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A Continuation of the 



The Library of the 

JAN 1 8 1979 





1. Two New Species of Palms from Nicaragua. By S. F. Glassman ... 1 

2. Tropical American Plants, VI. By Louis 0. Williams 11 

3. Agriculture, Tehuacan Valley. By C. Earle Smith, Jr 49 

4. Flora, Tehuacan Valley. By C. Earle Smith, Jr 101 

5. Preliminary Studies in the Palm Genus Syagrus Mart, and Its Allies. 

By S. F. Glassman 145 

6. Tropical American Plants, VII. By Louis 0. Williams 165 

7. Supplement to Orchids of Guatemala. By Donovan S. Correll .... 175 

8. Preliminary Notes on Scrophulariaceae of Peru. By Gabriel Edwin . . 223 

9. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 233 

10. Tropical American Plants, VIII. By Louis 0. Williams 247 

11. Notes on the Flora of Costa Rica, I. By William C. Burger 273 

12. A New Eurystyles from Nicaragua. By Alfonso H. Heller 279 

13. New Species in the Palm Genus Syagrus Mart. By S. F. Glassman . . 285 

14. A Revision of the Family Geastraceae. By Patricio Ponce de Leon . . 303 

15. Studies in American Plants. By Dorothy N. Gibson 353 

16. Two New Nicaraguan Juglandaceae. By Antonio Molino R 357 

17. Studies in the Palm Genus Syagrus Mart. By S. F. Glassman .... 363 

18. Tropical American Plants, IX. By Louis O. Williams 401 









VOLUME 31, NUMBERS 11, 12, 13, 14 


FEBRUARY 22, 1968 

1 1 ii -11 mr>n 



Assistant Curator, Cryptogamic Herbarium 



Published by 

FEBRUARY 22, 1968 


Library of Congress Catalog Card Number: 67-31602 


A Revision of the Family Geasteraceae 


I wish to acknowledge my indebtedness to Dr. Clark Rogerson of 
the New York Botanical Garden, Dr. Chester R. Benjamin of the 
National Fungus Collections, Dr. I. Mackenzie Lamb of the Farlow 
Herbarium, Harvard University, and Dr. Alexander H. Smith of the 
University of Michigan for making available to me on loan the neces- 
sary herbarium material, and for various kindnesses extended during 
my visits to study these collections. I am especially grateful to 
Dr. Alexander H. Smith who brought to my attention the partially 
completed manuscript and notes of Dr. Sanford M. Zeller. Gratitude 
is also expressed to the staff members of the Library and the Depart- 
ment of Botany of Field Museum of Natural History, who have 
helped me in numerous ways. 


The history of this family is intimately related to the history of 
its principle genus, Geastrum Pers. which was based on the Geaster 
of Micheli. Although Micheli placed only five species in Geaster, 
Persoon recognized six in his Geastrum. In 1842 Corda established 
his family Geastrideae, placing in it the genera Geaster Mich., Pleco- 
stoma Desv., and Myriostoma Desv. He included the species of 
Geaster of Micheli and Fries in the genus Plecostoma, and only Geaster 
hygrometricus in the genus Geaster. In the genus Myriostoma he 
placed M. coliformis, where it remains today. Subsequent authors 
included the genera Geaster and Myriostoma in the family Lycoper- 
daceae and reduced Plecostoma to a synonym of Geaster. 

In 1889 Morgan proposed the name Astraeus hygrometricus for 
the old Geaster hygrometricus and placed it in his order Lycoper- 
daceae. Astraeus hygrometricus is accepted today by all modern 
authors except Lloyd and Cunningham, who replaced it in Geaster. 
Coker and Couch, and, later, Fischer placed it in the family Calosto- 
mataceae. In 1936 Martin created the currently accepted family 
Astraeaceae for this genus. 



In 1933 Fischer established the family Geastraceae, including in 
it the genera Geastrum Pers., Myriostoma Desv., Geasteropsis Hollos, 
and Trichaster Czrn. In 1913 Long proposed the genus Geasteroides 
for a plant collected by him in Texas, which has a sterile base in 
the endoperidium. In this paper, Geasteropsis is placed in synon- 
ymy with Trichaster and Trichaster I consider to be a subgenus of 

GEASTRACEAE Fischer, Engl. & Prantl Nat. Pflanzenf . Aufl. 2(7a) : 
72.1933. GEASTRIDEAE Corda, Icones Fung. 5:25. 1842. GEASTRAE 
Lloyd, The Geastrae in Bull. Lloyd Lib. 5(2). 1902. GEASTREAE 
G. H. Cunningham, Proc. Linn. NSW. 52: 251. 1927. 

Basidiocarp epigeous or hypogeous, globose to acuminate; exoperidium of three 
well-defined layers, splitting at maturity in stellate lobes; endoperidium pedicellate 
or sessile, membranous, glabrous or variously roughened, opening by one or several 
mouths (when one, it may or may not have a peristome) ; capillitium threads simple 
or with short branches at the ends, long, tapering, with septa in the genus Geas- 
teroides; spores globose to subglobose, verrucose or spinose, 2 to 5jt in diameter. 


Endoperidium with a prominent sterile base Geasteroides. 

Endoperidium without a sterile base. 

Membrane of the endoperidium always persistent, with several mouths and 
several pedicels Myriostoma. 

Membrane of the endoperidium persistent or not, with only one mouth and one 
pedicel Geastrum. 


The principal characteristics of this genus are: the prominent 
corky, sterile base, the fragile, deciduous endoperidium, and the 
branched and septate capillitium. 

Long published the name Geasteroides in 1917. Later, in 1945, he 
replaced this name with Terrostella, saying that "The generic name 
Geasteroides is untenable since it is already preoccupied by Battarra's 
genus (1755) of the same name. ..." Since the starting date for 
Gasteromycetes is 1801, the use before starting date does not pro- 
hibit using it in 1917; consequently Terrostella is an illegitimate name, 
being a substitution for a valid one. 

Geasteroides Long, Mycologia 9: 271. 1917. Terrostella Long, 
Mycologia 37: 605. 1945. 

Peridium double; exoperidium splitting into stellate, reflexed, persistent seg- 
ments; endoperidium fragile, upper portion more or less deciduous, lower part 
persistent, consisting of a prominent sterile base; mouth indefinite, single; colu- 


mella present; capillitium branched and septate; spores globose, verrucose, some- 
times uni-guttulate. 

Type: Geasteroides texensis Long. 

Discussion: The presence of a sterile base in the endoperidium 
and the septate capillitium are characters of enough importance to 
establish a genus in this family, as Long did. 

This genus consists of two species: G. texensis Long and G. barbata 
(Dissing & Lange) P. Ponce. 


Endoperidium stipitate; columella deciduous G. texensis. 

Endoperidium not stipitate; columella persistent G. barbata. 

Geasteroides texensis Long, Mycologia 9: 271. 1917. Terro- 
stella texensis (Long) Long, Mycologia 37: 605. 1945. Geasteropsis 
texensis (Long) Fischer in Engler & Prantl, Nat. Pflanzenf. Aufl. 
2(7a):75. 1933. 

Unexpanded basidiocarp unknown, apparently hypogeous, becoming super- 
ficial and expanded at maturity, then 4-10 cm. diameter; exoperidium revolute, 
thick, rigid, coriaceous, subhygroscopic, splitting to about the middle into 7-10 
segments, concave below, convex above, rays unequal, recurved with involute tips; 
outer layer of arachnoid mycelium and dirt that peels off as the plant ages; fleshy 
layer adnate, dark brown, fissured and cracked when dry; endoperidium short 
stipitate, subglobose, drab gray, 15-25 mm. broad, very fragile, apparently with a 
very poorly defined mouth, upper part slowly dehiscing down to the sterile base, 
leaving it crowned with a subglobose columella and spores; sterile base corky, com- 
pact, wood brown to fawn color, 10-15 mm. across by 8-10 mm. tall; stipe terete 
to strongly flattened, stout, subligneous, 2-3 mm. thick by 3-15 mm. wide by 2 mm. 
high; gleba chestnut brown, in very old plants entirely disappearing and leaving 

FIG. 1. Gasteroides texensis Long. Long 
8787. Texas. Lloyd Myc. Coll. Smithsonian 

only the sterile base seated on the stipe; columella soft, weak, early deciduous; 
capillitium wine colored to light brown, threads very long, distantly branched, 
7-10/x thick, tapering, septate in thicker parts, breaking up into segments 800- 
l.OOOyu long, walls smooth, often appearing as if filled with minute pits, lumen 
very small or none; spores globose, 1-guttulate, 3-5^ in diameter, brown, faintly 


Habitat: Solitary or in small groups in rich, loose, sandy loam 
around bases of old rotting post oak stumps in open post oak woods. 
Distribution: Texas. 

Geasteroides barbata (Dissing & Lange) P. Ponce comb. nov. 
Geastrum barbatum Dissing & Lange, Cast, of Congo, Bull. Jard. Bot. 
Bruxelles32:382. 1962. 

Unexpanded basidiocarp hypogaeic, globose, 2-2.5 cm. broad, attenuated be- 
low in a short, stemlike base, with no debris incrusted; exoperidium, when open, 
7-8 cm. broad, split into 6-8 unequal, broad, bluntish-rounded rays; outer layer 
brown to fawn, squamose, rather thick, tending to peel off in flakes; medium layer 
greyish brown, hard, strongly wrinkled; inner layer first pale buff, thick; when 
dried, greyish-brown, persistent; endoperidium lacking; columella prominent, soft- 
leathery, roundish depressed, 2-5 cm. broad, 1 cm. high, with stiff radiating white 
brittle hairs, the columella attached to a compact, sterile base, 2.5 cm. high to 
1.5 cm. broad; spores 4.2-4.8/i broad, brown with dense, short warts; capillitium 
composed of 3.6-5.2/i broad, hyaline to yellowish tinted, thin-walled, septate, 
branching threads, and attached to the columella and the inner layer of the 

Habitat: Clayey sand of termitaries or attached to mud walls of 
cabins of the natives. 

Distribution: Congo. 

Discussion : Dissing and Lange mentioned the possible close rela- 
tionship of their Geastrum barbatum to the genera Geasteropsis Hollos 
(this an outright synonym of Geastrum) and Gasteroides Long. I con- 
sider that this species belongs to the genus Geasteroides Long, because 
of its sterile base and the branched septate capillitium. 


The genus Myriostoma includes only one species, M. coliforme, 
first mentioned by Doody in the appendix to Ray's Syn. 2nd Ed. 
1696, and illustrated by Dikson in Fasc. Plant. Crypt. Britanniae, 
who called it Lycoperdon coliforme. Persoon in Syn. Meth. Fung. 
1801, referred it to Geastrum, and in 1809, Desveaux proposed the 
genus Myriostoma for this species, which is accepted today. 

Myriostoma Desv. Obs. Gen. dans Champ. Journ. Bot. Paris 2: 
38. 1809. 

Basidiocarp subglobose; exoperidium coriaceous, splitting in a stellate form 
with segments expanded or reflexed; outer layer with adherent sand or debris; endo- 
peridium subglobose, compressed, with several short pedicels, membranous, papery, 
with several or many mouths, without peristome; gleba brown, with several colu- 
mellas; spores verrucose; capillitium simple. 


Only one species. 

Type: Myriostoma coliforme (Pers.) Corda. 

Myriostoma coliforme (Pers.) Corda, Anleitung zum Studium 
der Mycologie f. 16-17. 1842. M. anglicum Desv. Journ. Bot. Paris 
2: 104. 1809. Geastrum coliforme Pers. Syn. Meth. Fung. 131. 1801. 
Geaster coliforme Fr. Syst. Myc. 2: 12. 1829. G. columnatus Lev. 
Champ. Mus. Paris 161. 1846. 

Unexpanded basidiocarp subglobose, subterranean, becoming superficial when 
expanded, then 1.5-10 cm. wide, reflexed; exoperidium nearly smooth, splitting in 
stellate form in 5-7 lobes; outer layer with adherent sand or other particles; endo- 
peridium subglobose to compressed, with many pedicels, membrane minutely 
roughened, silver brown, with several to many small mouths; gleba brown, with 
several slender columellas; spores spherical, strongly verrucose, 4-5 M diameter; 
capillitium threads long, slender, tapering, 2-5^ thick, with thick walls. 

Habitat: Soil in woods, sandy soils. 

Distribution: Europe, North America, South America, Asia, and 
South Africa. 


The genus Geastrum Persoon is presumed to be based on Geaster 
Micheli. In it, Persoon included six species of Gasteromycetes in 
which the outer peridium opens in a stellate form. Two of these spe- 
cies today are known to belong to other genera. Geastrum coliforme 
is Myriostoma coliformis, and Geastrum hygrometricum is Astraeus 
hygrometricum. The four remaining species are basic, and I select 
G. coronatum Pers., from among the species originally given by Per- 
soon, as lectotype of the genus Geastrum. Clements and Shear would 
seem to have selected G. pectinatus for a genus which they call 
"Geaster (Michel.) Fr.", a name never published by Fries. Further- 
more, Micheli never knew a plant which Fries called G. pectinatus 
so the genus, even if valid (which it is not), could not be based on it. 

Geastrum is the principal and largest genus in the family Geastra- 
ceae to which it lends its name. In this family are two additional 
genera, Myriostoma Desv. and Geasteroides Long. The first has many 
mouths and many stipes in the endoperidium, and the second, which 
loses the endoperidium before maturity, has a sterile cup under the 
gleba and the capillitium is branched and septate. 

Regarding the division of the genus and the distribution of the 
species, the concept of Stanek is followed for species of the temperate 
zone; that of Dissing and Lange for the tropical species, with some 


Two subgenera are recognized in Geastrum, one based on the dis- 
appearance of the endoperidial membrane, and the second on the 
character of the stromatic mycelia and the epigeic habit. In order 
to separate and properly classify the species in these two subgenera, 
we must consider the mycelial layer, the mouth, the hygroscopic 
quality of the exoperidium, the texture of the endoperidial mem- 
brane, the surface of the spores, the form and ramifications of the 
capillitium, and the diameter of the spores as related to that of the 
capillitium threads. 

Geastrum Persoon, Synopsis Methodica Fungorum. 131. 1801. 
Plecostoma Desvaux. Journ. Bot. 2: 99. 1809. Geaster Micheli ex 
Fries, Syst. Myc. 3: 8. 1832. Cycloderma Klotzsch. Linnaea 7: 203. 
1832, in part. Coilomyces Berk. & Curt. Journ. Acad. Nat. Sci. 
Philadelphia (series 2) 2: 279. 1853. 

Basidiocarp in the button stage subglobose, with or without an apical point, 
hypogeous until dehiscence, or epigeous on soil, wood or organic duff; peridia 
double, the exoperidium splitting from the apex at maturity into rays which ex- 
pand more or less stellately or merely recurve, typically of three layers, the outer 
one delicate, of interwoven mycelial filaments or tomentum that may be scarcely 
discernible, sometimes a more or less perfect membranous film which remains at- 
tached to the middle layer or becomes more or less separated, sometimes separating 
completely and remaining as an empty sac or cup in the substratum when the endo- 
peridium is lifted by the arching of the next layer; the middle fibrous layer is more 
definite, thin, usually tough, of closely woven, homogeneous hyphae; the inner 
layer, splitting with the middle layer, is firm, fleshy at first, shrinking when dry, 
continuous or rimose, adnate to or peeling from the fibrous layer; endoperidium 
or spore sac thin, tough, membranous, pliable, stalked or sessile, opening by a 
single apical mouth that may be simply a smooth, torn hole or an elevated pore, 
sometimes surrounded by a circular, elevated or plane area (peristome) that is 
fimbriate, sulcate, or indeterminate; gleba composed of interwoven fibers that may 
be more or less laminate, or tubularly arranged, and which radiate from the colu- 
ini '11 ;i to the endoperidium; the tubular cavities are lined by the basidia, which are 
thick, elliptic or pyriform, each with 4-8 spores; columella usually of soft texture, 
clavate to spherical; capillitium abundant, simple or rarely branched; spores spher- 
ical, usually rough, warted or asperulate. 

Lectotype: Geastrum coronatum Pers. 

Distribution: Cosmopolitan, except not known in Antarctica; 
abundant in the temperate zones and in the tropics. 


Subgenus Geastrum 

Sect. Geastrum (Perimyceliata Stanek) 

Mycelium covering all the surface of the basidiocarp. 
Type: Geastrum coronatum Pers. 
Subsect. Geastrum (Glabrostomata Stanek) 


Peristome smooth, fibrillose. 
Type: Geastrum coronatum Pers. 

Group Coronatum. Peristome determinate, surrounded by 

a groove. 
G. coronalum 
Group Fimbriatum. Peristome indeterminate; basidiocarp 

not hygroscopic. 

G. fimbriatum, G. rufescens, G. hieronymii 
Group Floriforme. Peristome indeterminate; basidiocarp 

G. floriforme 
Group Minimum. Peristome determinate; basidiocarp not 

hygroscopic, fornicate or not. 
G. minimum 
Group Quadrifidum. Peristome indeterminate; basidiocarp 

not hygroscopic, fornicate. 
G. quadrifidum 
Subsect. Sulcostomata Stanek 

Peristome plicate-sulcate. 
Type: Geaslrum pectinalum Pers. 

Group Pectinatum. Endoperidium surface smooth, farinose 

or furfuraceus; basidiocarp hygroscopic or not. 
G. pectinatum, G. xerophilum, G. furfuraceum 
Group Campestre. Endoperidiujm surface granulose or ru- 
gose; basidiocarp hygroscopic. 
G. campestre, G. badium, G. ambiguum 
Group Lloydianum. Endoperidium surface granulose; basid- 
iocarp sub-hygroscopic. 
G. lloydianum 
Sect. Basimyceliatum Stanek 

Mycelium attached only at the base of the basidiocarp; outer layer not 


Type: Geastrum indicum (Klotz.) Rauschert 
Subsect. Basimyceliatum (Laevistomata Stanek) 

Peristome smooth, fibrillose, not outlined by a depressed border; 

basidiocarp not hygroscopic. 
Type: Geastrum indicum (Klotz.) Rauschert 

Group Indicum. Peristome orbicular, determinate; basidio- 
carp not fornicate, not hygroscopic or sub-hygroscopic. 
G. indicum, G. saccatum 

Group Smardae. Peristome determinate; basidiocarp pseu- 
dofornicate; mycelial layer nidiform, sometimes sepa- 
rate from the fibrous layer. 
G. smardae 
Subsect. Marginatum 

Peristomium sericeum, margine canaliculatis; fructificatio hygro- 


Peristome silky, outlined by a depressed border; basidiocarp hygro- 

Type: Geastrum recolligens (Sow.) Desv. 
Group Recolligens. 

G. recolligens 
Subgenus Myceliostroma Henn. 

Mycelial layer entire, persistent, not encrusted, with prominent basal mycelial 

tuft or subiculum. 

Type: Geastrum schweinitzii (Berk. & Curt.) Zeller 
Sect. Myceliostroma 


Epigeic or xylophilous; mycelial layer of fascicled threads, radiating from 
the fibrillose layer with subiculum or branched rhizomorphic system. 
More than one basidiocarp on a common mycelial stroma. 
Type: Geastrum schweinitzii (Berk. & Curt.) Zeller 

Group Schweinitzii. Xylophilous; many basidiocarps on one 


G. schweinitzii, G. schweinitzii var. stipitatum 
Sect. Subepigaea. sect. nov. 

Fructificationes juvenes hypogaeae; mycelium rhizomorphicum. 

Fructificationes unicae per mycelium. 

Basidiocarp hypogeic when very young, with prominent basal mycelial 

tuft. Generally one basidiocarp on each mycelium. 
Type: Geast rum javanicum (Lev.) P. Ponce 

Group Javanicum. Outer peridium splitting into two fibrous 


G. javanicum, G. javanicum var. welwitschii 
Subgenus Trichaster (Czern.) P. Ponce 

Endoperidium fugax, laceratum; gleba nuda. 
Endoperidium fugaceous; gleba naked. 
Type: Geastrum melanocephalum (Czern.) Stanek 
Sect. Trichaster. sect. nov. 

Fructificationes prima subterranea, demum epigaea. 
Basidiocarp hypogeic when young; becoming epigeal. 
Type: Geastrum melanocephalum (Czern.) Stanek 
Group Melanocephalum. 

G. melanocephalum, G. conrathii 
Sect. Lignicola. sect. nov. 

Fructification epigaeae, lignicola (ad ligna putrida). 
Basidiocarp epigeous, saprophytic on rotting wood. 
Type: Geastrum xylogenum Long & Stouffer 
Group Xylogenum. 
G. xylogenum 


Gleba exposed, not enclosed by the endoperidium [subgenus Trichaster (Czern.) 

P. Ponce] 
Columella prominent, sub-ligneous 

Fleshy layer of the exoperidium continuous G. melanocephalum. 

Fleshy layer of the exoperidium not continuous, transversely fissured. 

G. conrathii. 

Columella not evident G. xylogenum. 

Gleba enclosed by the endoperidium [subgenera Geastrum and Myceliostroma}. 
Mycelium subiculose, widely spread on the substratum [subgenus Myceliostroma 

Outer layer of the exoperidium splitting into 2 fibrous persistent layers. 

Exoperidium not fornicate G. javanicum. 

Exoperidium fornicate G. javanicum var. welwitschii. 

Outer layer of the exoperidium not splitting into 2 fibrous layers, not persistent. 

Basidiocarps not stipitate G. schweinitzii. 

Basidiocarps stipitate G. schweinitzii var. stipitatum 

Mycelium not subiculose, not widely spread on the substratum [subgenus 


Mycelium covering all the surface of the basidiocarp. 
Peristome indeterminate. 


Basidiocarps hygroscopic . G. floriforme. 

Basidiocarps not hygroscopic 

Exoperidium fornicate G. quadrifidum. 

Exoperidium not fornicate 

Endoperidium with setose hairs G. hieronymii. 

Endoperidium glabrous or subglabrous 

Reddish tinct on the exoperidium G. rufescens. 

No reddish tinct; exoperidium cream-colored 

Spores brown G. fimbriatum. 

Spores hyaline G. fimbriatum f . pallidum. 

Peristome determinate. 
Peristome sulcate. 

Endoperidium surface furfuraceous or granulose 
Endoperidium furfuraceous 

Basidiocarps hygroscopic G. furfuraceum. 

Basidiocarps not hygroscopic G. xerophilum. 

Endoperidium granulose. 
Basidiocarps hygroscopic. 

Endoperidium sessile, surface minutely granulose . . . G. badium. 
Endoperidium shortly pedicellate or sessile, surface with gran- 
ules well defined. 

Gleba umber, columella present G. campestre. 

Gleba ferruginous, columella not evident G. ambiguum. 

Basidiocarps not hygroscopic G. lloydianum. 

Endoperidium surface neither furfuraceous nor granulose. 

G. pectinalum. 
Peristome not sulcate. 

Peristome surrounded by a groove; basidiocarps pseudofornicate. 

G. coronatum. 
Peristome not surrounded by a groove; basidiocarps fornicate or not. 

G. minimum. 
Mycelium attached only at the base of the basidiocarp. 

Basidiocarps hygroscopic G. recolligens. 

Basidiocarps not hygroscopic. 

Basidiocarps pseudofornicate G. smardae. 

Basidiocarps not pseudofornicate. 

Basidiocarp always saccate, 1-2 cm. wide when open; fleshy layer not 
persisting as a collar around the base of the endoperidium. 

G. saccatum. 

Basidiocarp saccate or not, 4-5 cm. wide when open; fleshy layer fre- 
quently persisting as a collar around the base of the endoperidium. 

G. indicum. 

GEASTRUM subgenus Trichaster (Czern.) P. Ponce, comb. nov. 
Trichaster Czern. Bull. Soc. Nat. Moscow 18(2) : 149. 1845. Geaster- 
opsis Hollos, Novengt. Koslem 2: 72. 1903. 

This subgenus is based on Czerniaiev's genus Trichaster, made by 
him because he had observed that G. melanocephalum differed from 
the other Geastrums in the early loss of the internal peridium, leaving 
the gleba naked. However, it does not deserve generic status as the 
spores, capillitium, mycelia, and structure of the exoperidium are 
characteristic of Geastrum. 


Type species: Geastrum melanocephalum (Czern.) Stanek. 

This subgenus includes the following species: 
Geastrum melanocephalum (Czern.) Stanek. Europe and Africa 
Geastrum conrathii (Hollos) P. Ponce. Africa 

Geastrum xylogenum (Long) P. Ponce. North America 

Geastrum melanocephalum (Czern.) Stanek, Ces. Myc. 10: 
22. 1956. Trichaster melanocephalum Czern. Bull. Soc. Nat. Moscow 
18 (2): 150. 1845. Geaster fornicatus Hollos, Gast. Ung. 70. 1904. 

Unexpanded basidiocarp ovate, hypogeous until maturity, then 5-8 cm. tall 
and 3-8 cm. wide; exoperidium hard, rigid, coriaceous, splitting beyond the middle 
into 5-8 unequal, subhygroscopic rays which bend strongly backward and down- 
ward (not fornicate) ; fleshy layer 1-2 mm. thick, mummy brown, adnate, continu- 
ous; outer layer naked, smooth, dark brown, free of debris; base broad, concave 
with a cord-like rhizomorph in center; endoperidium sessile, apparently globose 
before dehiscence, a few fragments left at base of gleba; gleba naked, subglobose, 
with a round, thick, subligneous stipe 1.5 cm. broad; columella prominent, per- 
sistent, hard, subligneous, covered with a matted mass of capillitium and spores; 
spores 4.2-5.2 M in diameter, dark brown, globose, verrucose; capillitium 4.5-7/t 
thick, thicker than the spores, walls thin, unbranched, brown violet. 

Type locality: Ukraine. 

Habitat: Solitary or in small groups on the ground in deep forest. 

Distribution: Europe. 

Discussion: As I remarked in the description of the subgenus, the 
characteristics of this species are principally those of Geastrum, for 
which reason Hollos placed it in synonymy with G. fornicatus (G. 
quadrifidum), as the spores and capillitium are similar, but in my 
opinion it is quite different from that species. 

Geastrum conrathii (Hollos) P. Ponce, comb. nov. Gasteropsis 
conrathii Hollos, Nov. Kozlem. 2: 72. 1903. Trichaster conrathii 
(Hollos) Long, Myc. 37: 603. 1945. 

Unexpanded basidiocarp ovate, hypogeous, at maturity epigeous, 10 cm. in 
diameter; exoperidium revolute, thick, coriaceous, splitting to about the middle, 
into 10 unequal subhygroscopic rays; fleshy layer adnate, brown, transversely fis- 
sured into corrugations; outer layer clean, ocher colored, brown and white varie- 
gated, longitudinally striate; base broad, concave with the mycelium attached in 
the center; endoperidium sessible, globose, white, soft, flexible, only fragments 
remaining adherent to the fleshy layer; gleba stipitate, with an angular subligneous 
stipe, 12 mm. wide at top, 20 mm. wide at base, 10 mm. tall; the mass of the gleba 
dark brown, 3 cm. in diameter; columella firm, subglobose, persistent; spores glo- 
bose, some short pedicellate, 6-8/x in diameter; densely verrucose; capillitium sub- 
hyaline to dilute brown, rarely branched, non-septate, 4 M diameter, walls thick, 
lumen small. 


Type locality: Southern Africa. 

Habitat: In granitic soil on grassy slopes, associated with Wel- 
witschia mirabilis. 

Geastrum xylogenum (Long & Stouffer) P. Ponce comb. nov. 
Geaster xylogenus Long & Stouffer, Myc. 40: 563. 1948. 

Unexpanded basidiocarp not found; when expanded, 6-7 cm. broad, concave 
below; outer surface rather naked, almost smooth but faintly longitudinally striate, 

FIG. 2. Geastrum xylogenum (Long) P. 
Ponce. Long 10028. Type. New Mexico. 
Herb. Smithsonian Inst. 

pecan brown; exoperidium rigid, coriaceous, subhygroscopic, splitting nearly three- 
fourths the way to the center into 8 rays, which are unequal, 4-5 cm. long, recurved 
with strongly revolute, acuminate curled tips; fleshy layer usually adnate but some- 
times peeling off above; endoperidium short-stipitate, brittle, deciduous and break- 
ing away at the base, light brown; sterile base and columella not evident; stipe 
stout, subligneous, 5 mm. by 10 mm. thick, 2 mm. high; gleba liver brown, pul- 
verulent; capillitium subhyaline to light brown, 5-6/j. in diameter, unbranched; 
spores spherical, 3.2-5 M in diameter, dark brown, verrucose. 

Type locality: Near Cuba, Sandoval County, New Mexico, 6,400 
ft. elevation. 

Habitat: Solitary on decayed coniferous wood. 
Distribution : Known from type locality only. 

Discussion: We are including this species in the subgenus Tri- 
chaster with G. melanocephalum and G. conrathii from which it differs 
by the absence of columella and by the xylophyllous habit. 

GEASTRUM subgenus Myceliostroma Henn. Hedwigia 43: 185. 

The subgenus Myceliostroma, designated by Hennings, is based 
on the characteristic mycelium which is widely spread on the sub- 
stratum, forming a subiculum upon which one or several basidiocarps 
may develop. In addition, the species included in this division are 
always epigeaous in origin, the spore surfaces are all delicately and 
sparsely echinulate, the mycelium is always basal, the surface of the 


exoperidium may be smooth but more often is velutinate or strigose. 
They are almost constantly caespitose, and some of them develop a 
stipe. The inner peridium is sessile. The mouth is definite. Most 
of them are tropical or subtropical. 

Type species: Geastrum schweinitzii (Berk. & Curtis) Zeller. 
This subgenus includes the following species and varieties : 
Geastrum schweinitzii (Berk. & Curt.) Zeller. 
North and South America, Cuba, Jamaica, Africa, and Java. 

G. schweintzii var. stipitatum P. Ponce. 

Java, Brazil, Panama, and Republic of Congo. 

G. javanicum (Lev.) P. Ponce. 

Southern United States, South America, and Australia. 

G. javanicum var. welwitschii P. Ponce. 

Spain, eastern United States, and Bermuda. 

Geastrum javanicum (Lev.) P. Ponce, comb. nov.Geasterjavan- 
icus Lev. Ann. Sci. Nat. 5 (3): 161. 1846. G. readeri Cook & Massee 
Grev. 16: 73. 1888. G. velutinus Morg. Journ. Cinn. Soc. Nat. Hist. 
18: 38. 1895. G. lloydii Bress. & Pat. in Lloyd, Myc. Writ. 1: 50. 
1901. G. dybowskii Pat. Bull. Soc. Myc. France 18: 300. 1902. 
G. saccatus var. walkeri Coker & Couch, Cast. E. U.S. and Canad. 
113. 1928. Cycloderma ohiensis Cooke, Grev. 2: 95. 1882. 

Unexpanded basidiocarp ovate and bluntly pointed, 2.5 cm. broad, seated on 
the substratum, attached to the mycelium only at the base, surface dull and finely 
felted, tomentose, clean of debris, creamy yellow to flesh color; exoperidium split- 
ting to about the middle, into 5 or 7 reflexed or expanded rays which usually curl 
backward under the basal part which is convex or flat below; the outer layer split 
into two fibrous, persistent layers sometimes becoming semifornicate; fleshy layer 
thin, cream to fleshy color when fresh; endoperidium subglobose, sessile, smooth, 
appearing minutely felty under the lens, dark brown to light grey; mouth small, 
fibrilose, surrounded by a distinct fibrous, conical peristome, silvery-grey or light 
brown; gleba umber; columella obvious and clavate when young, obscure at ma- 
turity; spores globose, 2.5-3.5/x thick, fuscous, distinctly asperulate or finely verru- 
cose; capillitium threads 4-5/x thick, irregular, branched at the tapering tips. 

Distribution: Type locality: Java (and Australia for G. readeri; 
Ohio for G. velutinus), Australia, Africa, North America, Central and 
South America, West Indies. Not reported from Europe. 

Habitat: Epigeous under trees or shrubs. 

Discussion : It is difficult to separate this species from G. saccatum 
and G. fimbriatum. It differs from them in its epigeic habit and in 
the branched capillitium. Although G. fimbriatum also has the tend- 


ency to peel the outer layer of the exoperidium, this layer holds debris 
adhering to it, whereas in G. javanicum this layer remains clean. 
Generally there is a scar at the base of the exoperidium. 

Geastrum javanicum var. welwitschii (Mont.) P. Ponce 
comb. nov. Geaster welwitschii Mont. Syll. Crypt. No. 1048. 1856. 
G. radicans Berk. & Curt. Grev. 2: 49. 1873. G. glaber Lloyd, Myc. 
Writ. 1159. 1922. 

Unexpanded basidiocarp subglobose, not pointed above, reddish brown, 1-2 
cm. in diameter, seated on the substratum that is covered by the mycelial floccu- 
lence; exoperidium splitting into 6-8 rays, the two fibrous layers of which are sep- 
arable except at the tips of the rays, the outer left as a cup with lobed margins on 

FIG. 3 (left). Geastrum javanicum (Lev.) P. Ponce. Herb. Massee. Australia. 
(As type of Geaster readeri Cook & Massee.) N.Y.B.G. 

FIG. 4 (right). Geastrum javanicum var. weluntschii (Mont.) P. Ponce. H. C. 
Beardslee. Florida. N.Y.B.G. 

the substratum, attached to the mycelium by a basic point, the inner everted, 
adopting a fornicate form, its inner layer vaulted in the center elevating the spore 
sac; endoperidium globose, up to 2 cm. broad, with a narrow connective like a short 
pedicel; surface smooth but pitted, reddish brown, mouth sometimes crumpled, 
surrounded by a distinct peristome of radial, silky fibers; gleba ochre-brownish; 
spores spherical, warted, 3.4-5^ in diameter, brown; capillitium threads up to 7.5ju 
in diameter, very irregular with ramifications at the tapering ends. 

Type locality: Spain. 

Distribution: Spain, South Carolina, Florida, Bermuda. 
Habitat: Epigeous on humus, or on wood. 

Discussion : This plant is considered a subspecies of G. readeri by 
Lloyd and closely related to it by Coker and Couch. In my opinion, 


it is only a fornicate variety of G. javanicum. The two layers of the 
exoperidium that separate are fibrous, the mycelium is attached only 
to the basal point, the spores have the same kind of short, wide spines, 
and the capillitium is branched at the ends as in G. javanicum. 

Geastrum schweinitzii (Berk. & Curt.) Zeller, Myc. 40: 649. 
1948. G. mirabile Mont. Ann. Sci. Nat. 4 (3) : 139. 1855. G. rhizo- 
phorum Dissin & Lange, Bull. Jard. Bot. Etat. Bruxellas 32: 373. 

FIG. 5. Geastrum schweinitzii (Berk. & 
Curt.) Zeller. Herb. Massee. (As type of Coilo- 
myces schweinitzii Berk. & Curt.) N.Y.B.G. 

1962. Coilomyces schweinitzii Berk. & Curt. Jour. Acad. Nat. Sci. 
Phil. 2 (2) : 297. 1853. Geaster papyraceus Berk. & Curt. Proc. Amer. 
Acad. Arts & Sci. 4: 124. 1860. G. lignicola Berk. Journ. Linn. Soc. 
Bot. 18: 386. 1881. G. subiculosus Cooke & Massee, Grev. 16: 16. 
1887. G. juruensis Henn. Hedwigia 43: 184. 1904. G. tomentosus 
Lloyd, Myc. Writ. 1 : 185. 1904. G. trichifer Rick, in Lloyd, Myc. 
Writ. 2: 314. 1907. G. caespitosus Lloyd, Myc. Writ. 2: 315. 1907. 

Unexpanded basidiocarp 2-5 cm. broad, globose, obovate, densely caespitose, 
sometimes in considerable number, superficial, mostly epiphytic, arising from a 
white mycelium that covers branches of trees or binds together leaves and twigs; 
strigose-tomentose, whitish to reddish-brown or light olive; exoperidium split to 
about the middle, expanding into 6-8 broad lobes that tend to remain straight and 
upright rather than curl under, the basal part saccate 4-16 mm. broad; outer (my- 
celial) layer thick, persistent tomentose, light olive or whitish; fleshy layer thin, 
wood-brown, persistent, irregularly cracking in dried specimens, endoperidium 
sessile, 6-9 mm. thick, globose, surface smooth or delicately felted, grey with silvery 
lustre; mouth fimbriate, with a silky conical, concolored peristome with a definite 
ring; gleba brown; columella not evident, but present in some forms; spores globose, 
dark, delicately asperate or spinose, 3.2-3.8^ in diameter; capillitium sinuose, thick 
walled, with no lumen, much paler than the spores, about 3.4/z thick with tapering 
ends, not branching. 

Habitat: Epigean, frequently densely caespitose, sometimes soli- 
tary, upon a pallid mycelial subiculum, on decaying wood or other 
vegetable debris on the forest floor. 

Distribution: Type from French Guiana; North America, Brazil, 
Jamaica, Cuba, Ceylon, Japan, Africa, and Australia. 


Discussion: There is no doubt that Coilomyces schweinitzii Berk. 
& Curt, is Geastrum mirabile Mont, as Lloyd indicates in Myc. Notes 
17: 181. 1904 and Zeller formally published in Myc. 40: 649. 1948. 
The epiphytic behavior and the subiculous stromatic mycelia make 
it a very strongly characterized species, which Henning named as the 
type of his subgenus Myceliostroma. The tomentose mycelial layer 
of the exoperidium, the spinose spores, and the unbranched tapering 
capillitium complete the fundamental characteristics of the group 
which includes forms considered as separate species by some my- 

Geastrum schweinitzii var. stipitatum (Solms ex Fischer) P. 
Ponce, comb. nov. Geastrum stipitatum Solms ex Fischer, Hedwigia 
32: 50. 1893. G. congolense Dissing & Lange, Bull. Jard. Bot. Etat. 
Buxelles 32: 373. 1962. 

Unexpanded basidiocarp 2-2.5 cm. broad, globose, upper part obovate, taper- 
ing into a short, broad stipe, superficial, arising from a prominent, white mycelium, 
terrestrial or epiphytic; endoperidium the same as that of G. schweinitzii, but the 
peristome less conical; spores with the same kind of spines and the capillitium sim- 
ilar to those in the original species. 

Type locality: Java. 

Habitat: On wood. 

Distribution: Java, Brazil, Panama, Republic of the Congo; 

Discussion: This species collected by Solms in Java and described 
as a new species, has also been found in the tropics of America and 
Africa. A study of the principal characters of G. stipitatum proves 
it to be only a variety of G. schweinitzii. 

Geastrum floriforme (Vitt.) Cunn. New Zealand Journ. Sci. 
Tech. 23: 172. 1942. Geaster floriforme Vitt. Monogr. Lye. 23. 1842. 
G. spegazzinianus De Toni, Revisio Gen. Cast. 19. 1887. G. delicatus 
Morg. Am. Nat. 21: 1028. 1887. G. pazschkeanus Henn. Hedwigia 
39(B2) : 55. 1900. G. hungaricus Hollos, Gast. Ung. 64. 1904. G. simu- 
lans Lloyd, Lye. Austr. 17. 1905. G. sibiricus Pilat, Bull. Soc. Myc. 
Fran. 51:423. 1935. 

Unexpanded basidiocarp subglobose, depressed globose, sometimes pointed at 
the apex, dirty white, first subterranean, becoming superficial and expanded when 
2-6 cm. diameter; exoperidium split to about the middle into 5-8 subequal, narrow, 
acute, hygroscopic rays, folding over the endoperidium when dry, saccate with the 
tips expanded or revolute when moist; fleshy layer smooth or transversely cracked, 
tenuous, waxy-cartilaginous, pinkish brown; outer layer (mycelial layer) thin, 


whitish, at first with adherent sand, soon flaking away and leaving exposed the 
glabrous, ochraceous or brown fibrous layer; base slightly umbilicate or rounded 
with scar at point of attachment; endoperidium sessile, 1-1.5 cm. diameter, sub- 
globose, oblong, oval, albidocinereuous, to light brown, almost smooth, or covered 
at first with fine granules that later disappear; mouth only a puncture or slit with 
short radiating fissures, naked, with no definite peristome, fimbriate when old; 
gleba umber; columella tenuous, compressed, or none; spores globose, 5-6 M , warted, 
verrucose, reddish brown; capillitium threads varying in thickness but always 
slightly thinner than the spores, almost hyaline, simple or sometimes slightly 
branched near the ends. 

Habitat: In groups on the ground. 

Distribution: Europe, western North America, Australia, New 
Zealand, South Africa. 

Discussion : This species, first described from Europe by Vittadini 
as G. floriforme and later by Morgan from North Dakota as G. deli- 
catus, is characterized by its delicate hygroscopic exoperidium and its 
naked mouth. It may be separated from G. mammosum, to which it 
is very close, by the latter character. 

Geastrum quadrifidum Pers. Syn. Fun. 133. 1801. G. quadrifi- 
dum 7 fenestratum Pers. Syn. Met. Fung. 133. 1801. G. quadrifidum 

FIG. 6. Geastrum quadrifidum Pers. Cooper. N.Y.B.G. 


x majus Abb. & Schw. Consp. 79. 1805. G. quadrifidum DC. Fl. Fr. 
2:267.1815. Geaster fornicatus Fr. Syst. Myc. 3: 12. 1829. G. ma- 
cowani Kalchbr. Fungi Macowaniani, Grev. 10: 108. 1881. G. mar- 
chius P. Hemm. Nat. Pflanz. 1: 231. 1901. G. fenestratus Lloyd, 
Myc. Writ. 1: 70. 1901. G. fenestriatus (Pers.) Cunn. Proc. Linn. 
Soc. NSW. 51: 89. 1926. Plecostoma fornicatum Desv. Journ. Bot. 
Paris 2: 101. 1809. 

Unexpanded basidiocarp globose, at first submerged, medium or large size; 
exoperidium split to about the middle into 4-5 rays; the outer layer remaining 
as a hollow cup in the substratum; the inner fibrous and fleshy layers becoming 
strongly fornicate, attached at tips to the basal cup, rays firm, thick, brown; fleshy 
layer dark brown and adherent or partly peeled off, in old plants worn away; base 
strongly convex; endoperidium subglobose or urnshaped to depressed, dark brown, 
finely velvety on surface, constricted near the base so as to form a ring-like apophy- 
sis above the pedicel; pedicel about 2-3 mm. long; mouth naked, conical or mam- 
miform, tubular, apex fibrillose or lacerate; gleba ferruginous; columella long-ellip- 
tical, 2-3 mm. high; spores globose, 5-6^ in diameter, umber, verrucose, reticulate; 
capillitium threads 10-12^ thick, about as dark as the spores; walls roughened, 
tapering, with ramifications at the ends. 

Habitat: Solitary on vegetable debris on the ground. 
Distribution : Europe, North America, South America, Australia, 
North Africa, South Africa, and Hawaii. 

Discussion: This species is characterized by the fornicate exo- 
peridium and has been confused with G. minimum in Europe and 
North America, and with G. radicans in North America. It may be 
distinguished from them by its naked mouth. This species has been 
called G. fornicatus (Huds.) Fr., a name which is untenable. The 
species is very similar to G. rufescens in fundamental characters, such 
as mouth, surface at inner peridium, separating layers of outer perid- 
ium, and concave base when expanded. 

Geastrum hieronymii (Henn.) P. Ponce, comb. nov. Geaster 
hieronymii Henn. Hedwigia 36: 211. 1897. 

Unexpanded basidiocarp subspherical, slightly pointed, buried until dehis- 
cence; when open 3-7 cm. wide; exoperidium split to about the middle into 6-8 
rays, rigid, involute, acute; outer layer from pale to dark brown, covered with earth 
or debris and tending to split away from the fibrous layer which is chestnut or flesh 
color when fresh; fleshy layer thick when fresh, flesh to brown color; when dry, 
darker, adnate, often peeling away; endoperidium 2-3 cm. wide, subglobose, short 
pedicellate, the stalk flattened with a prominent apophysis, dark brown, harshly 
asperate with acute or subpyramidal spicules; mouth indeterminate, slightly ele- 
vated, fibrous, fimbriate or toothed; gleba sepia, columella globose; spores globose, 
brown, 3.5-6/x in diameter, minutely verruculose; capillitium threads simple, fas- 
ciate, light brownish, 3.5-7^t thick. 


Habitat: Sandy soil mixed with humus in moist forest. 

Distribution : Type from Argentina; also found in Brazil; has been 
collected in Mexico, New Mexico in North America, and in South 

FIG. 7. Geastrum hieronymii P. Henn. Long 9779. New Mexico. N.Y.B.G. 

Discussion : This plant resembles Geastrum rufescens, from which 
it differs by the asperate spore sac and the darker color. 

Geastrum rufescens Pers. Syn. Meth. Fung. 134. 1801. Geaster 
rufescens Fr. Syst. Myc. 18. 1829. G. schaefferi Vitt. Monogr. Lye. 
20. 1842. G. vulgatus Vitt. Monogr. Lye. 20. 1842. G. limbatus sensu 
Morg. Am. Nat. 18: 996. 1884. G. limbatus var. pacificus Morse, 
Myc. 33: 139. 1941. 

Unexpanded basidiocarp large, subspherical, not pointed, entirely buried until 
dehiscence; when open, 5-8 cm. wide; exoperidium split to about the middle into 
6-8 rays, the segments reflexed to form an arch; the outer layer covered with earth 
or debris and tending to split away in patches from the fibrous layer which is flesh 
colored when fresh; fleshy layer very thick when fresh, up to 5 mm.; when dry, 
forming a thinner crust with the appearance of rough leather; endoperidium up to 
4 cm. thick, depressed globose, with a low apophysis around the lower side; when 
fresh, flesh to greyish-flesh colored; when dry, brown to grey brown, the surface 
covered with very minute, scurfy dots; mouth slightly elevated, fibrous, fimbriated 
or toothed, indeterminate; gleba light brown in mass, columella globose; spores 
globose, brown, 3.5-4/x in diameter, minutely warted or asperulate; capillitium 
threads simple, 3.5-4.8/x thick, almost hyaline. 

Habitat: Usually at the base of old oak stumps [Persoon, "... in 
Pinetis." Fries, "Ad terram in pinetis, autumno."] 

Distribution: Europe, North America, Mexico, Japan. 



FIG. 8. Geastrum rufescens Pers. Harper collection 1733. Illinois. Field 

Discussion: This plant has been confused with G. smardae from 
which it differs by its reversed rays, light flesh-colored inner peridium 
and thicker, spongy, fleshy layer. 

Geastrum fimbriatum (Fr.) A. H. Smith, Puffballs and their 
Allies in Michigan, 92. 1951. G. rufescens var. minor Pers. Syn. 
Meth. Fung. 134. 1801. Geaster fimbriatus Fr. Syst. Myc. 3: 16. 1829. 
G. tunicatus Vitt. Monogr. Lye. 162. 1841. G. djakonense Schul. Kuj. 
Rada. Jugosl. Akad. Zwan. Mat. Period. Vazzeda 64: 6. 1882. 
G. infrecuens Lloyd, Myc. Writ. 2: 263. 1906. 

Unexpanded basidiocarp globose, submerged, becoming expanded when 3 cm. 
wide; exoperidium split to about the middle into 6-8 unequal, flaccid, pointed rays, 
whose tips turn under when completely open, leaving the base shallowly saccate; 
fleshy layer bay-brown, continuous, adnate; outer layer tending to split away from 
the others, wholly covered with debris held by the mycelial layer; base concave, 
plane or convex; endoperidium sessile, 0.5-1 cm. diameter, depressed-globose, 
dingy- white to umber brown, glabrous, smooth; mouth indeterminate with fibrous 


or lacerate margin, sometimes surrounded by a slightly depressed and lighter zone, 
sometimes approaching the fibrillose condition; gleba umber; columella inevident; 
spores globose, 3.4/z, fuscous, finely verrucose; capillitium threads unbranched, 
5/t thick, usually lighter than the spores, tapering, undulate. 

FIG. 9. Geastrum fimbriatum (Fr.) A. H. 
Smith. Harper collection S-142. Wisconsin. 
Field Museum. 

Habitat: In small groups on humus under cedar or in mixed woods. 

Distribution: Worldwide. 

Discussion: It is very difficult to separate this species from the 
small forms of G. saccatum, from which it differs externally only by 
the lack of a ring around the mouth area. The spores are smaller 
than those of G. saccatum and the spines are shorter and thinner. 

This species lives in the soil; in rich humus, much debris remains 
attached to the outer layer. The tendency of the outer layer to sep- 
arate from the exoperidium sometimes leads to confusion of this spe- 
cies with G. saccatum and G. readeri. G. fimbriatum is hypogeous, 
always has some adherent debris, and lacks the prominent umbilical 
scar at the base of the exoperidium. 

Geastrum fimbriatum f . pallidum A. H. Smith, Puffballs and 
their Allies in Michigan, 94. 1951. 

Differs from G. fimbriatum by its hyaline spores and capillitium. 
Habitat: On rich humus. 

Distribution: Known only from a single collection at Tahqua- 
menon Falls State Park, Luce County, Michigan. 

Geastrum furfuraceum P. Ponce, sp. nov. Geaster umbilicatus 
sensu Morgan, non Fries, Am. Nat. 18: 966. 1884. Coker & Couch, 
The Cast, of East. U. S. and Canad. 136. 1928. 

Fructificationes juvenes hypogaeae, ovatae. Exoperidium hygroscopicum 8-12 
radiis inaequis, involutis; stratum myceliale facile secedens. Endoperidium glo- 
bosum aut globoso-depressum, sessile, umbrinum, furfuraceum; peristomium coni- 
cum, plicato-striatum, furfuraceum, depreso-marginatum. Sporae 3.5-4 M diameter 


brumae, verrucosae, cum verrucis humibus. Capillitium 3.5-4 M diameter hyalinum 
non ramosum, tortuosum. 

Unexpanded basidiocarp small, globose, subterranean, becoming superficial 
and expanded when 4 cm. wide; exoperidium split about the middle into 8-12 acute, 
hygroscopic rays, unequal in breadth, which are involute when dry; fleshy layer 

FIG. 10. GeastrumfurfuraceumP.Ponce. Ellis N.A.F. 110. Type. New Jer- 
sey. (As Geaster mammosus Chev.) N.Y.B.G. left. 

FIG. 11. Geastrum xerophilum (Long) P. Ponce. Long 2858. Type. New 
Mexico. N.Y.B.G. right. 

thin, smooth, or more or less rimose, umber, adnate; outer layer delicate, floccu- 
lent, mixed with earth and gradually wearing away, leaving the rays smooth and 
glabrous, pale brown or tan, shining with some metallic reflection; base umbilicate; 
endoperidium sessile, 0.7-1.2 cm. diameter, globose or depressed-globose, dark 
brown, sometimes tan, minutely furfuraceous until old; peristome sulcate, furfura- 
ceous, concolorous, conical, seated on a depressed umbilicate zone; mouth fimbri- 
ate; gleba ferruginous; columella not evident; spores globose, 3.5-4/i diameter 
warted or verruculose, reddish umber; capillitium threads attenuated, wavy, 3.5- 
4ju diameter, hyaline. 

Type species: Ellis 110, North American Fungi. New Jersey. 

Habitat: Solitary on humus. 

Distribution: North America. 

Discussion: Although this species presents an umbilicate zone 
around the peristome, as do G. badium, G. campestre, and G. ambig- 
uum, it is separated from them by its smaller spores, more minutely 
warted than those of the other three, and its furfuraceous endope- 
ridium, which is spiny or granular in the others. 

This species, distributed by Ellis as G. mammosus (Ellis 110 from 
New Jersey in his North American Fungi) was considered by Lloyd 
(Geastrae 12. 1902) to be G. drummondii Berk. In the Lloyd Her- 
barium is a Florida collection annotated as G. smithii. 

Geastrum xerophilum (Long) P. Ponce, comb. nov. Geaster 
xerophilus Long, Myc. 34: 13. 1942. Geaster pluriosteus Long & 
Stouffer Myc. 40: 553. 1948. 

Unexpanded basidiocarp subglobose or strongly depressed-globose, small, 1-2 
cm. across, submerged, with the mycelium universal, becoming superficial when 
expanded; exoperidium split to about the middle into 7-12 acute rays which are 


pliable, not hygroscopic, with the tips involute around the endoperidium, saccate; 
fleshy layer cream-buff to cinnamon, adnate, rarely rimose; outer layer covered by 
sand held by the persistent, thin, strongly adnate mycelial layer; base concave 
below, elevating the endoperidium, and with a prominent scar; endoperidium usu- 
ally with a short pedicel, subglobose to strongly depressed on top, 1-2 cm. in diam- 
eter, light buff to drab grey when fresh, whitish with age, densely and minutely 
furfuraceous, lower part enclosed by the saccate exoperidium; peristome small, 
circular, acutely conic, sulcate with unequal ridges, without a surrounding groove, 
concolorous; gleba brown; columella cylindrical; spores spherical, 1-guttulate, 4.2- 
5/x in diameter. 

Habitat: In open sandy soil of semi-arid locations. 

Distribution: New Mexico. 

Discussion: According to Long, this is "the only sulcate-mouthed 
Geaster which is saccate and has a pedicellate spore sac." G. lloydi- 
anum has a sulcate mouth and pedicellate spore sac but is not saccate; 
G. archeri, as noted by Long, is saccate and has a sulcate mouth, but 
the spore sac is sessile. 

Geastrum badium Pers. Journ. Bot. Paris 2: 27. 1809. Geaster 
umbilicatus Fr. Syst. Myc. 3: 14. 1829. G. elegans Vitt. Monog. Lye. 
15. 1842. 

Unexpanded basidiocarp globose, subterranean, becoming superficial and ex- 
panded when 3-4 cm. wide; exoperidium split profoundly in 7-10 unequal hygro- 
scopic rays, involute when dry; fleshy layer thin, smooth, blackish; outer layer 
flocculent with abundant debris and earth adhering, splitting from the fibrous layer 
that shines metallically when exposed; endoperidium sessile, 1 cm. diameter, de- 
pressed-globose, blackish, with the surface wrinkled or minutely granular; peris- 
tome sulcate, concolorous, conical, on depressed umbilicate zone; apex fimbriate; 
gleba reddish-umber; capillitium threads tapering, simple, 6/i in diameter, hyaline; 
spores globose, 5.25-6/i diameter, reddish umber, verrucose. 

Habitat: Solitary on humus. 

Distribution: Europe, North America. 

Discussion: Although this plant was called G. umbilicatus from the 
umbilicate zone around the peristome, this feature is also found in 
G. campestre, G. ambiguum, and G. furfuraceum. It has been con- 
fused with G. furfuraceum from America, from which it is very diffi- 
cult to separate it. They differ in the size of the spores and capillitium, 
the darker color and more rugose endoperidium of the European plant. 

Geastrum campestre (Morg.) Stanek, Fl. CSR, B-l: 463. 1958. 
G. berkeleyi var. continentalis Stanek, Fl. CSR, B-l : 473. 1958. Geaster 
campestris Morg. Amer. Nat. 21: 1026. 1887. G. berkeleyi Massee, 
Amer. Bot. 4: 79. 1891. G. pseudomammosus Henn. Hedwigia 39 


(B2) : 54. 1900. G. asper Lloyd, Geastrae 18. 1902. G. clelandii Lloyd, 
Myc. Writ. 5: 794. 1918. 

Unexpanded basidiocarp globose, small, submerged, becoming superficial and 
expanded when 4 cm. wide; exoperidium split about the middle into 7-12 equal, 
acute, hygroscopic rays usually involute when dry, sometimes revolute; fleshy layer 
umber, adnate, continuous or rimose; outer layer covered with debris held by the 
adnate mycelial layer; base umbilicate; endoperidium shortly pedicellate, depressed 
globose or subglobose, 1.5 cm. diameter, dirty white, greyish, buff, or pale brown, 
minutely but densely covered with whitish or pale brown granules; peristome 
strongly sulcate, conical acute, seated on a depressed zone, concolorous or darker 
than the remainder of the endoperidium; gleba umber, columella present, small, 
spherical; spores 5-6 /t in diameter, verrucose, fuscous; capillitium threads 4-5^ 
diameter, pale, irregularly tapering, some threads with ramifications at the ends. 

Habitat: Solitary or in groups on the ground. 

Distribution: Central and western North America, Australia, 
Europe, and South Africa. 

Discussion : This species is characterized by the asperate or gran- 
ulate endoperidium and the fact that the capillitium threads are 
always thinner than the diameter of the spores. 

It has been collected in different parts of the world, but in North 
America is apparently confined to the area west of the Mississippi 

Geastrum ambiguum Mont. Fl. Boliv. 47. 1839. G. pouzarii 
Stanek, Ces. Mykol. 8: 100. 1954. G. kotlabae Stanek, Fl. CSR, B-l: 
474. 1958. Geaster drummondii Berk, in Hooker's Journ. 5: 1. 1846. 
G. striatulus Kalchbr. & Cke. Grev. 9:3. 1880. G. avellaneus Kalchbr. 
Keadja a Magy. Tud. Acad. 13(8) : . 1883. G. Schweinfurthii Henn. 
in Engl. Bot. Jahrb. 14: 361. 1891. G. involutus Massee, Grev. 21: 
3. 1892. G. smithii Lloyd, Geastrae 21. 1902. 

Unexpanded basidiocarp small, globose, at first submerged, becoming super- 
ficial and expanded when 3 cm. wide; exoperidium split to about the middle into 
8-10 equal, acuminate, hygroscopic rays which are involute over or under the endo- 
peridium when dry; fleshy layer umber, adnate, smooth or transversely rimose; 
outer layer dirty white, covered with debris, becoming partly smooth; base umbili- 
cate; endoperidium sessile or shortly pedicellate, subglobose, 0.8-1.5 cm. diameter, 
dirty white to brownish grey, finely asperate, often becoming smooth with age; 
peristome sulcate, conical, concolorous or slightly darker than the remainder of 
the peridium, seated on a depressed zone which may be wanting; gleba ferruginous, 
brown, columella inevident; spores globose or subglobose, 4.5-6/x diameter, brown, 
sparsely verrucose; capillitium threads attenuated, 3.5-4 M diameter, pale brown to 
almost hyaline, irregular. 

Habitat: In small groups on the ground. 


Distribution: Australia, Tasmania, South Africa, and Santo Do- 
mingo in the West Indies. 

Discussion: This species closely resembles G. badium Pers., from 
which it may be separated by the larger spores and the asperate 

FIG. 12. Geaslrum lloydianum (Rick) P. 
Ponce. Murrill 225. Brasil. (As Geaster 
harriotii Lloyd.) N.Y.B.G. 

Geastrum lloydianum (Rick) P. Ponce, comb. nov. G. saccatum 
var. lloydianum (Rick) Rick, Iherin. Bot. 9: 470. 1961. Geaster lloydi- 
anus Rick, Broteria 5: 26. 1906. G. violacens Rick, Broteria 5: 26. 
1906. G. harriotii Lloyd, Myc. Writ. 2: 311. 1907. G. reinkingii 
Lloyd, Myc. Writ. 7: 1280. 1924. 

Unexpanded basidiocarp globose, submerged, becoming superficial and ex- 
panded when 5 cm. wide; exoperidium split to about the middle into 7-9 subequal, 
acute, revolute or expanded, not hygroscopic, rays; fleshy layer umber, cracking 
and seceding in places; outer layer covered with debris held by the adnate mycelial 
layer; base concave below and elevating the endoperidium; endoperidium sub- 
pedicellate or sessile, 1.5-2 cm. in diameter, depressed-globose, dark brown to 
black, distinctly pitted and roughened but not truly warted or tomentose; peri- 
stome sulcate, conical acute, concolorous or darker; gleba umber; columella rudi- 
mentary; spores globose, 3-4 M in diameter, dark brown, finely spinose under high 
power; capillitium 5.5/i diameter, brown, irregularly tapering, some threads with 
ramifications at the ends. 

Habitat: Solitary on the ground. 

Distribution : Mostly from tropical America, reported in Australia 
and Ceylon. Lloyd mentions a specimen from Spain, in Kew. 

Discussion: The name G. lloydianus Rick (Broteria 5: 27. 1906) 
takes precedence over the name G. harriotii used by Lloyd and 
adopted by Coker and Couch and by Cunningham; I use it to make 
this new combination for this species. 

The species is characterized by the pitted and roughened en- 
doperidium and by the small and finely spiny spores. Although 


these characters were not mentioned in the original description of 
Rick or in the notes of Lloyd, they were observed in the material I 
examined, and were mentioned by Coker and Couch (1928, p. 135) 
as occurring in the specimens sent them by Patouillard. 

Geastrum pectinatum Pers. Syn. Meth. Fung. 132. 1801. G. 
coronatum woodwardii Pers. Syn. Meth. Fung. 132. 1801. G. stri- 
alum DC. Fl. Fr. 2: 267. 1805. G. nanum Pers. Mem. Jard. Bot. 
Paris 2: 27. 1809. G. nanum var. coniferarum Stanek, Fl. CSR, B-l: 
451. 1958. G. hollom Stanek, Fl. CSR, B-l : 467. 1958. Geaster mini- 
mus Chev. Fl. Env. Paris 1: 360. 1826. G. striatus Fr. Syst. Myc. 3: 
13. 1829. G. plicatus Berk. Ann. Nat. Hist. 3: 399. 1839. G. schmi- 
delii Vitt. Monogr. Lye. 157. 1841. G. biplicatus Berk. & Curt. Proc. 
Am. Acad. Art. & Sci. 4: 124. 1860. G. bryantii Berk. Outl. Brit. 
Fung. 300. 1860. G. tenuipes Berk. Fl. Tasm. 2: 264. 1860. G. calycu- 
latus Fuckel, Symb. Myc. 37. 1870. G. umbilicatus Quel. Mem. Soc. 
Em. Monthel. 2: (3) 1873. G. orientalis Hazsl. Grev. 6: 108. 1878. 
G. rabenhorstii Kunze, Mathern. Term. Kozlem 15: 9. 1878. G. turbi- 
natus Cragin, Bull. Washb. Coll. 1 (2) : 39. 1885. G. pseudostriatus 
Hollos, Mathern. Tern. Ert. 19: 505. 1901. G. pectinatus Lloyd, 
Geastrae 15. 1902. G. ellipticis Cunn. (sic) Proc. Linn. Soc. NSW. 
51:77. 1926. 

Unexpanded basidiocarp globose, submerged, with the mycelium universal and 
forming a soft, flocculent coat that holds debris to the entire outer surface of the 
plant, becoming expanded when 3.5 cm. wide; exoperidium split to about the 
middle into 5-12 subequal, acute rays, expanded or subrevolute, central region 
concave below and elevating the spore sac; fleshy layer brown, unequally flaking 
away in patches, leaving exposed the ochraceous fibrous layer; outer layer covered 
with debris held by the adnare mycelial layer, which is persistent but tends to 
flake away; endoperidium pedicellate, subglobose or urceolate, 1-2 cm. wide, brown 
or lead-colored, farinose, base tapering into the pedicel, striate or not, with or 
without apophysis; stalk typically slender, 3-6 mm. long; peristome sulcate, long, 
narrowly conical, concolorous; gleba ferruginous; columella indistinct; spores glo- 
bose, 4.5-5.5/x diameter, dark umber, verrucose, reticulate, with truncate spines; 
capillitium 5/j. diameter, irregularly tapering, with rudimentary ramifications. 

Habitat: Solitary or in groups on the ground among vegetable 

Distribution: Cosmopolitan. 

Discussion: This species, as its many synonyms show, presents 
many variations that have been taken for different species by some 
authors. As noted by Coker and Couch, this species runs into two 
extremes with many intermediate forms. G. schmidelii is the small- 


est and darkest; G. bryantii has a collar-like ring around the base of 
the spore sac; G. plicatus has the base of the endoperidium plicate. 

Geastrum coronatum Pers. Syn. Meth. Fung. 132. 1801. G. 
multifidum DC. Fl. Fr. 2 : 267. 1805. G. atratum Smarda, Ces. Mykol. 
1: 71. 1947. G. coronatum var. muelleri Stanek, Fl. CSR, B-l: 420. 
1958. Geaster limbatus Fr. Syst. Myc. 3: 15. 1829. G. pseudolim- 
batus Hollos, Math. Terem. Ertes. 19: 507. 1901. 

Unexpanded basidiocarp globose, submerged until expanded when 3-6 cm. 
wide; exoperidium split to about the middle into 7-10 unequal, acute rays which 
are expanded and revolute, or partially involute; fleshy layer bay-brown, contin- 
uous or rimose; outer layer covered with debris held by the persistent, adnate 
mycelial layer, in old specimens partially flaking away; base concave or plane; 
endoperidium pedicellate, depressed globose, obovate or subpyriforme, glabrous 
when old, farinose when young, grey to umber, up to 1.5 cm. diameter. Peristome 
depressed, acute, fibrillose, surrounded by a pallid or concolorous, fibrillose or silky 
zone; gleba chocolate; columella almost obsolete; spores globose, 4-4.5/1, fuscous, 
spines acute, densely and coarsely warted; capillitium dark, long, simple, 5-5.3 M , 
some of the threads with rudimentary ramifications. 

Habitat: On the ground in small groups. 

Distribution : Europe, North America, East Africa, Australia and 
New Zealand. 

Discussion : This description refers to the European concept of the 
species but differs from the description of the American material of 
G. limbatus by Coker (1928) by the persistent, universal mycelial 
layer. His plant, as noted by Cunningham (1927) has the exterior 
free of debris, is attached to the substratum by a central basal my- 
celial cord, and the mycelial layer frequently becomes separated from 
the fibrous layer, appearing fornicate. Therefore, the plant described 
by Coker is, in our opinion, G. smardae. 

G. coronatum has been confused in herbaria with G. rufescens; 
however, G. rufescens is pinkish and the fleshy layer of the exo- 
peridium is of a spongy texture. G. coronatum is almost black and 
is firm. 

Geastrum minimum Schw. Syn. Fung. Carol. 58. 1822. G. 
quadrifidum /3 minus Pers. Syn. Meth. Fung. 133. 1801. G. quadrifi- 
dum Ness. Syst. Pilze Schw. 135. 1817. G. minus (Pers.) Cunn. Proc. 
Linn. Soc. NSW. 512: 81. 1926. G. victorinii P, Ponce, Rev. Soc. 
Club. Bot. 3 (3) : 63. 1946. G. dissimile Bottom. Bothalia 4: 592. 1948. 
G. minimum var. fumosicollum Stanek, Fl. CSR, B-l: 435. 1958. 
Geaster marginatus Vitt. Monogr. Lye. 163. 1842. G. cesatii Rabenh. 
Bot. Zeit. 9: 628. 1851. G. granulosus Fuckel, Enumerat. 41. 1860. 


G. coronatus Schroet. Krypt. Fl. Schl. 3: 1889. G. arenarius Lloyd, 
Geastrae 28. 1902. G. leptospermus Atk. & Coker, Bot. Gaz. 36: 1903. 
G. calceus Lloyd, Myc. Writ. 2: 311. 1907. G.juniperinus Macbride, 
Myc. 4:85. 1912. 

FIG. 13. Geastrum minimum Schw. Harper 
collection 91. Michigan. Field Museum. 

Unexpanded basidiocarp subglobose, small, submerged; exoperidium split to 
about the middle into 4-8 unequal rays, commonly recurved or expanded or be- 
coming fornicate by means of the mycelial layer separating from the exoperidium 
in the middle but remaining attached at the tips of the segments, the mycelial 
layer remaining attached to the substratum; fleshy layer brown, rimose, frequently 
flaking away; endoperidium pedicellate, 3-5 mm. diameter, obovate or depressed 
globose, glabrous, farinose, or coated with glistening particles, greyish-brown; ped- 
icel 0.3 mm. long with an apical apophysis; mouth delicately fimbriate (not sulcate); 
peristome silky-fibrillose, the silky area generally outlined by a depressed groove; 
gleba ferruginose; columella inevident; spores 4.5 M , verrucose (spines very wide); 
capillitium 4.5-5^ diameter, fuscus brown, tapering, many with ramifications at 
the ends. 

Habitat: Solitary or in groups on the ground in open places or 
in woods. 

Distribution : Europe, North America, West Indies, Japan, South 
America, New Zealand, and Australia. 

Discussion: This species has been confused with Geastrum quad- 
rifidum, under the name Geaster fornicatus, both in Europe and Amer- 
ica. Geastrum quadrifidum is usually larger and has an indefinite 
mouth area and a smoother surface of the endoperidium. The spores 
are a little larger and the capillitium threads wider. This species is 
quite variable in the way that the exoperidium expands, as sometimes 
it becomes revolute and at other times is fornicate or saccate. Such 
revolute and fornicate forms have been considered as distinct species, 


Geaster coronatus Schroet. and Geaster minus (Pers.) Cunn., but both 
can be found growing together. 

A third form which is not fornicate, and which displays a short 
pedicellate endoperidium, Geaster arenarius Lloyd, is also placed in 
synonymy with Geasterum minimum, as the spores and capillitium 
are the same. 

Geastrum recolligens (Sow.) Desv. Journ. Bot. Paris 2: 102. 
1809. G. hygrometricum var. anglicum Pers. Syn. Meth. Fun. 135. 
1801. G. argenteus Desv. Journ. Bot. 2: 107. 1809. G. mammosus 
Chev. Fl. Paris 1: 359. 1826. G. panjabense Ahmad, Sydowia 4: 127. 
1950. Geaster mammosus Fr. Syst. Myc. 3: 17. 1829. G. lugubris 
Kalchbr. Ertk. Teremes. Korebol. 13: 10. 1884. G. argenteus Cooke, 
Grev. 17: 75. 1889. G. corollinus (Batsch.) Hollos, Magy. Gast. 57. 
1903. Lycoperdon recolligens Sow. English Fungi 4. t. 401. 1809. 

Unexpanded basidiocarp small, bulb-shaped, with a distinct tapering point, 
subterranean, 3-4 mm. long, covered with a thin, soft, buffy yellow coat; mycelium 
basal, at last exposed; exoperidium umbilicate, split almost to the base into 8-10 
equal, slender, rigid, hygrometric rays; fleshy layer thin, brown, smooth; outer 
layer free of debris and wearing off gradually, exposing the fibrous layer with its 
glabrous, shining, copper-brown color; endoperidium sessile, 0.5-1.5 cm. in diam- 
eter, depressed-globose, pale straw color at first, then brown, smooth to finely 
puberulent; peristome definite, silky-strigose, outlined by a depressed border; 
mouth elevated, fibriate; gleba brown; columella short, globose, evident or not; 
spores globose, 3.5-4/i, warted, verrucose, brown; capillitium simple, tapering, 
hyaline, often flattened, 3-4^ diameter (thinner than the spores). 

Habitat: On the ground in open or wooded places. 

Distribution: Europe, North America, South America, and South 

Discussion: This species is characterized by the conical, fimbriate 
mouth with a definite silky peristome outlined by a depressed zone. 
Two species, similar in appearance, G. badium and G. floriforme, may 
be easily separated, as G. badium has a sulcate mouth, and the peri- 
stome of G. floriforme is not clearly defined. Further, the spores of 
G. recolligens are always wider than the capillitium. 

Geastrum smardae Stanek, Ces. Mykol. 10: 18. 1956. G. coro- 
natum sensu Kambly & Lee (non Persoon), Univ. of Iowa Stud. Nat. 
Hist. 17(4): 158. 1936. G. smardae var. slovenicum Stanek, Fl. CSR, 
B-l: 525. 1958. Geaster limbatus sensu Coker & Couch (non Fries), 
Gast. East. U. S. and Canad. 107. 1928. 

Unexpanded basidiocarp ovate with a point up to 10-12 mm. long, half exposed 
or superficial, 1.5-3.5 cm. broad, 3-5 cm. high, with the mycelium attached to a 



FIG. 14. Geastrum smardae Stanek. Harper collection 1334. New Mexico. 
Field Museum. 

basal point, becoming expanded at 3-3.5 cm. wide; exoperidium split to about the 
middle into 5-9 reflexed, long, tapering, usually revolute rays; outer surface firm, 
glabrous, nearly free of debris, the outer layer not cracked into strips or flakes, but 
separating as a rule from the central region, and remaining convex below while the 
inner layer arches upward and elevates the spore sac to produce a fornicate or 
pseudofornicate form; fleshy layer when fresh, 3-4 mm. wide, pale yellowish or 
pinkish; endoperidium subglobose with a more or less obvious apophysis, with a 
short, thick stalk, superficial, pale brown, nearly glabrous; peristome definite, silky, 
broad, conical; mouth fimbriate; gleba ferrugineous to umber; columella subspher- 
ical, about 4-5 mm. thick, persistent; spores globose, 3.7-4.5^, dark, finely and 
closely verrucose, reticulate; capillitium threads brown, straight, tapering, not 
ramified, 4-5^ diameter. 

Habitat: Solitary or in groups on decaying vegetable debris. 

Distribution: North America, Europe, Australia, New Zealand, 
and eastern Africa. 

Discussion: This plant has been confused in North American her- 
baria, as Coker and Couch said, with G. rufescens and G. indicum, 
from which it may be distinguished by its habit of splitting the outer 
layer from the fibrous layer of the outer peridia, over the center and 
proximal part of the rays, which produces its fornicate appearance. 
The nomenclatural confusion with the mostly European G. limbatum 


Fr., a synonym of G. coronatum Pers., has been solved by Stanek's 
creation of G. smardae for the species which Coker and Couch ac- 
cepted as G. limbatus Fr. sensu Bresadola. 

Geastrum saccatum (Fr.) Fischer, Nat. Pflanz. 2(7) : 73. 1933. 
Geastrum saccatum var. cinereum Stanek, Fl. CSR, B-l: 493. 1958. 
G. saccatum var. wichanskyi Stanek, Fl. CSR. B-l : 494. 1958. Geaster 
saccatus Fr. Syst. Myc. 3: 16. 1829. 

FIG. 15. Geastrum saccatum (Fr.) Fischer. 
Banker 742. New York. N.Y.B.G. 

Unexpanded basidiocarp ovate with a point up to 4 mm. long, or with a rounded 
umbo, half-exposed or superficial, 2-3 cm. wide, 3 cm. high, with the mycelium 
attached to a basal point, becoming expanded when 3 cm. wide; exoperidium split 
to about the middle into 5-9 pliable, thin, expanded or revolute, equal, acute rays, 
leaving the base saccate; when open, the rays can attain a width of 1-2 cm.; fleshy 
layer brown, adnate, frequently rimose; outer layer ochraceous buff or cinnamon 
buff, spongy-felted, with very little adhering debris; when dried, irregularly rimose 
or pulling off over considerable areas; with a prominent umbilical scar in the middle 
of the base where the mycelium was attached; endoperidium sessile, 0.5-2 cm. in 
diameter, globose, glabrous, brown, partially enclosed by the saccate base of the 
exoperidium; peristome conic, fibrillose, delimited by a circular, raised or depressed 
line, silky, paler or more brownish than the spore-case wall; gleba umber or paler; 
columella indistinct, or a pseudocolumella present; spores globose, 3.4/x in diameter, 
globose, verrucose, with hyaline or yellowish warts (there are many poorly-formed 
spores in mounts); capillitium 4/1, pale yellowish to brownish, very thick-walled, 
tapering, undulate, often cracked, some threads having ramifications at the ends, 
4 M diameter, incrusted with much debris. 

Habitat: Solitary or in small groups, sometimes gregarious, on 
rich humus, often around old stumps; epigaean. 

Distribution: Cosmopolitan. 

Discussion: This species has been confused with G. fimbriatum in 
many herbaria as they have the same exterior aspect and are of equal 
size and shape. However, the mouth of G. saccatum is surrounded by 
a silky peristome limited by a groove, the button is pointed and free 
of debris, and presents a definite scar in the lower part where the 


mycelia was attached. Further, the spores of G. saccatum are more 
rugose and the capillitium is wider. 

Geastrum saccatum differs from G. indicum only in size, the latter 
being usually larger. The smallest forms of G. indicum and those of 
lageniform habit in the two species are difficult to separate. The 

FIG. 16. Geastrum indicum (Klotz.) Rauschert. Harper collection 3363. 
Michigan. (As Geaster triplex Junh.) Field Museum. 

presence of spores of different size and of abundant debris in the gleba 
of G. saccatus also help to separate these two species. 

Coker and Couch (Gast. E. U. S. & Can. 111. 1928) describe 
a "Geaster saccatus Fr., Northern Form" which seems to differ in no 
way from the typical form. 

Geastrum indicum (Klotz.) Rauschert, Zeits. Pilzk. 25 (2) : 53. 
1959. G. triplex (Jungh.) Fischer in Engler & Prantl, Nat. Pflanzenf. 
2 (7a): 73. 1933. G. triplex var. roseum Stanek, Fl. CSR, B-l: 480. 
1958. G. triplex var. pedicellatum Stanek, Fl. CSR, B-l: 480. 1958. 
Cycloderma indicum Klotzsch, Linnaea 7: 203. 1832. Geaster triplex 
Jungh. Tydsahr. Nat. Gesh. 7: 287. 1840. G. lageniformis Vitt. 
Monogr. Lye. 160. 1842. G. archeri Berk. Fl. Tas. 2: 264. 1860. G. 
australis Berk. Fl. Tas. 2: 265. 1860. G. mammosus var. galericulatus 
Kalchbr. Mather. Term. Kozlem. 2: 153. 1962. G. michelianus W. G. 
Smith, Gard. Chron. 608. 1873. G. cryptorhynchus Hazsl. Grev. 3: 
162.1874. G. dubius Berk. Journ. Linn. Soc. 14:130. 1875. G. kalch- 
brenneri Hazsl. Verhandl. Zool. Bot. Ges. Vien. 26: 76. 1876. G. ca- 
pensis Thumen. Myc. Univ. 715. 1877. G. vitiatus Kalchbr. Ung. 


Acad. Wiss. 17: 34. 1884. G. pittottii Roze, Bull. Soc. Myc. Fr. 4: 
34. 1888. G. coriaceus Col. Trans. N. Z. Inst. 22: 451. 1890. G. en- 
glerianus Henn. in Engl. Bot. Jahrb. 14 : 361. 1891. G. minutus Henn. 
Hedw. 34: 55. 1900. G. morganii Lloyd, Myc. Writ. 1: 80. 1901. 
G. squamosus Lloyd, Myc. Writ. 3: 339. 1907. 

Unexpanded basidiocarp ovate with a point up to 12 mm. long, half exposed 
or superficial, 1.5-3.5 cm. broad, 3.5 cm. high, with the mycelium attached to a 
basal point, becoming expanded at 3-3.5 cm. wide; exoperidium split to about the 
middle into 5-9 plane, or revolute, equal, narrowly acuminate rays, with a flat, 
convex, or, less often, arched base; when open the rays may attain a width of 4- 
5 cm.; fleshy layer thick, brown umber, rimose, frequently partially flaking away, 
sometimes a small portion persisting as a small collar around the base of the endo- 
peridium; outer layer with a very little adhering debris, firm, glabrous, dull yellow- 
ish, mostly adnate, cracking into radial strips or irregular areas; with a prominent 
umbilical scar at the point where the mycelium was attached; endoperidium sessile, 
1.5-2 cm. in diameter, depressed-globose, pulvinate, grey brown or umber, gla- 
brous, smooth, membranous; peristome conic fibrillose, seated on a broad, depressed, 
silky, pallid zone, usually outlined by an upraised margin; gleba ferruginous to 
umber; columella clavate and persistent; spores globose, 3.7-4.4 M , dark, finely and 
closely verrucose, reticulate; capillitium brown, thick-walled, tapering, sometimes 
undulate, some with ramifications at the ends, 4-5/* diameter. 

Habitat: Solitary or in groups on decaying vegetable debris. 

Distribution : Cosmopolitan. 

Discussion: It is often difficult to distinguish G. indicum from 
G. saccatum. However, G. indicum is generally larger. The charac- 
ter used by Junghuhn to establish the species, i.e., the collar-like 
form of the fleshy layer of the exoperidium around the base of the 
endoperidium, is not constant. The longitudinal cracks in the outer 
wall of the exoperidium are not constant either. 


Accepted Names in bold face 

Synonyms in italics 

ambiguum Mont. (Geastrum) 

anglicanum Desv. (Myriostoma) Myriostoma colif orme (Dicks, ex Pers.) 


archeri Berk. (Geaster) Geastrum indicum (Klotz.) Rauschert 

arenarius Lloyd (Geasier) Geastrum minimum Schw. 

argenteum Desv. (Geastrum) Geastrum recolligens (Sow.) Desv. 

argenteus Cooke (Geaster) Geastrum recolligens (Sow.) Desv. 

asper Lloyd (Geaster) Geastrum campestre (Morg.) Stanek 

atratum Smarda (Geaster) Geastrum coronatum Pers. 

australis Berk (Geaster) Geastrum indicum (Klotz.) Rauschert 

avellaneus Kalchbr. (Geaster) Geastrum ambiguum Mont. 


badiumPers. (Geastrum) 
barbata (Dissing & Lange) 

P. Ponce (Gasteroides) 
barbalum Dissing & Lange 

(Geastrum) Gasteroides barbata (Dissing & Lange) 

P. Ponce 

berkeleyi Massee (Geaster) Geastrum campestre (Morg.) Stanek 

berkeleyi var. continentalis Stanek 

(Geastrum) Geastrum campestre (Morg.) Stanek 

biplicatus Berk. & Curt. (Geaster). . . Geastrum pectinatum Pers. 

bryantii Berk. (Geaster) Geastrum pectinatum Pers. 

caespitosus Lloyd (Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

calceus Lloyd (Geaster) Geastrum minimum Schw. 

calyculatus Fuckel (Geaster) Geastrum pectinatum Pers. 

campestre (Morg.) Stanek 


campestris Morg. (Geaster) Geastrum campestre (Morg.) Stanek 

capensis Thumen (Geaster) Geastrum indicum (Klotz.) Rauschert 

cesatii Rabenh. (Geaster) Geastrum minimum Schw. 

delandii Lloyd (Geaster) Geastrum campestre (Morg.) Stanek 

coliforme (Pers.) Corda (Myrio- 


coliforme Pers. (Geastrum) Myriostoma coliforme (Dicks, ex Pers.) Corda 

coliformis Fr. (Geaster) Myriostoma coliforme (Dicks, ex Pers.) Corda 

columnatus Lev. (Geaster) Myriostoma coliforme (Dicks, ex Pers.) Corda 

congolense Dissing & Lange 

(Geastrum) Geastrum schweinitzii var. stipitatum 

P. Ponce 
conrathii (Hollos) P. Ponce 

conrathii Hollos (Gasteropsis) Geastrum conrathii (Hollos) P. Ponce 

conrathii (Hollos) Long 

(Trichaster) Geastrum conrathii (Hollos) P. Ponce 

coriaceus Col. (Geaster) Geastrum indicum (Klotz.) Rauschert 

corollinus (Batsh.) Hollos (Geaster) . Geastrum recolligens (Sow.) Desv. 
coronatum Pers. (Geastrum) 

coronatum var. mulleri Stanek 

(Geastrum) Geastrum coronatum Pers. 

coronatum sensu Kambly & Lee 

(non Pers.) (Geastrum) Geastrum smardae Stanek 

coronatus Schroet. (Geaster) Geastrum minimum Schw. 

cryptorhynchus Hazsl. (Geaster) . . . .Geastrum indicum (Klotz.) Rauschert 

delicatus Morg. (Geaster) Geastrum floriforme (Vitt.) Cunn. 

dissimile Bottom. (Geastrum) Geastrum minimum Schw. 

djaconense Schul. (Geaster) Geastrum fimbriatum (Fr.) A. H. Smith 

drummondii Berk. (Geaster) Geastrum ambiguum Mont. 

dubius Berk. (Geaster) Geastrum indicum (Klotz.) Rauschert 


duplicatus Chev. (Geaster) Astraeus hygrometricus (Pers.) Morg. 

dybowskii Pat. (Geasfer) Geastrum javanicutn (Lev.) P. Ponce 

elegans Vitt. (Geaster) Geastrum badium Pers. 

ellipiicis Cunn. (Geaster) Geastrum pectinatum Pers. 

englerianus Henn. (Geasier) Geastrum indicum (Klotz.) Rauschert 

fenestratus Lloyd (Geaster) Geastrum quadrifidum Pers. 

fenestriatus (Pers.) Cunn. (Geaster). .Geastrum quadrifidum Pers. 

fibrillosus Schw. (Geaster) Astraeus hygrometricus (Pers.) Morg. 

fimbriatum (Fr.) A. H. Smith 

fimbriatum f. pallidum 

A. H. Smith (Geastrum) 

fimbriatus Fr. (Geaster) Geastrum fimbriatum (Fr.) A. H. Smith 

floriforme (Vitt.) Cunn. 


floriformis Vitt. (Geaster) Geastrum floriforme (Vitt.) Cunn. 

fornicatum Desv. (Plecostoma) . . . .Geastrum quadrifidum Pers. 

fornicatus Fr. (Geaster) Geastrum quadrifidum Pers. 

fornicatus Hollos (Geaster) Geastrum melanocephalum (Czern.) Stanek 

furfuraceum P. Ponce 

glaber Lloyd (Geaster) Geastrum javanicutn (Lev.) P. Ponce 

granulosus Fuckel (Geaster) Geastrum minimum Schw. 

hariotii Lloyd (Geaster) Geastrum lloydianum (Rick) P. Ponce 

hieronimii (Henn.) P. Ponce 

hieronymii Henn. (Geaster) Geastrum hieronimii (Henn.) P. Ponce 

hollosii Stanek (Geastrum) Geastrum pectinatum Pers. 

huvgaricus Hollos (Geaster) Geastrum floriforme (Vitt.) Cunn. 

hygrometricum Pers. (Geastrum) . . . Astraeus hygrometricus (Pers.) Morg. 
hygrometricum var. anglicum Pers. 
(Geastrum) Geastrum recolligens (Sow.) Desv. 

indicum (Klotz.) Rauschert 

indicum Klotz. (Cycloderma) Geastrum indicum (Klotz.) Rauschert 

infreqwns Lloyd (Geaster) Geastrum fimbriatum (Fr.) A. H. Smith 

involutus Massee (Geaster) Geastrum ambiguum Mont. 

javanicum (Lev.) P. Ponce 

javanicum var. welwitschii 
P. Ponce (Geastrum) 

javanicus Lev. (Geaster) Geastrum javanicum (Lev.) P. Ponce 

juniperin us Macbride (Geaster) . . . .Geastrum minimum Schw. 

juruensis Henn. (Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

kalchbrenneri Hazsl. (Geaster) Geastrum indicum (Klotz.) Rauschert 

kotlabae Stanek (Geastrum) Geastrum ambiguum Mont. 

lageniformis Vitt. (Geaster) Geastrum indicum (Klotz.) Rauschert 


leptospermus Atk. & Coker 

(Geaster) Geastrum minimum Sehw. 

lignicola Berk. (Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

limbatus Fr. (Geaster) Geastrum coronatum Pers. 

limbatus sensu Morg. (Geaster) . . . .Geastrum rufescens Pers. 
limbatus sensu Coker & Couch 

(non Fr.) (Geaster) Geastrum smardae Stanek 

limbatus var. pacificus Morse. 

(Geaster) Geastrum rufescens Pers. 

lloydianum (Rick) P. Ponce 

lloydianus Rick (Geaster) Geastrum lloydianum (Rick) P. Ponce 

lloydii Bress. & Pat. (Geaster) Geastrum javanicum (Lev.) P. Ponce 

lugubris Kalchbr. (Geaster) Geastrum recolligens (Sow.) Desv. 

mammosum Chev. (Geastrum) Geastrum recolligens (Sow.) Desv. 

mammosus Fr. (Geaster) Geastrum recolligens (Sow.) Desv. 

mammosus var. galericulatus 

Kalchbr. (Geaster) Geastrum indicum (Klotz.) Rauschert 

macoivani Kalchbr. (Geaster) Geastrum quadrifidum Pers. 

marchius Henn. (Geaster) Geastrum quadrifidum Pers. 

marginatus Vitt. (Geaster) Geastrum minimum Schw. 

melanocephalum (Czern.) 
Stanek (Geastrum) 

melanocephalum Czern. 

(Trichaster) Geastrum melanocephalum (Czern.) Stanek 

michelianus W. G. Smith (Geaster). . Geastrum indicum (Klotz.) Rauschert 

minimum Schw. (Geastrum) 

minimum Chev. (Geaster) Geastrum pectinatum Pers. 

minimum var. fum,osicollum 

Stanek (Geaster) Geastrum minimum Schw. 

minus (Pers.) Cunn. (Geastrum) . . .Geastrum minimum Schw. 

minulus Henn. (Geaster) Geastrum indicum (Klotz.) Rauschert 

mirabile Mont. (Geastrum) Geastrum schweinitzii (Berk. & Curt.) Zeller 

morganii Lloyd (Geaster) Geastrum indicum (Klotz.) Rauschert 

multifidum DC. (Geastrum) Geastrum coronatum Pers. 

multifidus Hazsl. (Geaster) Geastrum fimbriatum (Fr.) A. H. Smith 

nanum Pers. (Geastrum) Geastrum pectinatum Pers. 

nanum var. coniferarum Stanek 

(Geastrum) Geastrum pectinatum Pers. 

ohiensis Cooke (Cycloderma) Geastrum javanicum (Lev.) P. Ponce 

orientalis Hazsl. (Geaster) Geastrum pectinatum Pers. 

panjabense Ahmad. (Geastrum) .. . .Geastrum recolligens (Sow.) Desv. 
papyraceus Berk. & Curt. (Geaster) . Geastrum schweinitzii (Berk. & Curt.) Zeller 
pazschkeanum Henn. (Geaster) Geastrum floriforme (Vitt.) Cunn. 

pectinatum Pers. (Geastrum) 

pectinatus Lloyd (Geaster) Geastrum pectinatum Pers. 


pillotii Roze (Geaster) Geastrum indicum (Klotz.) Rauschert 

pluriosteus Long & Stouffer 

(Geaster) Geastrum xerophilus (Long) P. Ponce 

plicatus Berk. (Geaster) Geastrum pectinatum Pers. 

pouzarii Stanek (Geastrum) Geastrum ambiguum Mont. 

pseudolimbatum Hollos (Geaster) . . . Geastrum coronatum Pers. 
pseudomammosus Henn. (Geaster) . .Geastrum campestre (Morg.) Stanek 
pseudostriatus Hollos (Geaster) . . . .Geastrum pectinatum Pers. 

quadrifidum Pers. (Geastrum) 

quadrifidum DC. (Geastrum) Geastrum quadrifidum Pers. 

quadrifidum Schw. (Geastrum) Geastrum quadrifidum Pers. 

quadrifidum Nees (Geastrum) Geastrum minimum Schw. 

quadrifidum majus Abl. & 

Schw. (Geastrum) Geastrum quadrifidum Pers. 

quadrifidum /3 minus Pers. 

(Geastrum) Geastrum minimum Schw. 

quadrifidum 5 fenestratum Pers. 

(Geastrum) Geastrum quadrifidum Pers. 

rabenhorstii Kunze (Geaster) Geastrum pectinatum Pers. 

radicans Berk. & Curt. (Geaster).. .Geastrum javanicum var. welwitschii 

P. Ponce 

readeri Cooke & Massee (Geaster) . . Geastrum javanicum (Lev.) P. Ponce 
recolligens (Sow.) Desv. 

recolligens Sow. (Lycoperdon) Geastrum recolligens (Sow.) Desv. 

reinkingii Lloyd (Geaster) Geastrum lloydianum (Rick) P. Ponce 

rhizophorum Dissing (Geastrum). . . Geastrum schweinitzii (Berk. & Curt.) Zeller 

rufescens Pers. (Geastrum) 

rufescens FT. (Geaster) Geastrum rufescens Pers. 

rufescens var. minor Pers. 
(Geastrum) Geastrum fimbriatum (Fr.) Smith 

saccatum (Fr.) Fischer 

saccatum var. cinereum (Stanek) 

(Geastrum) Geastrum saccatum (Fr.) Fischer 

saccatum var. lloydianum (Rick) 

Rick (Geastrum) Geastrum lloydianum (Rick) P. Ponce 

saccatus Fr. (Geaster) Geastrum saccatum (Fr.) Fischer 

saccatus var. walkeri Coker & 

Couch (Geaster) Geastrum javanicum (Lev.) P. Ponce 

schaefferi Vitt. (Geaster) Geastrum rufescens Pers. 

schmidelii Vitt. (Geaster) Geastrum pectinatum Pers. 

schweinfurthii Henn. (Geaster) Geastrum ambiguum Mont. 

schweinitzii (Berk. & Curt.) 
Zeller (Geastrum) 


schweinitzii Berk. & Curt. 

(Coilomyces) Geastrum schweinitzii (Berk. & Curt.) Zeller 

schweinitzii var. stipitatum 

P. Ponce (Geastrum) 

sibiricum Pilat (Geaster) Geastrum floriforme (Vitt.) Cunn. 

simulans Lloyd (Geaster) Geastrum floriforme (Vitt.) Cunn. 

smardae Stanek (Geastrum) 
smardae var. slovenicum Stanek 

(Geastrum) Geastrum smardae Stanek 

smithii Lloyd (Geaster) Geastrum ambiguum Mont. 

spegazzinianus DeToni (Geaster) . . .Geastrum floriforme Vitt. 

squamosus Lloyd (Geaster) Geastrum indicum (Klotz.) Rauschert 

stipitatum Solms. & Rick 

(Geastrum) Geastrum schweinitzii var. stipitatum 

P. Ponce 
striatulus Kalchbr. & Cooke 

(Geaster) Geastrum ambiguum Mont. 

striaium DC. (Geastrum) Geastrum pectinatum Pers. 

striatus Fr. (Geaster) Geastrum pectinatum Pers. 

subiculosus Cooke & Massee 

(Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

tenuipes Berk. (Geaster) Geastrum pectinatum Pers. 

texensis Long (Geasteroides) 

texensis (Long) Fischer 

(Geasteropsis) Geasteroides texensis Long 

texensis (Long) Long (Terrostella).. Geasteroides texensis Long 

tomentosus Lloyd (Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

trichifer Rick (Geaster) Geastrum schweinitzii (Berk. & Curt.) Zeller 

triplex Jungh. (Geaster) Geastrum indicum (Klotz.) Rauschert 

triplex (Jungh.) Fischer (Geastrum). Geastrum indicum (Klotz.) Rauschert 

triplex var. pedicellatum Stanek 

(Geastrum) Geastrum indicum (Klotz.) Rauschert 

triplex var. roseum Stanek 

(Geastrum) Geastrum indicum (Klotz.) Rauschert 

tunicatus Vitt. (Geaster) Geastrum fimbriatum (Fr.) A. H. Smith 

turbinatus Cragin (Geaster) Geastrum pectinatum Pers. 

umbilicatus Fr. (Geaster) Geastrum badium Pers. 

umbilicatus Quel. (Geaster) Geastrum pectinatum Pers. 

umbilicatus Fr. sensu Morg. 

(Geaster) Geastrum furfuraceum P. Ponce 

velutinus Morg. (Geaster) Geastrum javanicum (Lev.) P. Ponce 

victorinii P. Ponce (Geastrum) Geastrum minimum Schw. 

violasceus Rick (Geaster) Geastrum lloydianum (Rick) P. Ponce 

vittatus Kalchbr. & Cooke 

(Geaster) Geastrum indicum (Klotz.) Rauschert 

vulgatum Vitt. (Geaster) Geastrum rufescens Pers. 


welwitschii Mont. (Geaster) Geastrum javanicum var. welwitschii 

P. Ponce 

woodwardii Pers. (Geastrum) Geastrum pectinatum Pers. 

xerophilum (Long) P. Ponce 


xerophilus Long (Geaster) Geastrum xerophilum (Long) P. Ponce 

xylogenum (Long & Stouffer) 

P. Ponce (Geastrum) 
xylogenus Long & Stouffer 

(Geaster) Geastrum xylogenum (Long & Stouffer) 

P. Ponce 


aj finis Col. (Geaster) linkii Spreng. (Geaster) 

annulatus Lloyd (Geaster) menziesii Berk. (Geaster) 

bancroftii E. & E. (Geaster) persimile Rick. (Geastrum) 

bovista Klotz. (Geaster) pusillus Fr. (Geaster) 

corruptus Syd. (Geaster) queletii Hazsl. (Geaster) 

guilfoyleyi Mull. (Geaster) vellereus Morg. (Geaster) 

Nomen nudum : Geasteropsis stahelii Fischer in Engler & Prantl, Nat. Pflanzenf. 
2(7a): 75. 1933. 

FIG. 17. A, Geastrum ambiguum Mont. B, Geastrum saccalum (Fr.) Fischer. 
C, Geastrum badium Pers. D, Geastrum indicum (Klotz.) Rauschert. E, Geastrum 
campestre (Morg.) Stanek. F, Geastrum lloydianum (Rick) P. Ponce. 


FIG. 18. A, Geastrum schweinitzii (Berk. & Curt.) Zeller. B, Geastrum rufes- 
cens Pers. C, Geastrum recolligens (Sow.) Desv. D, Geastrum fimbriatum (Fr.) 
A. H. Smith. E, Geastrum smordae Stanek. F, Geastrum coronatum Pers. 


FIG. 19. A, Geastrum quadrifidum Pers. B, Geastrum furfuraceum P. Ponce. 
C, Geastrum minimum Schw. D, Geastrum xerophylum (Long) P. Ponce. E, Gea- 
strum floriforme (Vitt.) Cunn. F, Geastrum pectinatum Pers. 


FIG. 20. A, Geastrum javanicum (Lev.) P. Ponce. B, Geastrum hieronymii 
P. Henn. C, Geastrum xylogenum (Long) P. Ponce. D, Geastrum melanocephalum 
(Czern.) Stanek. E, Gasteroides texensis Long. F, Myriostoma coliforme (Dicks 
ex Pers.) Corda. 




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Publications 1036, 1037, 1038, and 1039