Gard. Herb.).

Massachusetts. Davis, coll. (N. Y. Bot. Gard. Herb, and U. N. C. Herb.). Spores 9.2-11.2 x 13-15/1.
Ohio. Lloyd, coll. (N. Y. Bot. Gard. Herb.).

PLATE 98

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66*

Cyathus rugispermus. No. 7489 [above]. Upper row dry; lower moistened
Cyathus strjatus. No. 5367 [below]

NIDULARIACEAE 177

Cyathus Poeppigii Tulasne

Plate 121

Cups 6-8 mm. high, goblet-shaped, the mouth up to 6 mm. wide, shrinking and
crumpled when dry, dark brown (about sepia), felted and fibrous to shaggy; inner sur-
face slate gray, shining, both surfaces strongly and closely striate-fluted in the upper
half. Peridioles 1.6-2 mm. wide, flat, black, dull or a little shining, the funiculi
conspicuous.

Spores short-elliptic, 11-20.3 x 22-4%.

This species is easily distinguished from others of its group by its large spores and
striated outer as well as inner surface. It is new to the United States, being known
heretofore only from northern South America and the West Indies. There is at the
New York Botanical Garden a collection of this marked "in bamboo pots" (Stevenson,
coll.) with no other locality data given, but probably from the tropics. The spores are
18-37 x 22-55*1.

Illustrations: Lloyd. Myc. Works, pi. 105, figs. 1-5.

Tulasne. Ann. Sci. Nat., 3rd ser., 1: pi. 4, figs. 23-25; pi. 5, figs. 3-4. 1844.
White. Bull. Torr. Bot. Club 29: pi. 14, figs. 1-4. 1902.

Florida. Gainesville. On earth in a yard, July 1924. Couch, coll. (U. N. C. Herb., No. 7282).
St. Croix. A. E. Ricksecker, coll. (N. Y. B. G. Herb.).

Cyathus stercoreus (Schw.) De Toni
Nidularia stercorea Schw.

Plates 100 and 121

Plants goblet-shaped, about 5-10 mm. high in our collections, almost sessile or
with a short stalk, gregarious in large numbers and usually crowded so that a group of
old plants looks very much like a wasp's nest; growing usually on manure or on earth
where manure has been spread. The mycelium is rufous as in all other species we have
seen except in C. Lesueurii, and it sometimes forms an obvious pad at the base of the
cup. Outer surface tawny when young, then tan to buff or gray-brown, covered with
thick, matted, woolly hairs, which with age become worn off, leaving the cups almost
smooth; inner surface smooth, even (not striate), dark grayish brown to nearly black
in age. On expanding above from the rounded, strigose stage of youth, the delicate
epiphragm is exposed, only to break and disappear in a short time; mouth smooth, not
fimbriate (the surface hairs that project above it soon after opening do not constitute a
fimbriate mouth). Peridioles nearly black, without a tunica, 1.5-2.5 mm. in diameter,
with a thick, hard wall, usually attached to the cup by funiculi, although often we
find some of the upper peridioles in the cups which show no signs of a funiculus and
wrhich drop out of the cup when it is inverted (as noticed by Lloyd). Outer removable
layer (tunica?) and denser layer below about 90-130^ thick; sclerotic layer about 37/*
thick around the edge of the peridiole, about 165yu thick on the sides near the center.

Spores (of No. 39) almost hyaline, subglobose to oval, thick-walled, varying greatly
in size, 18-30 x 22-35/j. Basidia (of No. 5916) club-shaped, distinctly swollen at the
distal end, 9-12.5 x 28^4ju, bearing 4-8 sessile spores, only 2 or 3 of which mature.
The spores may be attached to any part of the basidium, but are usually borne in a
circle near the end. After the spores are formed the basidia collapse, finally leaving
long threads among the spores, the layer just beneath the hymenium becoming sclerotic.

Not rare and often in large colonies on manured ground, dry manure, and much
more rarely on decidedly rotten wood. In the large spores, the species approaches

178 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

C. melanospermus, but that has distinctly larger spores on the average, and a fimbriate
mouth. Miss Andrews (1. c.) says that the spores of the type of Nidularia stercorea
in Schweinitz's herbarium are 15.8— 18.7ju, and thinks it perhaps the same as C. Z,e-
sueurii var. minor Tul. We find the spores of the type to be 16.5-22 x 18.5-27^.

Illustrations: Andrews. 1. c, pi. 17, figs. 2 and 6 (as C. stercoretts) ; figs. 1,3,5, and 7 (as C. Lesueurti).
Hard. Mushrooms, fig. 443.
Hollos. 1. c, pi. 28, figs. 11-14.
Lloyd. The Nidulariaceae, pi. 108.

Marshall. Mushroom Book, pi. opposite p. 130 (in error as C. vcrnicosus).
White. Bull. Torr. Bot. Club 29: pi. IS, figs. 17-20 and pi. 16, figs. 1, 2. 1902.

39. On heavily manured ground in Arboretum, Nov. 4, 1908.
5916. On soil in pasture, Nov. 21, 1922.
7516. On rotten place in a live sugar maple, Sept. 23, 1924. Spores 22-30 x 2 7-35/j. This is just like

plant on manure except in habitat.
Also Nos. 97, 377, 5417, 7494.

Asheville. Beardslee.

Linville Falls. Coker and party, No. 5728. On cow manure, August 24, 1922.

Haywood Co. On sawdust by Crawford's Creek, Aug. 2, 1926. (U. N. C. Herb.)

Florida. Couch, coll. (U. N. C. Herb., No. 7284). Spores 24-32 x 28-36M. Several of these plants

had an obvious brown pad at the base.
Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb.).
Mississippi. Underwood, coll. (N. Y. B. G. Herb.).
Virginia. Blacksburg. Murrill, coll. (N. Y. B. G. Herb.).
District of Columbia. Cook, coll. (U. S. Nat'l. Herb.).
New Jersey. (Schw. Herb.).

Newfield. Ellis, coll. (N. Y. B. G. Herb.).
New York. Bronx Park. Murrill, coll. (N. Y. B. G. Herb.).
Ohio. Lloyd, coll. (N. Y. B. G. Herb.).
Ilhnois. Urbana. McDougall, coll. (U. N. C. Herb.). Spores 20-30 x 25-35^.

Cyathus rugispermus (Schw.) De Toni

Plates 98 and 121

Cups short goblet-shaped, about 4—5 mm. high and up to 8 mm. broad, the mouth
in youth convex, at maturity not flaring or fimbriate except that the outer spongy
layer projects a little farther than the inner layer and is minutely eroded; base narrow
the light brown mycelium holding firmly to a bit of earth ; outer surface deep chestnut
brown, felted-tomentose to moderately strigose; inner surface deep brown, shining,
not at all striate. Peridioles 1.6-2 mm. wide, smooth, black, shining, not concave
below, the funiculi represented only by a little central plug at the time found.

Spores (of No. 7458) smooth, oval, thick- walled, 11.4-14.8 x 15-18. 5/*, borne as
usual in the genus.

While the spores of our plants run larger than in the type, as given by Miss White,
other characters seem to fit well except for the darker outer surface of the former.
The species is near C. stercoreus, and Lloyd thinks they are the same. In all of our
collections of C. stercorals the spores, though variable, run much larger than in the
present species. Lloyd finds the spores of Japanese specimens to run as low as 16-24/x,
while for American plants he gives the spores as 30-40/i. On account of the different

PLATE 99

Crucibulum vulcare. No. 7462. X 70
[above]

Cyathus striatus. No. 5911. X 260 [below]

Cyathus striatus. No. 5590. X 70 [above]
Disciseda Candida. No. 7507. X 6 [below]

NIDULARIACI \l 179

habitat and smaller spores, we are for the present retaining C. rugispermus as a species.
Cyaikus rugispermus has been known only from the original collection in Pennsylvania.

Illustrations. White. 1. c, pi. 16, figs. 3-6.

5911. On bank of branch, Xov. 20, 1922. Spores 11-14 x 14-1S.S^.

745S. On bare red . -lay soil by road, July 20, 1924.

7489. On sticks and bare clay. Aug. 8, 1924. Spores 11.2-15 x 15-21/x.

Cyathus Lesueurii Tul.

Plate 123

Plants very slender, about 6-8 mm. high and 3.5 mm. broad at the top, brownish
drab below, dull gray above; upper part fibrous to strigose, lower part felted; mouth
surrounded by long, scattered hairs. Inner surface smooth, blackish brown. Base
without a pad, but the concolorous mycelium holding a ball of earth. Peridioles about
1.7 mm. broad, very thin, nearly black.

Spores (of No. 7281) subspherical, very large, 18-29 x 20-31^-

Gregarious to crowded in fine, sandy soil. This seems to agree prefectly with
C. Lesueurii except that the plants are not quite so large and grow on poor soil without
obvious humus. The shape, however, is more that of the typical form than of the
variety minor. Both Miss White and Lloyd, as well as Hollos, consider C. Lesueurii
var. minor as a synonym of C. stercorals, and in this seem certainly correct. The first
two authors also consider C. Lesueurii as indistinguishable from C. stercoreus, and they
are probably right in this also in so far that intermediates occur. As the present plants
grew on sand, are more slender than in C. stercoreus, and have a peculiar appearance,
we think it well to retain the name for the present.

Tulasne gives the spores of the variety, which he describes from "Carolina," as
17.5-19.8 x 22.3M.

Illustrations: Tulasne. 1. c, pi. 5, figs. 5-10.

Florida. In poor sandy soil by roadside (no obvious manure). Couch, coll. (U. N. C. Herb., No.
7281).

Cyathus melanospermus (Schw.) De Toni

Plates 100 and 122

Cups gregarious or crowded, distinctly infundibuliform, or sometimes somewhat
long-campanulate, 9-12 mm. high by 6-8 mm. broad at the mouth, which is slightly
flaring; outer surface shaggy-hairy, the hairs collapsing and largely disappearing on
long exposure; color in our plants which are old, dark brown, about Mars brown of
Ridgway, in some with a slight tint of purple; inner surface even, smooth, and about the
same color as the outside. Mouth straight and distinctly fimbriate. Peridioles 1.8-2.3
mm. in diameter, almost black, with a rather thick outer layer of coarse, dark brown
fibrils with dark granules between. When wet this coat is easily torn off with a needle,
and underneath is a thin, dark, hard layer of densely packed threads and granules.
Inside this is the much thicker layer of pale sclerotic cells and within this is the hy-
menium which is recognizable only in the young condition. The rather ample "cavity"
in the center is filled with the immense spores lying in a horny (when dry), hyaline
matrix formed of greatly gelatinized threads. Funiculus large, strong, and very
elastic.

180 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

Spores hyaline, round to short-oval, very large and varying greatly in size, 24-35 x
28-48^, some with very thick walls, 3-4/1 thick.

The extremely large average size of the spores of this species is supposed to dis-
tinguish it from others of its group, and in our collection as well as in the type they do
run larger than in C. stercoreus. The average length of ten spores of our No. 23 measured
as one came to them was 34^, while ten of C. stercoreus measured in the same way was

24.9m-

The types in the Schweinitz Herbarium are a good lot of 16 plants densely set
together, shape and appearance about as in C. stercoreus. The only egg well seen was
smooth and black; spores subspherical to oval, 20-31 x 26-38^, which is very little
larger than in C. stercoreus. Miss White gives the spores of the type as 45-55 x 18-45/*.
Lloyd considers the species as a synonym of C. stercoreus and says he "can find no differ-
ence in the general size of the spores from the type material of C. stercoreus and C.
melanospermus. While in the latter there are many large spores, there are also small
ones and some we note measure not over 20 mic." The distinctly fimbriate mouth in
our No. 23 is probably of no specific importance, and in the type the mouth is smooth
or slightly fimbriate. While we retain the species at present, Lloyd is probably right
in regarding it as a form of C. stercoreus.

Illustrations: White. Bull. Ton. Bot. Club 29: pi. IS, figs. 1-4. 1902.

23. On soil and old male cones under pine trees, fall of 1908.

Asheville. Beardslee.

Pennsylvania. Bethlehem. (Schw. Herb., types, and Curtis Herb, from Schw. Herb.)

Cyathus verrucosus De Candolle
Cyathus olla (Batsch) Pers.
Cyathia lentifera (L.) White
Nidularia fascicularia Schw.

Plates 100 and 122

Cups long, goblet-shaped, or bell-shaped, 8-12 mm. high by 7-10 mm. broad at
the moderately or widely flaring mouth. Outer surface when young tomentose spongy
with appressed hairs, becoming nearly smooth in age; color when young brown to
tawny, later becoming paler (grayish brown to dull gray) ; inner surface even, smooth,
dark grayish brown to silvery brown. Mouth even, flaring, incurved and depressed
when young. Peridioles large, disc-shaped, averaging about 2-3 mm. in diameter,
about 0.5 mm. thick in center, concave above, convex below, attached to the cup by
large, strong, whitish funiculi.

Spores (of No. 1688) hyaline, ovate to ovate-elliptic, 5-7.4 x 8.6-12/j.

Wall of the peridium composed as usual of a dark outer region of three parts: a
surface layer (tunica?) about 20/u thick and composed of light brown filaments, mostly
collapsed; a loose median area of many cavities and scattered, nearly hyaline, thick-
walled filaments and numerous granules; an inner black layer about 37 n thick, composed
of dark brown, thick-walled, densely packed threads and numerous granules. The
pale, sclerotic, horny layer beneath is of the usual structure.

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NIDULARIACEAE 181

This plant is distinguished from all our other species by its smaller spores. We
have studied the type of Nidvlaria fascictdaria and find the spores to be oval, 6.5-8 x
8.5-12M.

Illustrations: Andrews. 1. c, pi. 17, figs. 4 and 8.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 35 (as C. olla).
Hard. Mushrooms, fig. 442.
Lloyd. The N'idulariaceae, pi. 110, figs. 1-4. 1906.
White. Bull. Torn Bot. Club 29: pi. IS, figs. 8-13. 1902.

1688. On soil under cedars in Arboretum, September 7, 1915.

Asheville. Beardslee.

Tennessee. Elizabethtown. Murrill, coll. (N. Y. B. G. Herb.).
New York. Westchester Co. Howe, coll. (N. Y. B. G. Herb.).
Wisconsin. Madison. (U. N. C. Herb, and Univ. of Wis. Herb.) Spores 6-8x9-11.5,*.

CRUCIBULUM Tul.

Plants cup-shaped or bell-shaped, sessile, gregarious on twigs and trash in general
or on the bark of logs, or rarely on living trees. Wall composed of a single thick layer
which in youth is velvety-tomentose externally, becoming almost smooth with age.
Inside of the cup covered with a thin, pale lining and without striations; the mouth in
youth covered with a thick, soft veil or epiphragm which disappears with maturity.
Peridioles numerous, almost filling the cup and attached to it by simple funiculi.
Tunica thick, white and quite conspicuous. The basidia forming a distinct and closely
packed hymenium surrounding a narrow central area which at maturity becomes
densely filled with spores. All the cells of the hymenium then become very thick-walled
and fuse together into a solid, hyaline, subhorny tissue, which seems almost structure-
less under low power. The spores are not embedded in cellular tissue, but stick together
by their surfaces and do not fall out when exposed. This organization of the fertile
part of the peridiole is quite different from that in Cyathus, which see.

There is but one species in the genus.

Crucibulum vulgare Tul.

C. criicibuliforme (Scop.) White
Nidularia Juglandicola Schw.

Plates 61, 99, 101 and 122

Plants bell-shaped or cup-shaped, sessile, varying greatly in size, even in the same
collection, from 3-7 mm. high by 4-8 mm. broad at top, but not so crowded as Cyathm
stercorals, growing on trash or twigs of various kinds, bark of decaying logs, or even
on the bark of living trees; when young covered with an ochraceous tawny, velvety
tomentum (much less coarse and shaggy than in Cyathus stercorals), then becoming
almost smooth, brownish, light tan or grayish with age. Inside of the cup smooth and
sometimes shiny, and usually silvery gray to white. The epiphragm, which closes
the cup until maturity, consists of two layers, the outer one felted-tomentose, colored
like the cup, and made up of short, thick hairs with many short, stubby branches;
the inner layer white and made up of an interwoven mass of slender, much branched
filaments. In withering, the outer felted layer of the epiphragm sloughs off first,
leaving the very thin layer which soon also tears and withers away. The peridioles
are lenticular in shape, 1-2 mm. in diameter, attached to the cup by cords and covered

182 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

with a rather thick white tunica which frequently ruptures. Under the tunica is the
thin black layer composed of slender filamentous dark brown cells mixed with a large
amount of granular, amorphous material. Beneath this is a thicker layer (95-1 15m)
composed of hyaline, thick-walled, sclerotic cells, hard and horny when dry but much
softer when soaked for several hours. Nest comes the hymenium which is peculiar
in that the old basidia and intermediate cells also become thick and sclerotic after spore
formation (see fig. by Sachs, 1. c, and our fig. 8). The funiculus is composed of a
delicate fascicle of long, thick-walled, hyaline threads which are rarely branched, and
which have at intervals peculiar joints. At maturity many of the funiculi may have
disappeared, and the peridioles be held in only by a small amount of mucus.

Spores (of No. 647) hyaline, elliptic or oval-elhptic, and varying greatly in size,
3.5-5.5 x 4.3-8.6/i. Basidia (of No. 5277) long-clavate, about 3.7yu thick at the swollen
tip, with 2-4 spores on distinct, slender sterigmata.

A very common and widely distributed plant, growing on wood, twigs and other
vegetable matter. Referring to Lloyd's remark that it rarely if ever occurs on large
logs, we have several collections from the bark of logs and living trees (Nos. 35, 1534,
2904, 2986 and 2987a). One collection (No. 2904) is on sound bark from a large
hickory log. The cups of this collection are a little larger than the average, but agree
in other respects. Lloyd has seen the type of C. fimetarius DC. and considers it the
same as Crucibulum vulgare (Myc. Notes, p. 291).

Illustrations: Eidam. 1. c, pi. 10, figs. 23-30.

Fries, Th. C. E. Sveriges Gasteromyceter, fig. 33.

Hard. Mushrooms, fig. 444.

Hollos. 1. c, pi. 28, figs. 23-26.

Lloyd. The Nidulariaceae, pi. 104. 1906.

Lloyd. Photogravure of Am. Fungi, No. 7.

Micheli. Nova Plant. Gen., pi. 102, fig. 3.

Michael. Fuhrer f. Pilzfreunde 2: No. 205. 1918 (2nd ed.).

Murrill. Mycologia 3: pi. 40, fig. 12. 1911.

Nees von Esenbeck. Syst. Pilze Schw., pi. 13, fig. 133A-B.

Sachs. 1. c, pis. 13, 14.

Sturm. Deutsch. Fl. Pilze 1: pi. 23 (as Cyathus crucibulum). 1814.

Walker. I.e., pis. 5, 6.

White. 1. c, pi. 16, figs. 10-15; pi. 18, figs. 7-13 and 16.

35. On dead bark of a live cedar tree about 2-3 feet from ground in Battle's Park, Jan. 15, 1909.
647. On pile of trash, Oct. 21, 1912.

2904. On sound bark of hickory log, Oct. 10, 1917. Spores 4-6.5 x 6.5-10yu.
7440. On f rondose twigs in Arboretum, July 20, 1924.
Also Nos. 67, 1534, 29S6, 2987a, 3375, 3865, 5277.

Salem. (Schw. Herb., as N. crucibulum.)

Asheville. Eeardslee.

Pink Bed Valley. Murrill and House, colls. (N. Y. B. G. Herb.).

Blowing Rock. Coker. No. 5711.

Waynesville. Standley, coll. (U. S. Nat'l. Herb.).

Alabama. Auburn. Earle, coll. (N. Y. B. G. Herb.).

Virginia. Mountain Lake. Murrill, coll. (N. Y. B. G. Herb.).

Pennsylvania. Bethlehem. (Schw. Herb., as type of N. Juglandicola, and as N. crucibulum.)

Ohiopyle. Ricker, coll. (U. S. Nat'l. Herb.) .
New York. Murrill, coll. (N. Y. B. G. Herb.).

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Ohio. Lima. Dawson, coll. (N. Y. B. G. Herb.).

Illinois. I'rbana. McDougall, coll. (U. N. C. Herb.).

Wyoming. Pitchfork. (U. N. C. Herb.)

Washington. Langlcy. Grant, coll. (V. N. C. Herb.).

Also from many other states in X. V. 15. G. Herb, and U. S. Natl. Herb.

New Zealand. Wellington. Atkinson, coll. (U. N. C. Herb, from Cunningham Herb.). Spores
smooth, oval-elliptic, 3.7-5 x 6.8-9^.

NIDULARIA Fries

Plants subglobose, the peridium wall of one layer and without an epiphram, not
opening regularly or forming a perfect cup, but thin and fragile and breaking up ir-
regularly, leaving the peridioles in an exposed pile on the substratum. Peridioles (in
our species) dark brown to reddish brown, shining, not attached by cords to the peridium
wall, but embedded in a mucus when fresh, the mucus drying down and sticking the
peridioles together after exposure.

Miss White uses the name Granularia Roth, but, as Fries and Tulasne use Nidu-
laria and the latter clearly defines the genus as now used we retain the latter name.
Both names antedate Fersoon.

Key to the Species

Plants minute, sporangioles less than half mm. broad JV. caslanea

Plants larger, sporangioles more than a half mm. broad N. pulvinata

Nidularia pulvinata (Schw.) Fries

Granularia pulvinata (Schw.) Kuntze
Granularia piriformis Roth
Nidularia pisiformis (Roth) Zell.
Nidularia alabamensis Atk.
Nididaria globosa (Ehr.) Fr.

Plate 122

Plants subglobose, small, up to about 6 mm. thick in specimen seen, surface cin-
namon brown, covered with a felted and somewhat powdery tomentum; peridium
excluding the tomentum only one layer thick, that is, there is no thin inner layer that is
stretched over the top as a diaphram before opening. Peridioles small, flattened,
only up to 0.8 mm. wide, dark brown, wrinkled when dry, densely packed, destitute
of a funiculus or other attachment to the wall, but embedded in a mucus when fresh
and damp and lightly stuck together by this mucus when dry. The surface of the
plants may be nodulated from the pressure of the peridioles within. After maturity
the peridium ruptures irregularly and does not form a perfect cup as in the other genera.

Spores (of No. 7588) ovate to subelliptic, hyaline, smooth, 4.2-5.5 x 6-7.4/i, mixed
with irregular, branched and nodulated, thick-walled cells. Basidia 4-spored (Maire,
Comp. Rend. 131 (2): 1248. 1900; as N. globosa).

A rare plant but widely distributed, growing gregariously on wood. It was first
described under this species name from North Carolina, but Lloyd is probably right in
thinking it the same as N. pisiformis of Europe, which has also been described under
many other names. We have examined the type of Nidularia alabamensis and find
that it agrees in all respects with N. pulvinata. The spores are oval with one end
distinctly pointed, 4-5.5 x 5-7.4/u.

184 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

For a long list of supposed synonyms, see Lloyd, 1. c, p. 8. For other cytological
information by Maire covering this and other species see under Scleroderma aurantium.

Illustrations : Fries, Th. C. E. Sveriges Gasteromyceter, fig. 32 (as N. pisiformis) .
Lloyd. Myc. Works, pi. 102 (as N. pisiformis).
Massee. 1. c, fig. 37 (as N. pisiformis) and fig. 81 (as N. conftuens).
White. 1. c, pi. 17, figs. 10-16, 20; pi. 18, fig. 19 (as Granularia puivinala).

7588. On old pine board fence near the power house, Feb. 4, 1925.

North Carolina. Asheville. Beardslee, coll. On old logs. (U. N. C. Herb.)
Alabama. Peters, coll. (Curtis Herb.).

Aubum. (N. Y. B. G. Herb., as type of N. alabamensis.)
Louisiana. Langlois, coll. On a decaying pine log. (N. Y. B. G. Herb.)
New York. Arkville. Murrill, coll. (N. Y. B. G. Herb.).

West Park. Earle, coll. (N. Y. B. G. Herb, and U. N. C. Herb.). Spores oval, 4-5.5 x 5.5-7.5/1.
Connecticut. Litchfield. White, coll. (N. Y. B. G. Herb.).
Maine. Piscataquis Co. Murrill, coll. (N. Y. B. G. Herb.).

Nidularia castanea Ell. & Ev.

Granularia castanea (Ell. & Ev.) White

Plate 122

This interesting little plant, remarkable for its minute size, is known only from the
original locality. Careful search would probably discover it in other parts of the
Atlantic seaboard. The types at New York have been studied and agree well with
Miss White's description. The peridioles as we find them, are dark reddish brown, not
"yellowish brown," and the spores are 4.5-7 x 6-9/x.

We take the following from Miss White (1. c, p. 276) :

"Peridia small, elongate-globose, or subspherical, sessile, sometimes confluent,
0.5-2 mm. long, 0.5-1 mm. high; peridial walls very thin, and brittle at first, pale
yellowish fawn-colored, gradually as the plant matures becoming grayish, and some-
what tuberculose from the inner pressure of the sporangioles, dehiscing irregularly,
the outer covering often breaking away completely, leaving the sporangioles seated
alone on the wood, barely visible without magnification; sporangioles yellowish brown,
very numerous, crowded, circular, flattened above and below, barely 0.25 mm. in di-
ameter, even or slightly creased under high magnification; spores subglobose, hyaline,
thick-walled, 4-7yu long, 3-6/i wide.

"Gregarious on wood."
Illustrations: White. Bull. Torr. Bot. Club 29: pi. 17, figs. 17-19; pi. 18, fig. 20. 1902.
New Jersey. Newfield. Ellis, coll. July 1883. (N. Y. B. G. Herb. Types.)

CALOSTOMATACEAE

Fruit body hypogean or partly so until maturity, attached by obvious basal strands;
with or without a stalk. Peridium highly specialized and formed of several layers.
Gleba without cavities or organized hymenium, separated into small blocks by delicate
tramal plates or strands. Capillitium or its rudiments arising from the whole inner
surface of the inner peridium. Basidia irregularly arranged, stout, club-shaped,
bearing several, usually more than 4, spores irregularly scattered and sessile around its
surface. Spore powder dry and escaping by an apical mouth.

ASTRAEUS Morgan

This genus, formerly included in Geaster and of the same general appearance, was
established by Morgan on the following essential differences: no open chambers and
therefore no organized hymenium ; threads of the capillitium long, much branched and
interwoven, scarcely different from the elemental hyphae of the peridium and con-
tinuous with them, in these respects agreeing with Tylostomd; columella entirely lacking ;
spores much larger than in any Geaster. The hygroscopic character of the rays is very
marked, as is their hard and woody texture when dry.

There is but one species known.

Morgan and Burnap call attention to the apparent relationship of this genus to
Tylostoma and Colostoma and Fischer puts it in the Calostomataceae, in which he has
been followed by others. We are following this precedent, although it is far from clear
that it belongs here rather than in the Tylostomataceae. It seems to us that it would
be better to establish another family for it, as its presence in any recognized one makes
the definition of that family awkward and unsatisfactory. Fischer's definition of the
Calostomataceae includes the statement that the capillitium is well developed. This
is not true of Colostoma, in which the rudimentary "capillitium" disappears entirely before
maturity (see p. 189). Relationship to Colostoma is shown mainly in the organiza-
tion of the gleba into stuffed areas and the stout basidia with a varying number of
scattered, partly lateral spores, and the highly complex organization of the coats. Its
relationship to Tylostoma is indicated by the absence of open chambers, the basidia and
spore characters, though the latter has only 4 spores, and by the highly developed
capillitium, which arises from the entire inner surface of the peridium.

Astraeus hygrometricus (Pers.) Morg.
Geaster hygrometricus Pers.
Geastrum fibrillosum Schw.
Geaster slellatus Scop.
Astraeus slellatus (Scop.) Fisch.

Plates 31, 77, 115 and 117

Plants of small to medium size, growing in colonies just beneath the soil or partially
exposed, arising from a group of black, hairlike, basal rhizomorphs. Outer peridium

185

186 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

thick and composed of several distinct layers (see below) ; surface hard, smooth (except
when very fresh or very old), at maturity splitting into six to many rays which expand
or recurve when wet and fold in again when dry, the inner layer soon becoming much
cracked. Inner peridium pliable, thin, sessile, depressed-globose, about 1.2-3 cm.
broad, surface felted and when in good condition reticulate with fibrous lines, whitish,
becoming gray or brownish, opening by an irregularly torn mouth which is without a
defined peristome. In the fresh condition the plant is tasteless and odorless. Gleba
pure white in youth, divided up into very small, more or less box-like regions by very
thin tramal plates, each chamber closely packed with the fertile, practically homogene-
ous tissue; this mass being made up of branching threads and small fibers bearing more
or less grape-like clusters of nearly spherical basidia, which are crowded amongst each
other, leaving only small unoccupied spaces here and there.

Spores 4-8 to a basidium, sessile on its top and sides, reddish brown, globose,
distinctly tuberculate, 7 .4-10.8/1 thick including the warts. Basidia irregularly ar-
ranged in long strings of clusters, subspherical with a stalk, 11-15.5 x 18-24/x. Capil-
litium threads arising from the entire inner surface of the peridium, including the base,
very pale, long, interwoven, branched, 3-5.5/j thick, smooth or encrusted with granules
and without any obvious tapering terminals; walls very thick, so as to leave no lumen
in many places; no cross walls obvious.

The plant is very persistent. Ripening in the fall, it may be found in good condi-
tion through the winter and spring and the shape is retained for more than a year of
exposure. Like many Geasters, it is loosened from the soil on expanding and may be
rolled about by the wind when dry, the outer layer of the rays slowly wearing away in
thin flakes. The range and adaptability of the species is astonishing. It is apparently
world wide in distribution and almost universal in habitat. Scores may be picked up
on a short walk among the sand dunes of the seashore, and it is among the commonest
fungi of the high mountains. We know of no other plant of which this can be said,
not even the fern Pteris.

In a fresh button practically fully grown but still with white gleba, a section shows
the following characters: a very thin outer flocculent layer of delicate, loosely woven
fibers, which as the plant approaches maturity or is exposed, collapses almost to in-
visibility and soon wears away; next a strong fibrous layer about 1 mm. thick, brownish
toward the surface, whitish inward; next the horny layer which in immature buttons
is very thin but thickens toward maturity to about 0.5-0.7 mm., translucent cartilage
color and sharply contrasting; next a soft, pure white layer which is thickest of all
(about 1.5-1.8 mm.), which splits in the middle at maturity and collapses down to a
thin, scurfy layer on the inside of the rays and the softly felted, irregularly reticulated
layer on the surface of the spore sac. Under the microscope it can be seen that the
outer part of this layer which goes to the rays is denser, firmer, composed of fibers
running parallel to the surface which in the central region turn inward and become more
loosely woven. At the point where the turn is made the splitting takes place. The
part remaining on the rays forms very thin reddish flakes that soon fall off. The in-
nermost layer is very thin (about 0.1 mm.), translucent white, fibrous and tough;
basal attachment of the spore sac 1.5 cm. wide, distinctly limited by the extent of the
fleshy layer. At the base all distinct layers disappear into a homogeneous white tissue
which is about 0.7-1.5 cm. wide. While the plant is often entirely subterranean until
dehiscence, the mycelial attachment is confined to a basal disk one cm. or more broad,
from which rather numerous, small dark fibrils extend into the soil. These are very

CALOSTOMATACEAE 187

easily broken off, and after dehiscence they are usually represented only by dark col-
lapsed lines over the region of attachment, giving this part a roughish appearance.

Microscopic structure of the peridia is as follows: the outer fibrous layer is com-
posed of small and densely packed threads, which in the outer region run parallel to the
surface and curving through a right angle continue through the horny layer perpen-
dicular to the surface. In this last layer the threads are straight, almost perfectly parallel
to each other and very densely packed; continuing into the spongy layer they at once
become looser and much more entangled. The thin inner layer of the spore sac is not
vcrv different from the spongy layer, and from its surface the threads composing it pass
into the gleba without obvious change to form the capillitium.

Maire makes the peculiar statement that the basidia are terminated by a "sort of
single compound sterigma at the top of which the spores are borne on very short pedi-
cels." He gets the number of spores right. Tulasne gives the basidial shape right,
but the number of spores as four. As they are of considerable interest, we quote below
in full Maire's observations on this species (Comp. Rend. 131 (2): 1248. 1900):

"Gcaster hygromctricus shows basidia with 2-8 spores. The fusion nucleus under-
goes two successive divisions; sometimes the divisions stop there, and there are formed 4
spores, each of which receives a nucleus. At other times one or two of the 4 nuclei
divided again, from which result 5 spores and 5 nuclei or 6 spores and 6 nuclei; often
the 4 nuclei divide and there result 8 spores and 8 nuclei. Sometimes the nucleus of the
basidium undergoes only one division, whence two nuclei and two spores. The most
usual case seems to be the 6-spored basidium. On the other hand, the formation of the
spores is, up to a certain point, independent of that of the nuclei: it happens, for
example, that a basidium having only 4 nuclei forms 5 spores, etc. These processes
are very variable, quite as in the basidia of the Cantharellaceae which we are studying
at this time. However that may be, in Geaster, the formation of the spores always
takes place in the following manner: at the tip of the basidium is sent out a single
prolongation, a sort of compound sterigma, at the summit of which the spores are
formed on very short pedicels. The nuclei, of definite number, are drawn all together
into the compound sterigma, to be separated afterwards, each in its spore. In Geaster,
the centrosomes and the kinoplasm are scarcely visible."

For other cytological information by Maire, covering this and other species, see
under Scleroderma auranlium.

Up to the present the attachment of this plant to the soil has been completely
misunderstood. This is hard to understand as the basal rhizomorphs are so conspicu-
ous. DeBary's figure (Fungi, Myc. and Bact, fig. 146) shows no basal attachment
and he says the button is attached to a felted mycelium which often spreads in the soil
for the distance of an inch all around. This description completely reverses the facts
and the error has never been corrected. The delicate flocculence of the surface of the
buttons is so little connected with the soil that they come up almost clean. This is
entirely different from the intimate attachment to the soil of the mycelial surface layer
of hypogeal Geasters. It will be noted that this is an additional character pointing
to a relationship with Colostoma and Tylosloma.

We have examined the type of Geastrum fibrillosum in the Schweinitz Herbarium,
and find that it is, as Lloyd says (Geastrae, p. 10), a weathered specimen of this species.
It is the flaky roughness of the outer surface that Schweinitz referred to in his name,

188 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

not the felted reticulations of the inner peridium, as stated by Lloyd. The spores of
this specimen are roughened, 7.4-10/j thick. Lycoperdon recolligens Woodward (Trans.
Linn. Soc. 2: 58. 1794) seems to include both the present species and Geaster mam-
mosits. He refers to Schmidel's plates 27 and 28 which are of A. hygrometricus.

Illustrations: Destrie. Cited on p. 104, pi. 10, fig. B.
Hard. Mushrooms, fig. 483.
Hollos. 1. c, pi. 10, figs. 26-29.
Marshall. Mushroom Book, pi. opposite p. 130.
Michael. Fiihrer f. PiMreunde 2: No. 204. 1918 (2nd ed.).
Micheli. Nova Plant. Gen, pi. 100, fig. 6.

Morgan. Journ. Cin. Soc. Nat. Hist. 12: pi. 2, figs. 8-11; Amer. Nat. 18: fig. 12.
Nees von Esenbeck. Syst. Pilze Schw., pi. 12, fig. 127.
Petri. Flora Ital. Crypt. (Gasterales), Fasc. 5: figs. 80-82. 1909.
Rea. Trans. Brit. Myc. Soc. 3: pi. 17. 1912.
Schmidel. Icon. Plant., pis. 27 and 28, figs. 19-31. 1793.
Vittadini. Monog. Lycoperd., pi. 1, fig. 8. 1842.

73. Mixed woods with cedars, Oct. 11, 1911.
673, 685. Old collections (Chapel Hill). No data.
7237. Underground in open, sandy field, Jan. 14, 1923.
7441. Sandy soil in woods, July 19, 1924.

Smith Island. Sandy soil, April 1918. Couch and Grant, colls. (U. N. C. Herb., No. 3007).

Spores 7.7-9.3/*.
Blowing Rock. Coker and party, Aug. 1922. Nos. 5786 and 5823.
Pink Bed Valley. Murrill and House, No. 400. July 1908. (N. Y. Bot. Gard. Herb.).

South Carolina. Ravenel. Fung. Car. Exs. No. 75. (Philadelphia Academy.)

North Island. Coker and party, colls. (U. N. C. Herb.). Plentiful among the sand dunes.
Florida. Alachua Co. Couch, coll. (U. N. C. Herb., No. 7301).

CALOSTOMA Desv.

Young fruit bodies formed just beneath the surface and basally attached; at
maturity pushed by the elongation of the very peculiar stalk which is composed of a
large number of anastomosing, firmly gelatinous strands woven together to make a
spongy, more or less cylindrical column. Before elongation the entire plant is enclosed
in a gelatinous layer called the volva by Burnap which at maturity becomes so watery
as to fall away, exposing the spore-bearing head. This is subspherical and is composed
of the outer peridium, the inner peridium, and the spore sac. The former is composed
of two layers, an outer thicker one which is much like the stalk in structure and in
most species becomes strongly or moderately gelatinized, and a thinner, denser inner
layer which does not gelatinize. This exoperidium breaks into pieces after the plant
emerges and falls off completely or in part, exposing the thin, dense, tough, and, when
dry, hard and horny inner peridium which remains inflated and is very permanent.
On the top of the inner peridium are elevated folds, like a puckered mouth, arranged in
a more or less radiating way into several lobes which open down their centers by narrow
slits leading into the spore sac within. This chamber is formed from the thin, soft and
papery innermost layer of the inner peridium which is at first in contact with the
next outer layer, but which soon begins to dry and contract away from the adjoining
layer except above, and gradually getting smaller while hanging pendent within the
firmer outer case it forces the dry spores and fragmentary remains of the accompanying
fibers out of the slits for dispersal. The gleba when young has a lobulated appearance
and is composed of solid masses of fertile hyphae with the basidia borne irregularly

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CALOSTOMATACEAE 189

throughout; between these masses run loosely woven fascicles or plates of branched
threads with broad annular thickenings that are not obviously connected with the
fertile hyphae and which entirely disappear before the maturity of the spores (Burnap,
p. 1S3). Basidia ovate or pyriform or short-clavate, irregularly arranged throughout
the fertile tissue. Spores sessile, variable in number (5-12), irregularly placed on
both ends and sides of the basidia (Burnap, p. 1S5), oblong-elliptic to globose, walls
usually pitted.

This peculiar genus is complex in structure and very striking in appearance.

The ephemeral threads with annular thickenings suggest a capillitium and are
almost certainly homologous to it. They are so-called by Fischer, but it seems best
to us to reserve this term for the threads found in mature fruit bodies. To say, as he
does, that Colostoma possesses a well developed capillitium is misleading. In the young-
est plant we have (C. cinnabarina) which is unexpanded but with spores about mature
there are only a few detached fragments of this earlier system. These fragments show
much narrower thickenings than those shown by Burnap.

Rev. Edward Hitchcock has published a non-technical article under the head of
Physiology of Gyropodium coccincum (Amer. Journ. Sci. and Arts 9: 56, pi. 3. 1825),
describing a Colostoma. He speaks of the genus and species as recently established by
Schweinitz, but the latter does not seem ever to have published this name, as noted
by Hitchcock farther on. It does not seem to be known how Hitchcock got the name
from Schweinitz. The species he described is doubtful. Burnap thinks it C. cinna-
barina; Saccardo, lutescens.

Mitrcmyccs Nees is the same as Colostoma.

Literature

Atkinson. A New Species of Colostoma. Journ. Myc. 9: 14. 1903.
Burnap. Notes on the Genus Colostoma. Bot. Gaz. 23: 180, pi. 19. 1897.
Fischer, in Engler and Prantl. Pflanzenfamilien l1: 339, fig. 177. 1900.
Lloyd. Myc. Notes, p. 123, pis. 8 and 9. 1903.

Lloyd. The Genus Mitremyces. Myc. Notes, p. 238, pis. 68 and 69. 1905.
Massee. A Monograph of the Genus Colostoma. Ann. Bot. 2: 25, pi. 3. 1888.
Morgan. Journ. Cin. Soc. Nat. Hist. 12: 20, pi. 2A. 1889.
Webster. Notes on Colostoma. Rhodora 1: 30. 1899.

Key to the Species

Spores 3.5-5 x 6-10 C. microsporum (p. 192)

Spores larger

Spores elliptic; exoperidium on dehiscing falling away entirely, the outer part forming a thick soft
jelly; entire inner surface of the exoperidium and outer surface of the endoperidium bright

red when fresh C. cinnabarina (p. 189)

Spores elliptic (a few may be spherical) ; exoperidium not gelatinous, the inner layer remaining as

scales on the endoperidium; red color confined to the mouth region. C. Ravenelii (p. 191)

Spores spherical; exoperidium composed of a firmly gelatinous (not watery) outer layer and a corky

inner layer, on dehiscing a large part remaining as a collar around the base of the

endoperidium; red color confined to the mouth region C. lutescens (p. 190)

Calostoma cinnabarina Desv.

Plates 100 and 123

Rooting stalks stout, rooting, usually short and often scarcely lifting the peridium
above ground, dull, sordid ochraceous and soft gelatinous. Exoperidium when fresh

190 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

and wet composed of a thick, slimy, transparent outer coat and a thin, bright red, non-
gelatinous inner coat, the whole breaking up together into large or small pieces which
curl inward and soon fall completely away, looking like red seed in a gelatinous pulp,
and leaving the thin inner peridium covered with a fine cinnabar red powder or scurf
which as it gradually wears away leaves the peridium surface paler until in old speci-
mens the red may be nearly gone. Mouth composed of about 5 elevated ridges which
are deep red until very old. Spore sac light clear yellow.

Spores (of No. 5521) pale yellow (about cream color), very distinctly pitted (pits
visible when x 670), oblong-elliptic, 6.3-8.5 x 14-20/x, rarely up to 24/^ long.

Distinguished by short, bulky stalk, cinnabar red color of the endoperidium and
inner layer of the exoperidium, completely deciduous exoperidium, and elliptic spores.
This is a very abundant plant at higher elevations in the eastern United States, rare
or absent in the coastal plain.

Illustrations: Burnap. Bot. Gaz. 23: pi. 19, figs. 3-10. 1897.
Lloyd. Myc. Works, pi. 8, figs. 1-10.
Marshall. Mushroom Book, pi. opposite p. 132.
Morgan. 1. c, pi. 2A (as C. lulescens).

North Carolina. Hillsboro. (Curtis Herb.).

Blowing Rock. Coker and party, August 1922. No. 5521 and No. 5615. In earth by paths.

Very common.
Little Switzerland. Miss Osborne, coll. No. 2835. Sept. 21, 1917.
Brevard. Miss Cornelia Love, coll. September 1922.
Hendersonville. Alma Holland, coll. (U. N. C. Herb.).
Winston-Salem. Schallert, coll. No. C-42. On soil in shady places.

Alabama. Montgomery. Burke, coll. (N. Y. B. G. Herb.).

Mississippi. Trice, coll. (Curtis Herb., as M. lulescens). Spores oblong in sketch.

Virginia. Apple Orchard Mountain. Murrill, coll. (N. Y. B. G. Herb.).

Pennsylvania. Buck Hill Falls. Mrs. Delafield, coll. (N. Y. B. G. Herb.).

Also represented in the Curtis Herbarium from the Schw. Herbarium. (No data.)

Calostoma lutescens (Schw.) Burnap

Plates 103 and 123

Rooting stalk long (up to 6 cm.), compactly woven, yellowish, firmly gelatinous
when fresh, clearer light yellow when dry. Exoperidium composed of two layers, an
outer, firmly gelatinous, fibrous-spongy one about 1 mm. thick when wet which is the
color and consistency of the stalk and continuous with its surface, and an inner one a
little less than 1 mm. thick which is clear pale yellow, opaque and much more compact
and homogeneous (about like orange peel), the two layers separable without much
difficulty, but dehiscing together and remaining united. After the elongation of the
stem this outer peridium breaks into several (usually) irregular apical segments which
fall away and into 5 or 6 or more irregular basal segments that curl outward and usually
remain expanded about the base of the endoperidium like a fluted collar or less often
break off close to the peridium and fall away. Endoperidium subglobose, pale orange-
yellow (not red) , smooth, dull, pliable when wet, horny when dry, 1-1 .5 cm. in diameter.
Mouth consisting of 5 or 8 elevated, tooth-like lobes which are bright red within when
fresh, the outside paler red and the whole fading somewhat with age. Spore sac
bright yellow.

Spores (of No. 1935) light yellow, globose, the walls thick and distinctly pitted,
giving the spore the appearance of a golf ball, 6-8,u thick.

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This is an elegant species and easily distinguished by the long stalk, light yellow
color throughout except the red teeth, fluted collar, and round spores. The surface
layer of the exoperidium is intermediate between that of the other two species, as it is
quite obviously gelatinous but thin, and does not form a thick, clear, slimy jelly as in C.
cintubarino.

Schweinitz's material labelled Milremyces lutescens consists apparently of two
species. There are two lots of two plants. The upper two with long stems are fused
and the spore sacs are largely broken off, with no spores to be found. These, so far
as they go, look just like if. lutescens. The other two consist of a smaller plant with the
top nearly gone and the upper part of another with perfect sac and spores. This looks
just like Rd'.rnclii, with warts on the surface of the sac, and, according to Lloyd (Myc.
Xotes, p. 395), with elliptic spores. We are glad to say, however, that the part of
Schweinitz's plants taken by Michener and now in his herbarium in Washington is
C. lutescens as generally accepted and agree with the description, the spores being
spherical, pitted, 6.5-8.2/x thick. The round spores and yellow spore sac and spores
of the description (Syn. Fung. Car. Sup. No. 345) indicate clearly the species as now
generally accepted. Schweinitz's figures do not show the fluted collar which is one of
the most marked characters of this species (though not always conspicuous) and they
may really represent C. Ravenclii.

Atkinson reported this species from Blowing Rock (this Journal 9: 85. 1892),
but later said that his plant was C. cinnabarina (Mushrooms, p. 213).

Illustrations: Burnap. Bot. Gaz. 23: pi. 19, figs. 1, 2. 1897.
Corda. Anleitung, pi. D, fig. 41.
Lloyd. Myc. Works, pi. 9, figs. 1-3.
Marshall. Mushroom Book, pi. opposite p. 132.
Schweinitz. Syn. Fung. Car. Sup., pi. 1, figs. 1-6.

1935. In mixed woods, Oct. 25, 1915.

1995. Under oaks south of campus, Nov. 23, 1915.

Asheville. Beardslee.

Pink Bed Valley. Murrill and House, colls. (N. Y. B. G. Herb.).

Alabama. Moulton. Peters, coll. (Curtis Herb., as Milremyces). Spores spherical.

Auburn. Underwood, coll. (N. Y. B. G. Herb.).

Mobile. Mohr, coll. (U. S. Nat'l. Herb., as Milremyces).
West Virginia. A. H. Curtis, coll. (Curtis Herb., as Milremyces).

Calostoma Ravenelii (Berk.) Massee

Plates 104 and 123

Rooting stem 2-5 cm. long, stout, rather compact, consisting of an interwoven
mass of thick, gelatinous fibers of a soaked clay color; exoperidium light clay or straw
color, not gelatinous at any stage, not obviously two-layered, surface spongy or fibrous
looking, less than 1 mm. thick; on exposure breaking up into small pieces except for a
larger, fluted, apical part which lifts up and falls away, the laiger particles on the sides
also falling away usually, but some of the smaller ones remaining attached to the
peridium for a long time until slowly worn away. In addition to these fragments
which are the full thickness of the exoperidium, the endoperidium is covered with much
thinner and smaller scales which are parts of the inner layer of the exoperidium from
which the outer part has broken away. These scales are very persistent and are found

192 THE GASTEROMYCETES OF THE UNITED STATES AND CANADA

on almost all herbarium specimens. When the exoperidial cap falls off, that part of
the inner surface which was touching the mouth ridges is seen to be red, other parts
of the inner surface are a distinct light yellow, as is also the freshly exposed surface of
the inner peridium, the color being due to minute, inherent, bran-like scales. These
scales get larger downward and merge into the larger scales above mentioned. Endo-
peridium tan or clay color, at length brown, the exposed surface dull and somewhat
glaucous-looking, 7-18 mm. in diameter when fresh, 5.5-11 mm. when dry; mouth
composed of about 5 rays, bright red when fresh, fading with age. Spore sac white.
Spores (of No. 1488) white, clear, with one or two large oil drops, variable in size
and shape, usually oblong-elliptic, a few nearly spherical, 6.5-7 x 10-1 7m; surface ap-
pearing smooth except under high power, then seen to be very minutely and closely
pitted, showing faint radial lines through the walls.

This species is easily distinguished by the non-gelatinous exoperidium, clay colored
endoperidium spotted with scales and particles, and the very faintly punctate spores.
Mitremyces Tylerii Lloyd (Myc. Notes, p. 240) is probably only a small form of this
species. One finds at times very small plants without obvious stalks mixed with more
typical ones.

Illustrations: Berkeley. Trans. Linn. Soc. 22: pi. 25 B.

Lloyd. 1. c, pi. 9, figs. 4-6; pi. 68, figs. 3S.

Marshall. Mushroom Book, pi. opposite p. 132 (good).
Massee. 1. c. (1903), pi. 3, figs. 26-28.

58. Among moss on hillside in open mixed woods, Sept. 27, 1910. Spores 6.5-8 x 8.6-14^.
1488. On ground in open woods, Dec. 7, 1914.
2872. By path in Battle's Park, Oct. 5, 1917. Spores variable in size and shape, distinctly pitted,

4.8-6.6 x 8.6-18m.
AkoNos. 119a, 1518, 1519.

Asheville. Beardslee, coll. Aug. 1918. Spores 6.2-7.4 x 11-17^.
Columbus Co. Barlow, coll. No. 49. July 1909.
Caesar's Head. Ravenel, coll. (Curtis Herb., co-types).

Alabama. Moulton. Peters, coll. (Curtis Herb., as M. lutescens) Spores oblong (as shown by sketch)

and case warted. •

Mobile. Mohr, coll. (U. S. Nat'l. Herb., us Mitremyces).
Kentucky. Conway. B. Fink, coll. (N. Y. B. G. Herb.).
Delaware. Wilmington. Commons, coll. (U. S. Nat'l. Herb. , as M. lutescens) .
Pennsylvania. Center Co. Adams, coll. (N. Y.|B. G. Herb.). Spores elliptic, obscurely pitted,

6-7.5 x 8-13/..

Calostoma microsporum Atk.

The following description is from Atkinson (Journ. Myc. 9: 16. 1903):

"Plants 4-7 cm. high, foot stalk 3-6 cm. by 1-2 cm., cylindrical or ventricose or
enlarged below, sometimes compressed, rarely two foot stalks joined throughout the
entire length. Peridium oval, 10-15 mm. broad; teeth 5-7, prominent, vermilion
colored on their inner faces; exoperidium separating into numerous small hard adherent
warts, covering the middle and lower surface of the endoperidium, usually entirely
separating from the upper surface, leaving a smooth area on the inner peridium around
the mouth. Spores white, smooth, oblong, some rarely elliptical, 6-10 x 3.5-5ji.
Protoplasm usually homogeneous, sometimes granular, often showing a tendency to
be constricted at the middle, perhaps because of a clear area at this point."

2
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8

lAl.OMOM VTAC'KAK 19-5

For a comparison between this species and C. Ravcnclii, see Atkinson's discussion,
which is rather long for quotation. The most important differences that seem to
distinguish the two species are the longer stalk (on the average), the smaller spores and
the vermilion color being confined to the inner faces of the stomal teeth in C. micro-
sporum. We have not seen Atkinson's plants and cannot give an opinion on the
validity of the species.

Tennessee. Rugby. Percival, coll. (Cornell Univ. Herb., t>pe).

LITERATURE

Berkeley. Sur la fructification des genres Lycoperdon, Phallus, et de quelques autres genres voisins.
Ann. Sci. Nat., 2nd ser., 12: 160, pi. 2. 1839. There is a larger work on this group in Russian
with a German resume printed in Moscow and St. Petersburg in 1902.
Bigeard. Flore des Champignons 2: 513, pis. 36, 37, and pi. 38, fig. 1. 1913.
Bucholtz. Beitrage zur Morphologie und Systematik der Hypogaeen (Tuberaceen und Gastromyceten).

Riga, 1902.
Bucholtz. Zur Morphologie und Systematik der Fungi Hypogaei. Ann. Myc. 1: 152, pis. 4 and 5.

1903.
Corda. Icon. Fung. 5: 22. 70, pis. 4, 6, 7, and pi. 8, fig. 51. 1842.
Cunningham. The Gasteromycetes of Australasia. Proc. Linn. Soc. N. S. W. 49: 97. 1924; SO:

245,367. 1925; 51: 72, 363, 627. 1926.
De Bary. Comparative Morphology and Biology of the Fungi, Mycetozoa and Bacteria, p. 308,

Oxford, 1887.
Fischer in Englerand Prantl. Pflanzenfamilien 1, pt. 1 : 276, figs. 126-182. 1900.
Fitzpatrick. A comparative study of the development of the fruit body in Phallogasler, Hysterangium,

and Gautieria. Ann. Myc. 11: 119, pis. 4-7. 1913.
Fries R. E. Einige Gasteromyceten aus Bolivia und Argentinien. Arkiv. for Botanik 8, No. 11:

1, pis. 1-4. 1909.
Fries. Th. C. E. Sveriges Gasteromyceter, 44 text figs. Arkiv. for Botanik 17: 1921.
Harkness. New Species of California Fungi. Bull. Cal. Acad. Sci. 1: 29. 1884.
Harkness. Californian Hypogaeous Fungi. Proc. Cal. Acad. Sci., 3rd ser., 1: 241, pis. 42-45. 1899.

Also in Rev. Myc. 22: 82. 1900.
Harvey. Contributions to the Gasteromycetes of Maine. Bull. Torr. Bot. Club 24: 71. 1897.
Hesse. Mikroscopische Unterscheidungsmerkmale der typischen Lycoperdaceengenera. Jahrb.

Wiss. Bot. 10: 383, pi. 28. 1876.
Hesse. D e Hypogaeen Deutschlands. Halle a. S., 1890-94.
Hollos. Die Gasteromyceten Ungarns. Leipzig, 1904.

Kauli'man. Outline of the Gasteromycetes of Michigan. Rept. Mich. Acad. Sci. 10: 70. 1908.
Lloyd. Myc. Notes Nos. 7-9, figs. 30-56. 1901-2.

Lloyd. The Genera of Gastromycetes, figs. l-14andpls. 1-11. Cincinnati, 1902.
Lloyd. The Curtis Collection. Myc. Notes No. 15: 152. 1903.
Lloyd. The Lycoperdaceae of Australia, etc., pis. 25-39 and figs. 1-49. 1905.
Lloyd. Sur Quelques Rares Gasteromycetes Europeen. Myc. Notes No. 22 : 261, figs. 98-111 and pis.

28, 35, 72, 87-89. 1906.
Lloyd. The Gastromycetes of Schweinitz's Herbarium. Myc. Notes No. 30: 395. 1908.
Lloyd. The Lycoperdaceae of Vittadini. Myc. Notes No. 70 : 1222. 1923.
Lohwag. Zur Stellung und Systematik der Gastromyceten. Verhandlg. der Zool.-Bot. GeselL in

Wien 74: 38. 1924.
Lohwag. Zur entwicklungsgeschichte und morphologie der gasteromyceten. Ein beitrag zur systema-
tik der basidiomyceten. Beih. Bot. Centralbl. abt. II, 42: 177-334, pis. 1-2. 1926.
Long. Notes on New Or Rare Species of Gasteromycetes. Mycologia9: 271. 1917.
Lotsy. Bot. Stammesgeschichte 1: 718. 1907.

Maire. Sur la cytologie des Gasteromycetes. Comp. Rend. 13 1 (2) : 1246. 1900.
Massee. A Monograph of the British Gasteromycetes. Ann. Bot. 4: 1, pis. 1-4. 1889.
Micheli. Nova Plantarum Genera. 1729.

Morgan. Gastromycetes. Journ. Cin. Soc. Nat. Hist. 11: 141, pi. 3. 1889; 12: 8, pis. 1, 2. 1889;
12: 163, pi. 16. 1890; 14: 5, pis. 1, 2. 1891; 14: 141, pi. 5. 1892.

194

LITERATURE 195

Petri. Flora Italics Cryptogama (Gastcrales), Fasc. 5. 1909.

Rea. British Basidiomycetac, p. 21. 1922.

Rchsteincr. Bcitr. zur Entwickclungsgeschichte der Fruchtkorper ciniger Gastcromyceten. Bot.

Zeit. SO: 761, 777, 800, 823, 843, 865. 1892.
Saccaido. Sylloge Fungorum 7: 1, 469. 1888; 9: 262. 1891; 11: 152. 1895; 14: 254. 1899;

16: 224. 1902.
Schroeter. In Cohn's Kryptogamen Flora von Schlesien 3, pt. 1 : 689. 1888.
Trcleasc. Morels and Puff-balls of Madison. Trans. Wis. Acad. Sci. Arts and Letters 7: 105, pis.

7-9. 1888.
Tulasne, L. R. and Ch. De la fructification des Sc'eroderma, compart a cclle des Lycoperdon et des

Batista. Ann. Sci. nat., 2nd. ser., 17: 5. 1842.
Tulasne, L. R. and Ch. Champignons hypog£s de la famille des Lycoperdacees. Ann. Sci. Nat.,

2nd. ser., 19: 373. 1843.
Tulasne, L. R. and Ch. Fungi nonnulli hypogaei, novi vel minus cogniti. Parlatore, Giorn. Bot. 2:

55. 1844
Tulasne. L. R. Fungi Hypogaei. Histoire et monographic des champignons Hypoges. Paris, 1851.
Vittadini. Monographic Tuberacearum. Mediolani, 1831.

Vittadini. Monographia Lycoperdineorum. Mem. Accad., Torino, 5: 145, pis. 1-3. 1843. Re-
viewed by Tulasne in Ann. Sci. Nat., 2nd ser., 19: 277. 1843.
Winter. Gastcromycetes. In Rabenhorst's Krypt.-Fl. Deutsch. 1, Abt. 1: 864. 1884.
Zopf Die Pike. p. 362. 1890.

PLATE 105

PLATE 105

Fig. 1. Clathrus columnatus. No. 4949.

Fig. 2. Simblumsphaerocephalum. No. 1427.

Fig. 3. Mutinus Ravenelii. No. 741.

Fig. 4. Mutinus Curlisii. No. 5113.

Fig. 5. Ithyphallus Ravenelii. No. 41a.

Fig. 6. Ithyphallus Ravenelii. No. 649.

Fig. 7. Dictyophoraduplicata. No. 5195.

Figs. 8 and 9. F 'roto phallus jamaicensis. Cinchona, Jamaica.

Figs. 10 and 11. II ysterangiumdathroides. No. 7416.

Figs. 12-14. Hysterangium pompholyx. No. 7490a. Fig. 14 shows one of the flocculent threads on

the peridium.
Figs. 15 and 16. Hysterangium sloloniferum\a.r.americanum. Ithaca, New York.
Figs. 17-19. Gymnomyces vesiculosa. No. 7470. Fig. 18 shows 4 spores held together by a sort of

gelatine; fig. 19 shows cross section of a wall of the glebal chambers.
Fig. 20. Gautieria monticola. California.

Figs. 1-5, 7, 9, 10, 12, 15, 18, 20 X 1620; figs. 6, 8, 11, 13, 16 X 1013; figs. 14, 17 X 810; fig. 19 X 188.

PLATE 105

PLATE 106

PLATE 106

Figs. 1-6. Sclerogastcr minor. No. 7474.

Fig. 7. Rhizopogon rubescens. No. 6003.

Fig. 8. Rhizopogon rubescens. No. 7450.

Fig. 9. Rhizopogon rubescens. No. 7204.

Fig. 10. Rhizopogon rubescens. No. 58.

Figs. 11 and 12. Rhizopogon atlanticus. No. 5999.

Fig. 13. Rhizopogon truncatus. Type.

Figs. 14 and 15. Rliizopogon luteolits. No. 1607 (vonHohnel).

Figs. 16 and 17. Rhizopogon luleolus. No.7359 (Fitzpatrick).

Figs. 1-5, 7-9, 11, 15, 17 X 1013; others X 1620.

PLATE 106

PLATE 107

PLATE 107

Figs. 1-4. Rhizopogon nigrescens. No. 7582 (showing bits of hymenium and basidia) .

Figs. 5 and 6. Rhizopogon nigrescens. No. 7372.

Fig. 7. Rhizopogon nigrescens. No. 1910.

Fig. 8. Rhizopogon nigrescens. No. 976.

Figs. 9-11. Rhizopogon roseolus. No. 7207.

Fig. 12. Rhizopogon roseolus. No. 598 (Whetzel).

Fig. 13. Rhizopogon roseolus. No. 7214.

Fig. 14. Rhizopogon piceus. No. 6059.

Figs. 15 and 16. Rhizopogon maculatus. No. 955.

Figs. 1, 2 X 825; figs. 3-7, 9, 10, 12, 13, 15 X 1013; Figs. 8, 11, 14, 16 X 1620.

PLATE 107

PLATE 108

PLATE 108

Figs. 1 and 2. Melanogaster nauseosus. Sketch of a section showing peridium, tramal plates and
cavities, some partly filled with spores and others with spores just beginning to form. X 34.
Fig. 2, basidia with spores, basidia beginning to gelatinize. X 900.

Fig. 3. Nigropogon asterosporus. Sketch showing continuity of the tramal plates with the peridium.
X34.

Figs. 4-6. Sections of hymenium and subhymenium of the above, showing young basidia, basidia with
spores and collapsed basidia. X 900.

Fig. 7. Spores of above. X 1440.

Fig. 8. Leucogaster carolinianus. Young and mature basidia. X 900.

Fig. 9. Mature spores of above, showing heavy reticulum. X 1440.

PLATE 108

wSJ^.-i- w,^-.*^—1 !*??#?^

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W*

f« '■ ...... .,-.■■'

PLATE 109

PLATE 109

Figs. 1 and 2. Melanogaster rubescens (?). No. 5804.

Fig. 3. Melanogaster variegalus. Type.

Figs. 4 and 5. Melanogaster variegalus. Ithaca, N.Y.

Fig. 6. Melanogaster variegalus. Hungary (Hollos).

Figs. 7 and 8. Melanogaster ambiguus. Syracuse, N. Y.

Fig. 9. Uymenogasler Thwailesii. Syracuse, N. Y. (Povah.No. 832).

Fig. 10. Hymenogaster Thwailesii. Type.

Figs. 1, 4, 7 X 1013; others X 1620.

TLATE 109

PLATE 110

PLATE 110

Fig. 1. Hymenogasterfoetidus. No. 7473.
Figs. 2-4. Hymenogasterfoetidus. No. 7467.
Fig. 5. Octaviania Ravenelii. No. 7200.
Fig. 6. Octaviania Ravenelii. No. 6002.
Fig. 7. Octaviania Stephensii. Type.
Fig. 8. Octaviania asterosperma. Ithaca, N. Y.

Figs. 1 and 2 X 1013; figs. 4, 6 X 503; others X 1620.

PLATE 110

PLATE 111

PLATE 111

Figs. 1-11. Odaviania purpurea. No. 7468. Figs. 4-11 show nuclear behavior in the basidia.

Figs. 12, 13. Secotiumagaricoides. No. 7091.

Fig. 14. Secotium arizonicum. Type.

Fig. 15. Rhopalogastertransversarium. Alabama (Earle, coll.).

Figs. 1, 13-15 X 1620; figs. 2-11 X 1013; fig. 12 X 810.

PLATE 111

PLATE 112

PLATE 112

Figs. 1-3. Cahatia rubro-flam. No. 7154 (showing basidia and capillitium thread with thin places

in the wall).
Fig. 4. Cahatia maxima-. Wisconsin.
Figs. 5 and 6. Cahatia cyathiformis. No. 550.
Figs. 7 and 8. Cahatia data. No. 2843 H.
Fig. 9. Cahatia cretacea. Abisco, Lapland.
Fig. 10. Cahatia fragilis. No. 4783. Canada.
Figs. 11 and 12. Cahatia craniformis . No. 538.
Fig. 13. Cahatia craniformis. Type.
Figs. 14 and 15. Cahatia caelala. New York.
Fig. 16. Lycoperdon atropurpitreitm. No. 1019.
Fig. 17 '. Lycoperdon pulcherrimum. No. 7560.
Fig. 18. Lycoperdon echinatum. No. 7421.
Fig. 19. Lycoperdon subvelatum. No. 7263. Florida.
Fig. 20. Lycoperdon rimulatum. No. 715.

Figs. 1, 2, 11 X 1013; figs. 3-10, 12-20 X 1620.

ri.Aii: 11.'

PLATE 113

PLATE 113

Fig. 1. Lycoperdon umbrinum. No. 1004.

Fig. 2. Lycoperdon umbrinum. West Albany, N. Y.

Fig. 3. Lycoperdon umbrinum. Bres. Herb. (as£. hirlum).

Figs. 4 and 5. Lycoperdon Turneri. Type.

Fig. 6. Lycoperdon molle. Persoon Herb, (now at N. Y. B. G. Herb.).

Fig. 7. Lycoperdon acuminatum. No. 680.

Fig. 8. Lycoperdon pyriforme. No. 998.

Fig. 9. Lycoperdon subincamatum. No. 5754. Linville Falls, N. C.

Fig. 10. Lycoperdon subincamatum. No. 3171a.

Fig. 11. Lycoperdon muscorum. Type.

Fig. 12. Lycoperdon gemmatum. No. 3159.

Fig. 13. Lycoperdon Peckii. No. 5749. Linville Falls.

Fig. 14. Lycoperdon pedicellatum. No. 5668.

Fig. 15. Lycoperdon eximium. No. 1002.

Fig. 16. Lycoperdon fuscum. No. 5525. Blowing Rock.

Fig. 17. Lycoperdon marginatum. No. 482.

Fig. 18. Lycoperdon marginatum. No. 5225.

Fig. 19. Lycoperdon cahescens. Co-type (Wright, No. 6366).

Fig. 20. Lycoperdon Curtisii. No. 552.

Fig. 21. Lycoperdon Curtisii. Co-type.

Fig. 22. Lycoperdon Wri'Jitii. Co-type.

Fig. 23. Lycoperdon candid um. Persoon Herbarium.

Fig. 24. Lycoperdon oblongisporum. No. 497.

Fig. 25. Lycoperdon pusillum. No. 2209.

Fig. 26. Lycoperdon pusillum. No. 1003.

Fig. 27. Lycoperdon pusillum. No. 5390.

Fig. 28. Lycoperdon pusillum. No. 5211.

Fig. 29. Lycoperdon polymorphum. Pitchfork, Wyoming.

Figs. 1-17, 19-26, 29 X 1620; figs. 18, 28 X 1013; fig. 27 X 810.

PLATE 113

PLATE 114

Fig
Fig
Fig
Fig
Fig,
Fig.

Fig,
Fig
Fig
Fig
Fig
Fig

PLATE 114

1. Bovislellaradicata. No. 5919.

2. Bovistella radicata. No. 5200.

3. Bovistellaradicata. Torrend, No. 77.

4. Bovistellaradicata. No. 1545. South Carolina.

5. Bovistella cchinella. Abisco, Lapland.

6. Bovistella eckinella. Type.

Figs. 7 and 8. Bovisla plumbea. No. 5358.

9. Bovistapila. No. 7999.

10. Bovistapila. Co-type.

11. Bovistapila. No. 5359.

12. Bovisla nigrescens. Cavelante. Bresadola Herbarium.

13. Bovisla minor. No. 2416. Canada.

14. Bovistella dcalbata. Lloyd Herbarium.

Figs. 15-17. Myccnastrumcorium. Lloyd Herbarium.

Figs. 1, 3, 5, 6, 8, 10-14, 17 X 1620; figs. 2, 16 X 810; 4, 7 X 47; fig. 9 X 503; fig. 15 X 188.

PLATE 114

PLATE 115

PLATE 115

Fig. 1. Geaster triplex. No. 682 (lower spore drawn in 7% KOH).

Fig 2. Geaster limbatus, sense of Bresadola. Burlington, Wis.

Fig. 3. Georfer/imiaftw, sense of Hollos. England (?). N. Y.B.G. Herb. (Massee, coll.).

Fig. 4. Geaster Morganii. Ohio. N. Y. B. G. Herb. (Underwood, coll.) .

Fig. 5. Geaster saccatus, northern form. Asheville, N. C.

Fig. 6. Geaster saccatus. New Haven Gap, Jamaica.

Fig. 7. Geaster velutinus. No. 4095.

Fig. 8. Geaster Readeri. Type.

Fig. 9. Geaster radicans. Co-type (Ravenel, No. 953).

Fig. 10. Geaster mirabilis. No. 7077.

Fig. 11. Geaster papyraceus. Kiu Siu, Japan (Curtis Herb.).

Fig. 12. Geaster caespitosus. Ohio. N. Y. B. G. Herb. (Lloyd, coll.).

Fig. 13. Geaster trichifer. Morce's Gap, Jamaica.

Fig. 14. Geaster subkulosus. Jamaica (Earle, No. 184).

Fig. 15. Geaster mammosus. Canada.

Fig. 16. Geaster arenarius. Florida (Lloyd, coll.).

Fig. 17. Geaster mfescens. No. 985.

Fig. 18. Geaster fornlcatus. Smith Island, N. C.

Fig. 19. Geaster fimbrialus. No. 7097.

Fig. 20. Geaster floriformis. Colorado (Clements, No. 613.).

Fig. 21. Geaster coronatus. No. 75.

Fig. 22. Geaster coronatus. No. 684.

Fig. 23. Geaster coronatus. No. 38S1.

Fig. 24. Geaster minimus. Type.

Fig. 25. Geaster Cesati. No. 1634 (Cesati; Path. & Myc. Herb.).

Fig. 26. Geaster leptospermus. No. 2028.

Fig. 27. Geaster pcclinatits. Jamesville, N. Y.

Fig. 28. Geaster plicalus. France (Patouillard).

Fig. 29. Geaster biplicatus. Bonin Islands. Co-type.

Fig. 30. Geaster Bryanlii. England (?). N. Y. B. G. Herb. (Massee, coll.).

Fig. 31. Geaster Schmidelii. Calloway, Nebraska (Bartholomew, coll.) .

Fig. 32. Geaster Rabenhorslii. Type distribution.

Fig. 33. Geaster Hariotii. Brazil (Patouillard Herb.).

Fig. 34. Geaster asper. Lincoln, Neb.

Fig. 35. Geaster umbilicatus. Newfield,N. J.

Fig. 36. Geaster Drumnwndii. Brisbane, Australia.

Fig. 37. Aslraeus hygromelricus. No. 685.

All figs. X 1620.

PLATE 115

PLATE 116

PLATE 116

Fig. 1. Geaster limbatus, sense of Bresadola. Kansas. No. 8252.

Fig. 2. Geaster elegans. France (?). Tulasne Herbarium.

Fig. 3. Geaster radicans. Sarasota, Florida.

Fig. 4. Geaster caespitosus. Type.

Fig. 5. Geaster arenarius. Type.

Fig. 6. Geaster mammosus. North Dakota. No. 254.

Fig. 7. Geaster subiculosus. Type.

Fig. 8. Geaster lignicola. Type.

Fig. 9. Geaster Hieronymi. No. 244. Jalapa, Mexico.

Fig. 10. Geaster floriformis. North Dakota. Sydow, Fungi exot. exs. No. 152.

Fig. 11. Geaster calceus. Type.

Fig. 12. Geaster granulosus. Austria (Fuckel, coll.).

Fig. 13. Geaster argenteus. Type.

Fig. 14. Geaster Archeri. Type.

Fig. 15. Coilomyces Sehweinitzii. Co-type.

Fig. 16. Geaster terreus. Waste Lands, S. Dakota.

Fig. 17. Geaster argentalus. Type.

Fig. 18. Geaster coronatus, fornicate form. No. 6065.

Fig. 19. Geaster eoronatus, minimus form. Ymala, Mexico.

Fig. 20. Geaster involutus. Type.

All figs. X 1620.

PLATE 116

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7

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16

10

15

11

14

12

13

17

18

19

20

PLATE 117

PLATE 117

Fig. 1. Geaster saccatus. No. 7350.

Fig. 2. Geaster Lloydii. Florida.

Fig. 3. Myriostomacoliformis. Lloyd Herbarium.

Figs. 4-7. Astraeushygrometricus. No. 7441.

Figs. 8 and 9. DiplocyslisWrightii. Bahama Is.

Figs. 10-14. Sphaerobolusstellalus. No.5112. Fig. 13 shows a gemma from the gleba.

Figs. 1-3, 8, 14 X 1620; figs. 4-6 X 810; figs. 7, 9 X 503; figs. 10-13 X 1013.

0©

PLATE 117

§ m

PLATE 118

PLATE 118

Figs. 1 and 2. Disciseda Candida. No. 6092.

Fig. 3. Disciseda Candida. No. 7507 (showing section of trama with basidia).

Fig. 4. Disciseda Candida. No. 5969.

Fig. 5. Disciseda Candida. No. 7115.

Fig. 6. Disciseda Candida. Co-type.

Fig. 7. Bovista Candida. Type.

Fig. 8. Disciseda subterranea. North Island, S. C.

Fig. 9. Disciseda pedicellata. Hartsville, S. C.

Fig. 10. Disciseda pedicellata. No. 15 (Ravene\,as Bovista nigrescens).

Fig. 11. Arachnion album. No. 5209.

Figs. 12 and 13. Arachnion album. No. 5224.

Fig. 14. Arachnion album. No. 730.

Fig. 1, f nat. size; figs. 2, 4, 11, 12 X 1013; fig. 3 X 466; figs. 5-10, 14 X 1620; fig. 13 X 298.

PLATE 118

PLATE 119

PLATE 119

Fig. 1. Tylostoma m^mmosum. North Dakota.

Fig. 2. Tylostoma mammosum. No. 926 (Fungi bavarici).

Fig. 3. Tylostoma brumale. New York Botanical Garden (Schlesien).

Fig. 4. Tylostoma simulans. No. 7504.

Fig. 5. Tylostoma simulans. No. 87.

Fig. 6. Tylostoma fioridanum. Type.

Fig. 7. Tylostoma verrucosum. Ohio (Lloyd).

Figs. 8 and 9. Tylostoma Finkii. No. 6060.

Fig. 10. Tylostoma volvulatum. No. 142. Porto Rico.

Fig. 11. Tylostoma pocidatum. No. 18S9(E.&E., Fungi Columbiana).

Fig. 12. Tylostoma obesum. Co-type.

Fig. 13. Tylostoma granulosum. No. 424 (Saccardo, Myc. italica).

Fig. 14. Tylostoma campestre. No. 861 Ip. Canada.

Fig. IS. Tylostoma Berkeley ii. Folly Island, S.C.

Figs. 16 and 17. Quelelia mirabilis. Trexlertown, Pa. (Fig. 16showsabitof capillitium) .

Figs. 1-3, 5-8, 10-15, 17 X 1620; fig. 4 X 1013; fig. 9 X 278; fig. 16 X 503.

HATE 119

PLATE 120

PLATE 120

Fig. 1. Scleroderma geasier. No. 1761.

Fig. 2. Scleroderma flavidum. No. 660.

Fig. 3. Scleroderma flavidum. No. 1698 (E.& E.).

Fig. 4. Scleroderma bovista, sense of Hollos. No. 725.

Fig. 5. Scleroderma bovista, sense oiBvesa.do\3.. No. 5920.
Fig. 6. Scleroderma aurantium. No. 5622.
Fig. 7. Scleroderma cepa. No. 916.
Fig. 8. Scleroderma tcnerum. Type.
Fig. 9. Sclerodermalycoperdoides. No. 5201.

Fig. 10. Sclerodermalycoperdoides. No. 5253.

Fig. 11. Sclerodermalycoperdoides. Type.

Fig. 12. Scleroderma verrucosum. Bresadola Herbarium.

Figs. 13-17. Pisolithuslinclorius. No. 8170.

Figs. 18, 19. Pisolithus linctorius. No. 5402. One spore in fig. 9 shows appearance when first put into
water; the others after the gelatinous material has soaked off.

Figs. 1-9, 12 X 1013; figs. 11, 19 X 1620; figs. 10, 13-18 X 810.

PLATE 120

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PLATE 121

PLATE 121

Fig. 1. Cyathus slriatus. No. 5590.

Figs. 2-4. Cyathus slriatus. No. 5280.

Fig. 5. Cyathus slriatus. No. 2432.

Fig. 6. Cyathus Poeppigii. No. 7282.

Figs. 7-12. Cyathus stercoreus. No. 5916. Fig. 12 shows section of wall of young peridiole: lower

layer the tunic; middle, the sclerotic layer; upper, the hymenium with basidia and delicate

threads.
Fig. 13. Cyathus stercoreus. No. 39.
Fig. 14. Cyathus stercoreus. No. 5417.
Fig. 15. Cyathus rugispermus. No. 7458.

Figs. 1, 5, 6, 13-15 X 503; figs. 2-4 X 1013; figs. 7-11 X 810; fig. 12 X 278.

PLATE 121

PLATE 122

PLATE 122

Figs. 1-3. Cyathus melanospermus. No. 23.

Fig. 4. Cyathus melanospermus. Type.

Fig. 5. Cyathus vernicosus. No. 1688.

Fig. 6. Crucibulum vulgare. No. 7440.

Fig. 7. Crucibulum vulgare. No. 647.

Figs. 8, 9. Crucibulum vulgare. No. 5277.

Figs. 10, 11. Nidularia pulvinata. Asheville, N. C. Fig. 10 shows one of thejsclerotic cellsjrom the

subhymenial layer.
Fig. 12. Nidularia pulvinata. No. 7588.
Fig. 13. Nidularia castanea. Newfield, N. J.

Figs. 1, 2, 8, 9 X 1013; figs. 3-7, 10 X 503; 11-13 X 1620.

PLATE 122

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PLATE 123

PLATE 123

Figs. 1-3. Cyathus Lesueurii. No. 7281 (showing spores embedded among gelatinizing threads).
Fig. 4. Colostoma cinnabarina. No. 5521.
Fig. 5. Colostoma lutescens. No. 1935.
Fig. 6. Colostoma Ravenelii. No. 1488.

Figs. 1-3 X 503; others X 1620.

PLATE 123

INDEX

ma, 139
Agaricus tabularia, 63
Anthurus, 3, 8

aseroeformis, S

borvalis. 4. 8

boreali* van Klitzingii, 9

MuUerianus, 8
Ar.ichniaoar. 1. 144
Arachnion. 58, 144

album, 144, 146
Arachniopsis. 144
Arcangcliella. 23
Astraeus, 105. 139, 185

hygrometricus, 124, 185

stcllatus, 185

Battaiea, 149

Boletus parasiticus, 165
Bovista. 58, 94, 96, 97, 101, 139
ammophila. 95
brunnea, 100
Candida, 140
circumscissa, 139
gigantea, 62
lilacina, 63
minor, 95, 97, 99
montana, 97
nigrescens, 9S, 142
ovalispora, 99
pila, 97
plumbea, 90, 95, 97, 98, 93

radicata, 95
Bovistella. 58, 75. S6, 94, 100

ammophila, 95

Davisii, 96

dealbata, 100

echinella, 95, 96

ohiensis, 95

radicata, 75, 95, 14"

tomentosa, 100
Broomeia, 143

congregata. 145

guadaluper.sis. 143

Calostoma, 139, 185, 187, 188
cinnabarina, 189. 191
lutescens, 189, 190
microsporum, 189, 192
Ravenelii, 189, 191

Calostomataceae, 1 . 185
Calvatia, 1, 5S, 59, 60,90,94

arctica, 66

aurca, 60

borealis, 66

Candida, 61

caelata, 60, 61, 64, 68

craniformis, 60, 67, 68

cretacea, 6(1, 66

cyathiformis, 59, 60, 62, 63, 64

elata, 60, 65, 81, 82, 83

favosa, 68

fragilis, 60, 64

lepidophorum, 60

lilacina, 67

lilacina var. occidcntalis, 65

maxima, 59, 60, 61, 62

novae-zelandiae, 63

pachyderma, 60, 64, 65

polygonia, 63

rubro-flava, 1, 59, 60, 93

saccata, 65, 83
Carpobolus stellatus, 147
Catastoma circumscissum, 139

pedicellatum, 142
Cauloglossum transversarium, 56
Chlamydopus, 149, 150
Claustula Fischeri, 3
Clathreae, 3
Clathrella, 6
Clathrus, 5, 18

albus, 6

cancellatus, 4, 5, 6

columnatus, 4, 5, 6

flavescens, 6

ruber, 6

volvaceus, 6
Coilomyces Schweinitzii, 117
Colus, 6

Garciae, 7

hirudinosus, 6

javanicus, 7
Mulleri, 6

Schellenbergiae, 4, 7
Crucibulum, 40, 173, 174, 175, 181
crucibuliiorme, 181
vulgare, 175, 181
Cyathia hirsuta, 175

197

198

INDEX

Cyathus, 173, 174, 175

emodensis, 173

fimentarius, 182

hirsutus, 166

intermedius, 175

lcntifera, 180

Lesueurii, 175, 177, 179

Lesueurii var. minor, 178, 179

melanospermus, 175, 178, 179

olla, 180

Poeppigii, 175, 177

rufipes, 175

rugispermus, 175, 178

stercoreus, 173, 175, 177, 178, 179, 180, 181

striatus, 175

vernicosus, 175, 178, 180
Cycloderma ohiensis, 113

Dendrogaster connectens, 46
Dictyocephalos, 149
Dictyophora, 11, 13, 16

duplicata, 5, 13

phalloidea, 14

Ravenelii, 11
Diplocystis, 58

Wrightii, 143
Disciseda, 1, 58, 59, 97, 139

Candida, 139

circumscissa, 139, 141

pediceUata, 139, 141,142

subterranea, 139, 141, 142

verrucosa, 141

Elasmomyces, 53, 55

Gallacea scleroderma, 51

violacea, 51
Gautieria, 15, 20, 21, 23

graveolens, 22

morchelliformis, 22

Trabuti, 23
Geaster, 1, 59, 80, 102, 185

ambiguus, 132, 136, 138

Archeri, 110, 135

arenarius, 103, 105, 120

argentatus, 120, 138

argenteus, 119

asper, 105, 128, 135, 137

australis, 114

avellaneus, 138

badium, 137

biplicatus, 132, 133

Bryantii, 105, 129, 132, 133

caespitosus, 116

calceus, 130

campestris, 135

Cesatii, 129

columnatus, 138

corallinus, 119

coronatus, 103, 105, 115, 121, 125, 129, 132,

134, 136
Curtisii, 115
delicatus, 127

Drummondii, 103, 105, 120, 137
elegans, 134, 137
nbrillosum, 185
fenestratus, 125
fimbriatus, 105, 106, 111, 113, 121, 123, 124,

125
floriformis, 103, 105, 120, 121, 127
fornicatus, 102, 103, 105, 106, 115, 124, 129, 130
granulosus, 130
Hariotii, 103, 105, 135
Hieronymi, 105, 123, 124
hygrometricus, 166, 185
involutus, 137
juniperinus, 130

lageniformis, 106, 111, 112, 126, 128
leptospermus, 78, 105, 131, 136
lignicola, 116

limbatus, 104, 107, 122, 124, 127, 132
Lloydii, 105, 123
lugubris, 119

mammosus, 103, 105, 111, 119, 136, 137, 188
marchicus, 124
marginatus, 129
Michelianus, 106
minimus, 103, 127, 129
mirabilis, 105, 110, 116, 118
Morganii, 103, 104, 109
multifidum var. a, 132
papyraceus, 116, 117
pectinatus, 105, 108, 132, 133, 134, 135
plicatus, 133
pseudo-limbatus, 109
pseudomammosus, 135
quadrifidum, 129
Rabenhorstii, 134
radicans, 103, 104, 115, 124
Readeri, 113
rufescens, 103, 105, 106, 108, 109, 121, 122, 123,

125, 126
saccatus, 104, 106, 109, 110, 116, 126
saccatus, northern form, 111
saccatus var. Walkeri, 113
Schaefferi, 109

Schmidelii, 105, 132, 134, 136, 137
Schweinfurthii, 137
Smithii, 132, 136
stellatus, 185

INDEX

199

ttipitatus, 103

striatulus. 136

striatum 1(17. 134

btriatus var. minimus, 137

subieulosus. 104, 114, 116, 118

tentrfpes, 133, 135

teneus, 130

trnhifcr. 105. 116,118

triplex, 104, 106, 10S, 109, 111, 112, 114, 115,
126

tunicatus, 125

umbilicatus, 105, 136

velutinus, 104,112,113, 115, 124

Welwitschii, 115
Geastrum fibrillosum, 185

minimum, 137
Granularia, 183

. i<tanea, 184

pisiformis, 183

pulvinata, 183
Gymnomyces, 15, 21, 23

vesiculosus, 23
Gyrophragmium decipiens, 55

Delilei, 55
Gyropodium coccineum, 189

Holocotylon, 144
Hydnangium, 25, 47

cameum, 48

galathejum, 49

hysterangioides, 25

liospermum, 25

monosporum, 51

Ravenelii, 48

Stephensii, 49

Stephensii var. Ravenelii, 48
Hymenogaster, 24, 45, 46

anomalus, 44, 45

decorus, 45, 46, 47

foetidus, 45

Thwaitesii, 45, 46, 47

utriculatus, 46
Hymenogastraceae, 1, 24, 26, 173
Hypoblema, 60
Hysterangiaceae, 1, 3, 15, 26
Hysterangium, 15, 17, 21, 25, 36, 46, 51

calcareum, 18

clathroides, 15, 17, 19, 20, 21, 26

coriaceum, 18

fragile, 18

membranaceum, 18

nephriticum, 18

pompholyx, 17, 18, 19

rubricatum, 18

stoloniferum, 18, 21

Btoloniferum var. americanum, 17, 19, 20

stoloniferum var. mutabilc, 21
ThwaiUsii, 18

Ithyphallus, 11

imperialis, 12
impudicus, 5, 12, 13
Ravenelii, 5, 11, 12
rubicundus, 5, 13

Jaczewskia, 16

Key to the Families, 1

Key to the Phalloids of the United States, 4

Laternea columnata, 5
Leucogaster, 24, 42, 45

anomalus, 42, 44

araneosus, 42, 43, 44

badius, 42, 44

carolinianus, 42, 43

fulvimaculosus, 42, 44

luteomaculatus, 42
Leucophleps, 51

magnata, 51
Lycoperdaceae, 1, 58

Lycoperdon, 1, 58, 59, 69, 84, 86, 93, 94, 139, 144,
145

abscissum, 89

acuminatum, 70, 78, 132

atropurpureum, 70, 71

atropurpureum var. stellare, 72, 74, 75

Berkeleyi, 67

bovista, 62

caelatum, 68, 166

calvescens, 87

candidum, 90

calyptriforme, 78

caudatum, 85

cepaeforme, 61, 75, 92

coloratum, 71, 93

compactum, 74

constellatum, 73, 85

cretaceum, 66

cruciatum, 87

cupricum, 87

cupricum var., 76

Curtisii, 71,89

cyathiforme, 63

delicatum, 62, 67, 68

depressum, 69

dryinum, 87

echinatum, 70, 72, 73, 85

elatum, 65

elegans, 77

200

INDEX

excipuliforme, 82, 166

eximium, 71, 85, 86

favosum, 68

Fontanesii, 68

Frostii, 72

furfuraceum, 93

fuscum, 71, 85, 87

giganteum, 62

gemmatum, 65, 71, 82, 85, 88, 89, 166

glabellum, 76

hirtum, 76

lepidophorum, 60

leprosum, 78, 113

macrogemmatum, 82

marginatum, 66, 71, 73, 87, 90

Missouriense, 67

molle, 77, 78, 81, 82

muscorum, 65, 70, 71,81, 102

nigrescens, 83, 85

oblongisporum, 71, 91, 92

papillatum, 87

Peckii, 71, 84

pedicellatum, 71, 85, 89

perlatum, 83, 84

polymorphum, 61, 69, 71, 92, 94

polytrichum, 81

pratense, 88, 89

pseudoradicans, 70, SI

pulcherrimum, 70, 72, 74, 85

pusillum, 69, 71, 90, 91, 93, 94

pusio, 119

pyriforme, 70, 71, 79, 83

recolligens, 188

rimulatum, 70, 75, 76

saccatum, 65

sculptum, 70

separans, 87

Sinclairii, 68

subincarnatum, 71, 80

subvelatum, 70, 76

Turneri, 77

umbrinum, 70, 71, 76

Warnei, 55

Wrightii, 89
Lysurus, 8

australiensis, 9

Clarazianus, 9

Gardneri, 9

Mokusin, 8

Texensis, 8

Macowanites, 23, 53
Marasmius oreades, 63
Melanogaster, 24, 35, 36, 38, 41

ambiguus, 38, 41

mollis, 38

nauseosus, 38, 39

odoratissimus, 40

rubescens, 38, 40

variegatus, 38, 40, 41
Mitremyces, 189

lutescens, 191

Tylerii, 192
Mutinus, 9

bovinus, 10

caninus, 4, 10

Curtisii, 4, 10

elegans, 10

Ravenelii, 4, 9, 10
Mycenastrum, 58, 97, 101

corium, 101

corium var. Kara-Kumianum, 101

radicatum, 101

spinulosum, 101
Myriostoma, 1, 58, 59, 138

coliformis, 138

Nidula, 173, 174
Nidularia, 174, 183

alabamensis, 183

castanea, 183, 184

fascicularia, 180

globosa, 166, 183

Juglandicola, 181

pisiformis, 183

pulvinata, 183

stercorea, 177
Nidulariaceae, 1, 173
Nigropogon, 24, 37

asterosporus, 37

Octaviania, 24, 25, 47, 173
asterosperma, 25, 48, 51
compacta, 25, 26
purpurea, 38, 48, 50
Ravenelii, 48
Stephensii var. Ravenelii, 48

Peziza striatus, 176
Phallaceae, 1, 3
Phalleae, 3
Phallogaster, 3, 15, 16, 20

saccatus, 16

Whitei, 17
Phallus aurantiacus, 13

imperialis, 13

impudicus, 11

indusiatus, 14

Ravenelii, 11

rubicundus, 11

togatus, 14

l\!>|-\

201

rhcllorin.i, loll
Phlyctogpoia hisca, 160
PisoKthus, 114, loO, 170

areaarius, 170

tinctorius, 170
Podaxaceae, loO
Podaxon, 56, 160
Polj saccum, 1 70

i rassipes, 1 70

pisocaipium, 170
Protophallus, 3, 15

jamaiccnsis. 16, 21
Protubcra. 3, 1<)
Pseudocolus, 6

Quclctia. 140, 158
mirabi'is, 158

Rhacophyllus, 53
Rhizopogon, 24, 26, 33, 37, 38, 41
aestivus, 41

albus, 29

atlanticus. 27. 2l>, 35, 36
diplophloeus, 41
graveolens, 27, 30
lutcolus, 27, 28, 29, 33, 35
maculatus, 27, 35
nigrescens, 27, 29, 30, 35
pachyphloeus, 41
parasiticus, 26. 27
piceus, 27, 31, 34, 35
provincialis, 31
roscolus, 26, 27, 29,31,32
albescens, 27, 28, 31, 32, 3i, 34, 41, 4S
rubescens var. Vittadini, 29
truncatus, 27, 36
Rhopalogaster, 53, 56
transversarium, 56

Sclerangium polyrhizon, 161
Scleroderma, 38, 101, 160, 170

aurantium, 66, 161, 165

bovista, 161, 163, 164, 167

cepa, 161, 163, 165, 167, 169

flavidum, 161, 162, 165, 167

fuscum, 160

geaster, 161, 163

lycoperdoides, 160, 161, 165, 168

lycopcrdoides var. reticulatum, 161, 170

pteridis, 160, 162

tenerum, 168, 169

Texense, 164

verrucosum, 161, 168, 169

vulgare, 165
Sclerodermataccae, 1, 160
Sclerogaster, 24. 25

compactus, 26

lanatus, 26

minor, 25

lai eae, 1, 53
tiuni, 53
acuminatum, 54, 55
agaricoides, 54

arizonicum, 55

australe, 55

decipiens, 55

krjukowinsi , 53

rabigenum, 54, 55

russuloides, 55

tenuipes, 55

Warnei, 54, 55
Simblum, 5, 7

rubescens, 7

rubescens var. Kansensis, 7

sphaerocephalum, 4, 7

texense, 4, 7
Sphaeria herculea, 56
Sphaericeps, 149
Sphaerobolaceae, 1, 146
Sphaerobolus, 146

carpobolus, 147

impaticus, 146

iowensis, 146, 148

stellatus, 146, 147, 148

stellatus var. giganteus, 146, 148
Stella americana, 161

Trichastcr melanocephalus, 102
Tulasnodea, 150

fimbriata, 158
Tulostoma, 150
Tylostoma, 149, 158, 174, 185, 187

albicans, 154

americanum, 154

Berkeleyii, 150, 157

brumale, 151

campestre, 150, 156, 157

fimbriatum, 156

Finkii, 150, 154

floridanum, 150, 151, 153

gracile, 154

granulosum, 155, 156, 157

kansense, 154

Lloydii, 150, 154, 156

Longii, 154

mammosum, 150, 151, 152, 153, 158

minutum, 154

obesum, 155

pedunculatum, 151, 152

poculatum, 150, 155, 156

punctatum, 157

pygmaeum, 153

simulans, 150, 161, 153

squammosum, 154, 158

verrucosum, 150, 163

volvulatum, 150, 166
Tylostomataceae, 1, 149

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