Title: Lycoperdaceae_and_Geastraceae
Author: Ian Gibson
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Trial field key to the PUFFBALLS without long stalks and EARTHSTARS of the Pacific Northwest
Prepared for the Pacific Northwest Key Council
By Robert Ramsey, Puget Sound Mycological Society
Copyright  © 1978, 1985, 1998, 2003, 2019 Pacific Northwest Key Council
Minor revisions by Ian Gibson 2003 and 2019

TABLE OF CONTENTS
Note on revisions	1
Introduction	1
Key to genera of mature stages	3
Key to species	4
	Bovistella	4
	Geastrum and Astraeus	4
	Disciseda and Abstoma	6
	Bovista	7
	Lycoperdon	8
	Mycenastrum	11
	Vascellum	11
	Calbovista	12
	Calvatia	12
Glossary	17
Acknowledgements	18
Selected References	19
Index		20

NOTE ON REVISIONS

This key was reformatted, but is otherwise the same as the key Robert Ramsey revised in 1998 for Lycoperdales, apart from a few added REMARKS, indicated as such, a list of Selected References, and a few items added to the Glossary. Lycoperdaceae had been absorbed into Agaricaceae and any comments about families was updated. Abstoma was added. Lycoperdon pyriforme was changed to Apioperdon pyriforme.

INTRODUCTION

The Family Agaricaceae now contains the true puffballs, and the Family Geastraceae contains the earthstars in the genus Geastrum. The earthstars in the genus Astraeus are covered here because of their similarity to Geastrum, but are also considered along with the earthballs in the separate Key to Sclerodermatacae. They are in the Family Diplocystidiaceae. Stalked puffballs are considered in the Key to Desert Fungi. The following key will attempt to guide the investigator in making accurate separations and ultimate identifications.

There are over 250 species of gasteromycetes reportedly identified from Western Montana, Idaho, Oregon and Washington. The Family Agaricaceae contributes over 50 species to this total. When you pick up a fungal fruiting body which appears to form its spore mass internally, there will be certain other characteristics to compare before you can settle on the Family Agaricaceae.

The fruiting body is a relatively simple structure, more or less globose in shape. The name, gasteromycete, means "stomach fungus", in reference to its shape, although the appellation hardly fits some of the more specialized species. The mature fruiting body consists of a spore case filled with spores and parts of sterile modified hyphal threads called capillitium. The whole mass is referred to as the gleba. Some genera have significant volume in a sterile base, or sub-gleba, where no spore formation takes place. The surrounding wall is known as the peridium. The structure is usually anchored to the substratum by rootlike structures which vary from fine fibers to a thick cord. The base of the fruiting body may be elongated into a stemlike structure, as in Lycoperdon perlatum, or the spherical character may be scarcely interrupted by a few rhizoids, as in Bovista.

One of the difficult problems in identifying within this group of fungi is the time span between the condition of a white interior, necessary for eating, and the differentiation necessary for maturity on which good identification can be made. However, we will attempt to make identifications, for this publication, on the basis of mature visual characteristics. In any case, having immature and mature specimens on hand for each species will improve the chances for identification. Occasionally, it will be possible to identify on the basis of locality and habitat, though this may not be infallible since the distribution of many of the species is not well known. Spore sizes and shapes and the microscopic appearance of the capillitia are not given since the macro characteristics are distinctive enough to make identification. In fact, it is often harder to identify by spores than by visual external and internal features. If this does not prove to be the case, then it will be our description which has failed.

To determine that a mushroom is applicable to this key, it must fit each of the following characteristics:

1.	The fruiting body, prior to maturity, is fleshy, spongy or soft rubbery to pressure. Three species become hard rather early.

2.	Upon maturity, it is visible on soil, turf, duff or rarely on well-rotted wood. It may start beneath the surface but will always mature visibly.

3.	The gleba becomes disorganized by maturity, becoming a gooey mass then drying to become powdery to cottony.

4.	The peridium opens in a more-or-less characteristic manner, for each genus, ranging from a specialized apical pore in a Geastrum to simply falling apart in chunks to expose the spore mass as in some of the Calvatias.

5.	The peridium does not contain a gelatinous layer.

6.	The peridium is generally of 2 layers; in Geastrum the outer layer (exo) is sometimes subdivided into 3 layers. In some species the exo may well have disappeared prior to discovery; in others, it may not be obvious even when present. Inner peridium (endo) is usually papery and thin, sometimes thicker and somewhat corky.

7.	Capillitium (sterile threads) will always be present but only visible under a microscope.

The species included here represent the recorded species found in the Pacific Northwest. However, it is certain that other species will be identified within the next few years as more specimens are brought to our attention and as our identification techniques and experience improve.

TRIAL KEY TO GENERA OF MATURE STAGES OF PUFFBALLS 
WITHOUT LONG STEMS IN AGARICACEAE & GEASTRACEAE

1a	Spore case opens by pore or fissure at apex	2
1b	Spore case opens by other means	6

2a	Base extending downward from pear-shaped body in a taproot-like pseudorhiza (radicating)	Bovistella A
2b	Bases of various shapes but not radicating	3

3a	Fruiting body outer layer splitting and opening to form a star-like structure with a persistent spore sac at the center	Geastrum and Astraeus B
3b	Fruiting body loses part or all of outer layer but form does not become star-like	4

4a	Upper portion of sandy outer case falls away to reveal blue-gray spore sac sitting in lower cup, similar to acorn; small	Disciseda and Abstoma C
4b	Fruiting body sheds outer layer in various ways but does not retain conspicuous sandy basal portion	5

5a	Spores release slowly from persistent cottony / felty cushion inside of spore case; body spherical	Bovista D
5b	Spores loose and powdery inside of spore case. Any fibers much looser and softer than Bovista; body with extended base on most specimens	Lycoperdon E

6a	(1b)  Spore case opens by longitudinal slits which form more or less stellate reflexed lobes, exposing gleba (not endoperidium)	Mycenastrum F
6b	Opens by large radial tears, or general fragmentation with some sterile bases persistent, or decomposition of peridia to form very large pore	7

7a	Exo- and endoperidium decomposing simultaneously across top to form very large pore, longer than wide, often to full diameter of top; distinct separation between sterile base and gleba	Vascellum G
7b	Peridium opening by large radial tears or by general fragmentation	8

8a	Sterile base well developed; endoperidium a delicate shiny membrane; exoperidium broken into 5-8 mm thick, truncated, 3 to 6-sided pyramids with transverse parallel lines along edges	Calbovista H
8b	Sterile base present or absent; peridia either not separable or endoperidium not shiny (except the distinctive C. rubroflava); ornamentation of outer layer in some species similar to Calbovista but different in detail	Calvatia I

A.   BOVISTELLA

Only one species reported in Northwest to date but a very small (5-8 mm) Bovistella is recorded just east of the Rocky Mountains and may at some time be discovered on the west side.

Bovistella radicata
Generally 3 to 7.5 cm broad but occasionally up to 14 cm. Higher than broad, turbinate, with a thick well-developed pseudorhiza. Smooth outline to plicate. Color white, to yellowish near maturity. Peridium duplex with exo upper surface covered with a coating of slender spines, many of which are united at the tips into fascicles. Exo sloughs away irregularly and endo eventually opens by an apical pore or slit which gradually enlarges to expose most of gleba. Well-developed sterile base, occupying most of narrowed lower part of body and extending up sides to form a cup in which the gleba is situated. Turfed areas, range lands, idle cultivated fields. Late summer and fall. Widespread but not common.

B.   GEASTRUM & ASTRAEUS

Nine to fourteen species recorded in Northwest to date. Mostly formed below the surface until mature when thrust upward by the reflexing sections of the 3-layered exo. Apical pore is a preformed structure known as a peristome, not resulting from a rupture or tear, except in 2b below. Spore sac papery but relatively tough.

1a	Rays hygroscopic (expanding when wet - curving upward over spore sac when dry)	2
1b	Rays not hygroscopic (remaining open or curved away from spore sac, wet or dry)	5

2a	Spore case opening by distinct peristome	3
2b	Spore case opening by slit, rupture or irregular hole. (Astraeus is included here because of its similarity to Geastrum)	4

3a	Spore case whitish; peristome with faint, short radiating lines to its base, not grooved	Geastrum floriforme
(also known as Geastrum delicatus Morg.)
Specimens typically small with expanded width of 2.5 cm, submerged until mature; rays 8 to 10, unequal, hygroscopic and expanding quickly when put into water, covered at first with sandy soil held on by mycelium, this flaking off by degrees to leave the surface smooth and pale after exposure, fleshy layer of rays dark brown, adnate, continuous then rimose. Spore case 6-8 mm across, subglobose or depressed, sessile, pale, covered with fine granules or furfuraceous particles at first which soon wear away in large part; peristome short, not well defined, with short radiating lines, with no defined or silky area around it, after a time the margin becoming fimbriate. Sandy soil, eastern Washington (Cheney). Small size, low visibility make collection difficult. REMARKS Geastrum corollinum (Batsch) Hollós (with which G. recolligens often synonymized) has a comparatively well-delimited peristome, a mycelial layer that does not bind debris, and smaller spores (Pegler et al. 1995) – I. Gibson

3b	Spore case brown, peristome lacerate / fimbriate	Geastrum recolligens
To 3.5 cm across opened rays. Exterior surface of rays whitish, bleaching to near white in desert sun. Interior (upper) surface of rays dark brown. Spore sac brown and sessile. Rays 3-layered, outer a thin felt soon glabrous. Peristome lacerate / fimbriate. Range lands in southeastern Oregon and Southwestern Idaho. REMARKS 1) This name is often synonymized with Geastrum corollinum. 2) There is a herbarium collection labeled Geastrum campestre, and another labelled Geastrum umbilicatum, both deposited by Paul Kroeger and from British Columbia. Both might key out here. Both have distinct, plicate peristomes, but only G. campestre has a pedicel. – Ian Gibson

4a	(2b)  Small (1-4 cm open), outer surface of rays with black fibrils		
		Astraeus hygrometricus
To 4 cm across rays when open, arising from black hairlike rhizomorphs, some of which may adhere to the base of the fruit body as appressed hairs. Exo splitting into 7-15 pointed rays which bend backward when wet and curve inward over the spore sac when dry. Spore sac sessile, pallid grayish, opening by a slit, tear or irregular pore (not a peristome). Gleba white when young, cocoa brown at maturity. Gregarious to scattered on sandy areas and old fields, widely distributed across North America.

4b	Large (10-15 cm open), surface of rays with brown fibrils	Astraeus pteridis
Up to 15 cm across expanded rays, outer surface with a thin coating of matted brownish fibrils which wear away or remain as a rough exterior coating around the base. Rays 3-6 mm thick, hard and woody when dry; coriaceous to semi-woody when moist; sordid brownish over exterior surfaces. Interior surfaces of rays with thin softer whitish layer which soon becomes conspicuously checked or areolate. Spore case globose, 2.5 to 4 cm diameter, delicate and fibrillose, sordid gray brown not opening by pore but by rupture. Gleba dull purplish brown. Single or in small groups on thin soil or humus in fall.

5a	(1b)  Spore sac opens by a slightly elevated but not sharply defined opening, lacking a definite peristome	Geastrum rufescens
To 6 cm across when open, outer surface with much trash not separating from it cleanly. Spore case on short thick stalk. Fawn colored exterior surface. Exo with thin separable mycelial layer which peels off along with the attached layer of debris. Cortex thick, coriaceous, splitting about halfway to base into 5 to 9 lobes with acute tips, occasionally split into secondary lobes. Fleshy layer 5 mm + when fresh, forming a thin continuous crust on the surface or breaking up somewhat as in G. triplex. Spore sac unpolished to granular velvety, mouth slightly elevated, fibrous and becoming fimbriate. More or less buried in duff near hardwoods, gregarious to clustered or in gregarious clusters. Rare in Idaho and Washington in late summer and fall. (An eastern species for the most part.)

5b	Spore sac opens by a distinct cone-shaped projection or peristome	6

6a	Spore sac sits up on short pedicel	7
6b	Spore sac sessile in cup	8

7a	Area of apical pore paler than remainder of spore sac; outer layer of rays clean of trash	Geastrum limbatum
6 cm or more across opened rays. Short pallid pedicel. Rays dull cinnamon on upper surface. Spore sac dark wood brown to light drab. Outer surface drying hard, separating cleanly from medial layer over the central part and almost to the tips of the rays, leaving a cup in the ground to which the tips of the rays are attached. On range lands in Idaho. REMARKS This is G. limbatum Fr. sensu Bres. – Ian Gibson

7b	Area of apical pore not paler than remainder of sac; outer layer of rays covered with dense coat of white mycelium and debris	Geastrum quadrifidum
(Synonyms include G. coronatum  (Schaeff.) J. Schroet.)
To 2 cm across opened rays; ordinarily much smaller than G. limbatum; acorn-shaped spore sac dark wood brown to purplish drab. Outer wall splitting into 4 to 6 strongly recurved rays from which mycelial layer may become stripped and remain as an indistinct cup below or else continue to adhere to and remain attached to mycelial cover only at the tips. Fleshy inner layer of rays often scaling more or less completely from the whitish fibrillose mid-layer which remains. Idaho (Owyhee County) sagebrush hills, all months except freezing weather. REMARKS This species is “fornicate” (see Glossary). A larger fornicate species, G. fornicatum, has been reported from California and occasionally from the Pacific Northwest. – Ian Gibson

7c	Not fitting the criteria in 7a or 7b, distinct, plicate peristome, short pedicel		
		consider Geastrum pectinatum, Geastrum striatum, and Geastrum schmidelii
The key lead 7c was added after the original key was written – there are herbarium specimens with these three labels from the Pacific Northwest. G. schmidelii is smaller than the other two, with open diameter 1.4-3.7 cm (may be a dried measurement) and 4-8 rays. G. striatum (2.8-6.5 cm open diameter, 6-9 rays) has a basal collar. G. pectinatum (3-7.5 cm open diameter, 6-9 rays) does not have a basal collar. – Ian Gibson

8a	(6b)  Inner layer (upper) of rays thick and breaking transversely so as to form a cup around the spore sac	Geastrum triplex
Largest of the Geastrums, to 9 cm across opened rays. Gray brown to wood brown, spore sac pale to dark avellaneous, unpolished. Outer surface (under) rough but with little trash. Base marked by a distinct scar at point of attachment. Spore sac sessile in cup. Peristome 6 to 18 mm long, conic, surrounded by a paler ring, distinctly radially fibrillose, mouth lacerate / fibrillose. Gregarious to clustered, late summer and fall, mixed forests in Idaho and Oregon.

8b	Expanded rays not breaking to form a cup, surface remaining relatively smooth	Geastrum saccatum
To 5 cm across open rays. Sessile spore sac sitting in a bowl-like depression of the reflexed rays. Outer surface ochraceous buff to cinnamon buff; inner layer (upper) pallid to avellaneous when fresh, drying to near wood brown; outer surface holding little trash. Spore sac purplish drab to wood brown, unpolished. Inner surface smooth and continuous except rimose along edges. Peristome delimited by circular raised to depressed line, often paler or more brownish than spore sac wall, fibrillose. Gregarious on rich humus, often around old stumps in late summer to early fall, sometimes under junipers. Has been found in Cispus drainage, Copper Creek, Pierce Co., Washington, and McNeil Island in Washington. REMARKS G. fimbriatum Fr. is a widely distributed species that according to Bessette et al. (1997) is nearly identical to this species but with smaller round spores that measure 3-3.5 microns (instead of 3.5-5 microns) and an apical pore that is fimbriate and not delimited by a distinct circular paler zone. There are collections from BC at the University of British Columbia and it is listed for the Pacific Northwest by Helene Schalkwijk-Barendsen, but requires confirmation. – Ian Gibson

C.   DISCISEDA and ABSTOMA

Disciseda: Fruit bodies depressed - globose, peridium 2-layered; exo in form of a membrane or a sand case, fragile and deciduous in part so that lower portion of case remains and spore sac reminds one of a flattened acorn. Only one species in the Northwest.

Abstoma: According to Coker, Abstoma differs from Disciseda “essentially only in the absence of a definite mouth, both peridia wearing away from the attrition of wind-blown sand, etc.

Disciseda subterranea
To 2 cm across, 1 cm high. Cup is sand or soil color; spore sac unpolished bluish to glaucous gray over a dull ground color. Spore sac papery, thin, tough, opening by a special pore with elevated, entire to lobed margin. Exo deciduous to full diameter, exposing upper half of spore sac. On sandy soil in pastures; summer and fall; gregarious to solitary, mostly eastern North America but reported from Idaho.

Abstoma reticulatum
To 6 cm across, depressed spherical. Exo very thin or up to 0.2cm thick, fragile, fleeting, whitish or brownish, composed of hyphae intermixed with earth or sand, sometimes merely a meshy layer. Endo tough, membranous, dehiscing by irregular rupture, smooth, dark colored, mostly brown. Spore mass yellowish, then olivaceous with purplish tints, powdery, capillitium tinted. Under or on duff under conifers. There are collections from Idaho, California and Colorado.

Abstoma townei
To 6 cm across, 2-3.5cm high. Exo and Endo adherent and together forming a leathery spore case opening widely by radial tears across the upper part; surface smooth apart from small flecks and patches, brown. Spore mass with tints of yellow and purplish when young, dark sepia when mature. On soil in pastures and waste land. There collections from Idaho and Oregon. 

D.   BOVISTA

Surface at first covered with a thin, smooth to unpolished covering which flakes off at maturity, exposing the smooth polished wall of the spore sac, opening at times by a pore enlarging to a rupture, or developing several openings; becoming detached from the ground and blown about. Sterile base lacking. Four species in the Northwest.

1a	Attached to soil by small patch of fibers	2
1b	Attached to soil by a single small cord-like rhizomorph which breaks at maturity	3

2a	Mature spore case bluish gray to purplish umber; exo peeling in sheets	B. plumbea
1 to 3 cm across; depressed globose to globose; attached to soil by drab patch of fibers. White outer layer peels off in sheets to expose spore case of bluish gray to purplish umber. Spore case thin, tough, persistent; sometimes becoming detached and blown about. On turfed areas in summer and fall, throughout Northwest. Solitary to few clustered.

2b	Mature spore case red-brown to bronze brown. Exo peeling in small plaques	B. leucoderma
To 2 cm broad, spherical or somewhat depressed base attached to substrate without conspicuous mycelial strands. Exo smooth, later indistinctly areolate, finally peeling off from endo in small plaques from the base upward; pure white. Endo papery, red-brown to bronze-brown, dull to slightly glistening. Opening 1 to 3 mm with star-shaped flaps. Spore powder olive brown to umber brown.

3a	(1b)  Mature spore case copper color	B. pila
3 to 9 cm diameter; globose to subglobose. Attached to ground by cord-like rhizomorph which is easily broken. Color of mature spore case bronze metallic or coppery sheen. Outer layer white, thin, flaking off to expose spore case which is thin, papery, tough and persistent. Opening at apex by an irregular fissure or pore which becomes enlarged and torn. Spore case may become dislodged and blown about. Solitary, scattered to gregarious on turfed areas, stables, corrals, feedlots, pastures; in late summer and fall. Widely distributed.

3b	Mature spore case leaden to smoky gray	B. nigrescens
3 to 6 cm across; globose to subglobose; attached to soil by cordlike rhizomorph. Color white at first, exo thick, fragile, smooth, fleshy; breaking up as fungus matures. Spore case thin, tough, white at first, turning leaden or smoky gray. Opens at top with relatively large jagged apical pore 1/3 to 1/2 diameter of body. On turfed areas in summer and fall, gregarious to solitary in dry places. Well known in Europe, reported from Montana, needs verification.

E.   LYCOPERDON

Fruiting body typically globose to pyriform; peridium composed of two layers, the outer one consisting of spines, warts, granular or felty floccose material which usually sloughs off at maturity; inner wall forming a papery, thin but tough spore case; typically opening at the apex by a pore but occasionally by irregular fissures. Sterile base prominent in most species.

1a	Sterile base present and chambered (lacunae)	4
1b	Sterile base absent or, if present, not chambered	2

2a	Sterile base present, context felty without chambers	L. ericetorum
(also known as L. cepaeforme)
1 to 3.5 cm broad, 2-4.5 cm high, turbinate to pyriform, attached by rhizomorph quite strongly. Whitish to light brown at maturity; surface covered by a sparse coating of short spines and minute warts and granules; becoming glabrous, apical pore round. Gleba olive-brown when mature, cottony; sterile base well developed, 1/3 of fruit body, yellow-brown, felty with no chambers. First reported specimen found Dec. 1980 at Ocean Shores, Washington.
2b	Sterile base absent	3

3a	Exo closely covered with white spines, appearing furry, later peeling away from endo	L. curtisii
(also known as Vascellum curtisii)
1 to 2 cm broad, less high; depressed globose to pulvinate, plicate below; without conspicuous rhizomorphs; exo white; cracking and peeling away in chunks as in L. marginatum, revealing smooth endo which is ultimately medium brown and opens by a small pore. Sterile base absent. Gregarious in grass (Leavenworth, Washington), later summer and fall.

3b	Exo scurfy, with small warts and granules, some falling off, some drying and remaining, and not separating from endo	L. pusillum
(also known as Bovista californica)
1 to 3.5 cm broad, 1 to 3 cm high; depressed globose, somewhat pointed at base, holding small mass of rhizoids and soil. White to grayish white becoming red-brown to dull brown with obscure spotting at maturity; first smooth then breaking into areolae from center of top out and down. Spore sac relatively smooth but retaining persistent remains of exo as minute particles seen under a hand lens. Pore crenulate. Gleba olive brown; powder olive brown. Sterile base none. On pastures and idle lands; summer and fall; densely gregarious to clustered. Rare in Oregon; two colonies at Snake Lake Nature Center in Tacoma Washington, Aug. 1979. REMARKS It is unclear what is being described. L. pusillum of some authors is L. dermoxanthum, and both L. pusillum of other authors and Bovista californica may be different. – Ian Gibson

4a	(1a)  Exoperidium peeling away from spore case	5
4b	Exoperidium not peeling away from spore case	6
5a	Fruiting body stipitate / capitate	L. nettyana
Gasterocarp 2 to 3 cm broad, 3 to 4 cm high, stipitate-capitate with upper portion hemispherical, medium brown, covered with small, (more or less) regular granular patches that impart light grayish brown color; granular exoperidium interspersed with short conical spines and rounded buttons about 2 mm apart. Exoperidium at maturity splitting irregularly across the apex and peeling or flaking away. Endoperidium, when exposed, lustrous, smooth, light grayish yellowish brown, persistent; apical pore soon forming, circular, less than 1 cm diam. Pseudostipe tapering to base, attached to substrate by white rhizomorphs; surface pallid with scattered dark granules and reduced gland-like projections.
     Subgleba whitish, pigmented in uppermost 5 mm, with empty loculae less than 1 mm across; distinctly separated from gleba by a thin layer of collapsed loculae which is central elevated about 2 mm at the base of the pseudocolumella. Gleba grayish brown to moderate brown near or at maturity, with a distinct pseudocolumella rising to near the apex.
     Collected on soil in woodland clearings, scattered to cespitose, autumn. Cispus watershed, Lewis County, Washington.

5b	Fruiting body depressed globose	L. marginatum
1 to 5 cm broad, less high; depressed globose, plicate below with occasional projection downward. Exo white; cracking and flaking off at maturity in large chunks to reveal a spore sac which is smooth to obscurely pitted and opens by a small apical pore. Sterile base well-developed with chambers 1 mm across. On sandy soil; summer and fall; scattered to numerous, even cespitose. Idaho and western Montana.

6a	(4b)  Growing on wood or wood debris	Apioperdon pyriforme
	(also known as Lycoperdon pyriforme)
1.5 to 3 cm broad; 2 to 3 cm high; pyriform to subglobose; may be plicate at juncture of enlarged portion with stipe-like base. Connected to substrate by numerous white rhizomorphs. Pallid to tawny brown immature, darker rusty brown at maturity -- some yellowish; areolate patches darker. Exo breaking into areolate patches that divide into smaller units, which on drying form minute granules. At times exo over apex consists of small spines and granules. Remains of exo relatively persistent and rough to touch. Apical pore slow to form, often irregular. Sterile base slight to prominent depending on shape. On wood or sawdust on ground, cespitose to scattered, September to heavy frost. Old cases persist to next summer. REMARKS Morganella subincarnata (Peck) Kreisel and Dring (= Lycoperdon subincarnatum Peck) also grows on wood but according to Arora has cinnamon-buff to brown spines when young, conspicuous pits at maturity (after spines have worn off), and sterile base that is well-developed to practically absent. It is found mainly in eastern North America but has been reported from BC: confirmation is needed. – Ian Gibson

6b	Growing terrestrially	7

7a	Opening considerably larger than a pore, with stellate lobes or uneven margins, widening almost full breadth of body which is mostly turbinate or urn shaped, as broad as high	See VASCELLUM
7b	Opening consisting of classic pore less than 1 cm across, rarely more; fruit bodies mostly stipitate	8

8a	Exo spines lavender	L. peckii
1.5 to 4 cm across; to 5 cm high; obovoid to turbinate, sterile base often stipe-like (shape same as L. perlatum). Spiny coat is purplish lavender to near maturity then changing to buffy brown. Background and lower part white until near maturity. Covered with tapering spines to 1.5 mm long which split at base into 2-4 parts and remain joined at tips. Easily rubbed off, leaving pale, smooth, circular spots surrounded by minute granular warts and dots. Opening by apical pore. Sterile base occupying 1/2 of body. Terrestrial, fall, normally eastern species, rare in Northwest. Needs confirmation.

8b	Exo blackish, brown, gray or pallid	9

9a	Exo fuscous to black	L. foetidum
(also known as Lycoperdon nigrescens)
1 to 3 cm broad, 2 to 5 cm high; pyriform to globose. Spines fuscous to black, background yellowish; very fine and pointed, densely spaced over upper part, surrounded by granular material; not cone-shaped but more polygonal, with tips becoming dried, fine and curled. Spore sac grayish tan, smooth, papery, opening by apical pore. Sterile base well developed with obvious chambers. On duff and debris in dense conifer forests, Pacific Coast. At least 3 varieties. Common at times.

9b	Exo brown, gray or pallid	10

10a	Exo across top consisting of fine spicules which degrade to a mealy covering over the margin and down the stipe	11
10b	Exo with stout spines to 6 mm long, at least over top	12

11a	Exo spicules and granules soon wearing away; fruit body large (to 12 cm high); capitate; rhizomorphs large and strong	L. floccosum
Fruiting body stipitate / capitate to pyriform. In the specimen described: head to 5 cm broad, stipe below head 2 cm, stipe above base compressed laterally, 3.5 cm by 5.5 cm across; height to 12 cm. Sulcate below head and perhaps at base. Rhizomorphs large and strong. Whitish, with surface over top covered with very small hairy projections less than 0.5 mm long, degrading to a mealy covering over the margin and down onto the stipe. Peridium 0.5 to 1 mm thick in fresh specimens most of which is exo; endo very thin; the two layers not separating on maturing and drying. Spicules and mealiness of head soon wearing away. Sterile base well-developed, lacunae to 1 mm, subgleba extending up into head convexly. Dark gray-brown.
     Sandy soil along river bank, under trees, one cluster of several sporocarps. Federation Forest on White River, Pierce County, Washington, 1980. Only report in Northwest to date.

11b	Exo spicules and granules persistent, reducing on drying; fruit body rarely over 6 cm high; non-capitate; rhizomorphs present but small	L. umbrinum
2.5 to 5 cm broad, 3 to 8 cm high, base 1 to 4 cm across; pyriform, tapering or sometimes almost globose; often sulcate to plicate below. White to yellowish; spore sac yellowish to dull brown in age. Exo with scurfy or granulose coating mixed with very slender short spines which are relatively persistent or finally breaking up into concentric zones on base or areolate patches over the top. Eventually disappearing to leave smooth spore sac with apical pore. Gleba medium brown to deep brown showing evidence of pseudocolumella, cottony, confined to upper third of interior. Sterile base occupying entire tapered base and extending up sides to widest part. Violaceous medium gray; lacunae open, irregular, less than 1 mm; fusing into fertile tissue without demarcation. Solitary to gregarious or subcespitose on leaf mold under conifers, occasionally on wood; Sep. to Nov.; variable; W. Montana and Oregon and W. Washington.

12a	(10b)  Some spines joined at tips; spines fall away from spore sac but remain on stipe	L. pulcherrimum
2 to 5 cm broad, 3 to 4.5 cm high; pyriform to sub-globose over a narrowed, often plicate base; attached by white rhizomorphs. Exo white, spore sac dark purple brown. Exo consists of dense coating of spines 4-6 mm long, slender, fascicled, joined at the tips. Exo falls away at maturity to expose smooth papery tough spore sac, opening by apical pore which enlarges. "Stipe" remaining covered with spines. Well-developed sterile base turning purple brown in age. On humus under hardwoods in fall. Reported from Washington and W. Montana. Needs confirmation.

12b	Spines solitary, larger ones falling away readily, leaving round white scars; stipe without spines	L. perlatum
(also known as L. gemmatum)
2.5 to 6 cm broad, 3 to 7 cm high on stipe-like sterile base half as wide; typically obovoid, turbinate or pyriform, top narrowed abruptly to "stipe", often sulcate to plicate from widest point down onto base. White with spines brown in age before dropping off. A variety west of Cascades with brown spines in immature stage. Another variety is gray overall. Spore sac yellowish to dark avellaneous. Exo consists of sharp soft spines with round bases (cones) between which appear smaller cones, warts or granules. Larger cones deciduous, leaving pale round spots, more so on top. Spore sac becoming almost smooth, opening with apical pore. Sterile base well-developed, with chambers 1 mm across, chocolate brown in age. Solitary to cespitose on humus, often in arcs or fairy rings; in conifer or hardwood forests or in pastures or other turf; Aug. to late Oct.

F.   MYCENASTRUM

Only one species in the Northwest; widely distributed but not common except Rocky Mountain states.

Mycenastrum corium
6 to 15 cm (sometimes 20 cm) in diameter, 5 to 12 cm high; globose with subsessile "stipe" (like light bulb), at the end of which is a fibrous mycelial area of attachment. White changing to deep brown as exo sloughs off, sometimes purplish brown. Exo cracks into block-like plates, falling away to reveal smooth spore case which is hard and about 2 mm thick. Opens at apex by irregular fissures, becoming extended toward base. Segments curve back, giving appearance of giant Geastrum at times. Interior becomes a powdery or flocculent mass. No sterile base. Gregarious to solitary on soil where cattle are fed, in pastures; summer and fall; at maturity breaks loose from soil and blows about.

G.   VASCELLUM

Two species in the Northwest. The first species common along the Northwest coast on lawns, golf courses and other turfed areas. Most often dismissed as a Lycoperdon.

1a	Peridium opening across top longer than wide by simultaneous decomposition of exo and endoperidium, edge of opening ragged, less than full diameter of body; separation between gleba and sterile base distinct but not a separable membrane	V. lloydianum
2 to 5 cm broad and as high; turbinate to pyriform. Exo white to pallid, endo medium shiny brown; exo with mixture of soft spines and granules, both deciduous. Opening by simultaneous decomposition of exo and endo across the top, longer than wide and ragged along its edge. Gleba dark brown, becoming powdery. Sterile base in lower constricted portion, lacunae less than 1 mm, extending somewhat up sides; separation of sterile base from fertile tissue distinct but inseparable layer of flattened lacunae. W. Washington.

1b	Peridium opening across top to widest diameter by stellate lobes; separation between gleba and sterile base by distinct parchment-like membrane	V. depressum
(also known as Lycoperdon depressum)
2 to 4 cm broad, 2.5 to 5 cm high; turbinate, bowl or urn-shaped. Exo white to pale buff, endo shiny metallic brown. Exo with soft spines and granules, both deciduous. Opening by stellate lobes across the top, later enlarging and breaking away until lower half persists to form a bowl. Sterile base present and separated from gleba by separable papery to parchment-like membrane, chambers large. Montlake fill, University of Washington campus. REMARKS Vascellum concepts have been the subject of some debate. Both V. depressum and V. lloydianum are considered by some to be synonyms of Vascellum pratense (Pers.: Pers.) Kreisel. – Ian Gibson

H.   CALBOVISTA

Only one species in the Northwest. Generally found above 3,000 foot elevation, occasionally in alpine areas of disintegrated rock and soil mix. Idaho pastures down to 2,000 feet.

Calbovista subsculpta
8 to 16 cm broad, to 9 cm high; globose to pear shaped, often plicate and contracted toward a rooting base. Whitish to dingy, tips of scales grayish. Exo thick, felty, leathery, broken into irregular 3 to 6-sided, low truncated pyramids, usually blunt but sometimes pointed. Pyramids 5-8 mm high at top of body but thinner toward bottom. Sides of pyramids show parallel transverse lines. Sterile base 1/4 to 1/3 of gleba. Single to gregarious in subalpine open areas near forest edge; April to August.

I.   CALVATIA

Typical puffballs; peridial wall consisting of two layers which are adherent in some species and separating in others; sometimes the outer layer not easily seen or understood. Depending on the species, may have an interior sterile base below the spore mass (gleba) or may lack one. Forms are generally globose to depressed, many with bases extending downward. Nineteen species in the Northwest, several apparently rare.

1a	Sterile base lacking or rudimentary	2
1b	Sterile base present, although in a few species it becomes less evident at maturity	10

2a	Fruiting body usually over 10 cm broad	3
2b	Fruiting body usually not over 10 cm broad	4

3a	Globose to subglobose; exo thin and fragile; found in brushy or grassy bottomlands	C. gigantea
20 to 50 cm diameter (up to 12 kilograms in Idaho); globose to depressed globose. White to grayish, changing to yellowish to olivaceous. Exo smooth like kidskin glove, ultimately breaking up and falling away from inner layer in pieces. Endo thin and fragile, breaking up after maturity. Gleba white to yellowish to olivaceous. Solitary to gregarious on low ground under brush. East of Cascades in Washington, Oregon, and Idaho in late summer and fall.

3b	Depressed globose; exo thick and breaking up in raised polygons; found in arid lands under sagebrush and juniper	C. booniana
To 60 cm across, 7 to 30 cm high; depressed globose but more or less circular from above. White to dingy pale brown. Exo thick and white, subfloccose, breaking up into 4-6 sided areolate polygons to 2 cm deep, falling away from inner layer which is whitish then buff, with slightly punky texture, unpolished and felt-like. Gleba dingy olive when mature. On soil under sagebrush and juniper, in summer in arid regions of Idaho, Oregon and Nevada. Also Pend Oreille Valley, Washington.

4a	(2b)  Spore mass cottony	5
4b	Spore mass powdery	6

5a	Exo similar to C. gigantea, scaling off to reveal pliant endoperidium; opens by radial tears from top to near bottom	C. lacerata
About 7 cm diameter, globose. White to pallid changing to brownish. Exo present as a thin felty layer, sloughing off in patches, leaving a thin, pliant, pallid to brownish inner layer. Gleba very dark brown at maturity. Found only near Rhododendron, Oregon, in young Douglas-fir area. Solitary in fall.

5b	Exo adheres to endo but cracks into small polygons; opens by endo wall cracking into large plate-like areas	C. lycoperdoides
2 to 5 cm broad, 1.5 to 4 cm high; globose to pulvinate. Whitish, changing to brown. Exo delicately furfuraceous to spinulose over all, the upper surface often breaking into small areolae becoming raised polygons up to 3 mm across, low and rounded: occasionally tips pointed and surface then spinose. Warted but furfuraceous to spinulose below and not areolate. Opening occurring by wall cracking into large plate-like areas; spines deciduous, leaving a thin tan endo. Similar to Lycoperdon but not opening by a pore. Gleba dark olivaceous brown to umber and purplish at base. In coniferous woods in late summer and fall in Washington and Idaho. REMARKS Calvatia lycoperdoides A.H. Sm. is an illegitimate name because the homonym Calvatia lycoperdoides Koscielny and Wojt was published earlier. – Ian Gibson

6a	(4b)  Peridial wall hard at maturity	7
6b	Peridial wall soft at maturity	9

7a	Peridial layers separating; surface roughened with warts	C. subcretacea
2 to 7 cm broad, 1.5 to 4 cm high; depressed globose to pulvinate, base tapering to a point when young but lobed and broad to almost flat when mature; attached by white mycelium. Chalky white with polygonal warts smoky gray. Warts up to 7 mm across, lined horizontally or vertically. Endo thin, firm, furfuraceous, dull chamois to honey yellow, breaking irregularly above. Gleba white to buff to burnt umber. Scattered to solitary, under spruce / fir late summer and fall; Oregon, Idaho and Washington. REMARKS C. subcretacea  is now usually known as Lycoperdon subcretaceum or Gastropila subcretacea. – Ian Gibson

7b	Peridial layers adherent; surface more or less smooth	8

8a	Sporocarp malodorous, hard and persistent	C. fumosa
3 to 10 cm broad, less high; subglobose; with cordlike rhizomorph; smooth to slightly furfuraceous, somewhat wrinkled in drying, rarely lightly areolate. Whitish but soon smoke gray to umber brown where exposed to light but chalky white under the surface. Peridia inseparable, 1 to 2 mm thick, hard and persistent, as Scleroderma. Gleba very dark brown and powdery. Odor as a combination of sour milk, diesel oil and pit toilet. Sterile base lacking to rudimentary. Idaho variety with yellow brown mature gleba. Solitary to gregarious in duff under spruce / fir forests in mountains. “Variety C.f.idahoensis in Hell's Canyon, Idaho.” [The original key underlined any italics.] REMARKS 1) C. fumosa  is now usually known as Gastropila fumosa. 2) Var. fumosa was examined from WA, OR, and ID by Zeller. 3) In transferring C. fumosa to Handkea fumosa, Kreisel rejected var. idahoensis on the basis of the microscopic description. – Ian Gibson 2019

8b	Sporocarp with fungoid odor, hard leathery	C. hesperia
4 to 8 cm broad, less high; depressed globose; smooth to roughened leathery surface; white to pearly gray. Peridia adherent, cracking and breaking up slowly to release spores. Gleba clay color to greenish yellow and finally chocolate; becoming powdery at maturity. Sterile base absent. Four collections in Idaho (2 Canyon County farmland and 2 Owyhee County desert country with sagebrush and cheatgrass). Farmland in June and August; desert in December and January. Formerly known only from California. REMARKS C. hesperia is now usually known as Gastropila hesperia. – Ian Gibson 2019

9a	(6b)  Opens by radial tears widely across the top; exo somewhat leathery, with glabrous to rimose surface	C. paradoxa
2 to 6 cm broad, 2 to 3.5 cm high; pulvinate to depressed globose. White to dull brown. Peridial layers adherent forming a somewhat leathery spore case opening widely by radial tears across the top. Empty case persistent. Gleba white then dark sepia but not purplish. Sterile base lacking or rudimentary. On soil in pastures or range lands in mountainous areas of Idaho.

9b	Opens by breaking of polygons which form evenly and smoothly across upper surface except each with a small central spine	C. owyheensis
5 cm broad and 3 cm high; depressed globose to pulvinate, with mycelioid rooting base. Lower surface pallid and furfuraceous; upper surface dingy pallid with patches stained brown. Peridial layers adherent, outer evenly areolate / warty and more or less polygonal and smooth, giving effect of an embossed surface except polygons have a small central wart or spine. Polygons neither large nor lined as in C. sculpta. Inner layer thin. Gleba white, eventually olive brown. Sterile base limited to a small area of chambers at the top of the rooting mycelium-and-soil column. Late spring, Owyhee County, Idaho.

10a	(1b)  Gleba becoming cottony	11
10b	Gleba becoming powdery	14

11a	Sterile base cellular, purplish brown or grayish brown	12
11b	Sterile base fine textured, neither purplish nor grayish brown	13

12a	Exo remaining granular to furfuraceous; sporocarp higher than broad, somewhat stipitate	C. elata
Stipe to 5 cm wide, head to 7 cm wide; the whole to 12 cm high. Exo whitish to pale brown to leather brown at maturity, remaining somewhat granular to furfuraceous. Endo thin, breaking into fragments. Gleba brown to olive-brown, becoming cottony during spore dispersal. Sterile base well-developed with fine-textured chambers not over 0.5 mm across. On low ground or stabilized sand dunes at the edge of wet areas. First located in Northwest in Nov. 1980 under Hooker Willows at Ocean Shores, Washington.

12b	Exo areolate with obtuse spines and prominent warts, not separating from endo; sporocarp wider than high, depressed globose to turbinate	C. lloydii
3 to 8 cm broad; depressed globe to subturbinate, rounded or flattened above, pinched below and attached by several white rhizomorphs. Creamy white changing to chamois to reddish brown. Peridial layers adherent, outer dull to silky areolate above, becoming increasingly areolate and forming more or less prominent warts or obtuse spines which become smaller and furfuraceous to smooth below. Opening irregular; peridia more or less persistent around sterile base. Gleba from light to dark olive buff to olive brown. Sterile base concave and forming shallow cup around gleba; small cells with purplish brown, metallic luster. Dry coniferous woods, mountains of Idaho.

13a	(11b)  Exo surface dull reddish brown; sterile base concave, forming shallow cup	C. ochrogleba
7 to 9 cm broad, 7 to 9 cm high; turbinate to depressed globose to misshapen. Dull reddish brown. Peridial layers adherent. Exo very thin, and smooth to furfuraceous. Endo medium thick and ochraceous. Gleba very fine texture, pale yellow-orange to ochraceous buffy. Known only from meadows near Aurora, Oregon.

13b	Exo surface drying to orange red or bay and finally metallic brown; sterile base deeply cupped	C. rubroflava
2 to 10 cm broad, 1.5 to 5 cm high; flattened above, subglobose, usually with short abruptly pointed base, centrally attached by one or more root-like rhizomorphs. White becoming pink or lavender to pinkish leather brown. All parts changing at once to chrome yellow-brown. Exo represented only by a delicate furfurescence, deciduous, revealing smooth, shiny endo, cracking into irregular areas or scales and falling away after drying. Sterile base homogeneous without cells and extending up sides as a thin layer; yellowish to chocolate brown. On cultivated soils in Oregon.

14a	(10b)  Gleba distinctly violaceous at maturity	15
14b	Gleba other colors at maturity	16

15a	Body higher than broad, substipitate to turbinate, persistent sterile base without membrane	C. cyathiformis
7 to 16 cm broad, 9 to 20 cm high; globose to pyriform, sulcate to deeply wrinkled from base to broadest dimension. Pallid becoming brownish to dingy purplish drab. Exo thin, smooth, becoming areolate over upper part, often developing flat scales. Endo thin, delicate, slowly scaling away with exo to expose gleba. Gleba white to yellow to purple drab. Sterile base chambered. On soil in grasslands in fall; eastern Oregon. REMARKS C. fragilis (or C. cyathiformis forma fragilis) is said by Zeller to be the common purple-spored Calvatia of western Oregon. (She noted C. cyathiformis, which also has purple spores, from the Willamette Valley in western Oregon as well as from eastern Oregon.) Arora says that it is very similar to C. cyathiformis, but has only a rudimentary sterile base. – Ian Gibson

15b	Body lower than broad, base not extending downward; persistent sterile base convex, with membrane	C. rubrotincta
4 to 7 cm broad, 3 to 4 cm high. Depressed globose, sulcate below, base with several cordlike rhizomorphs. White becoming stained red in spots to dark reddish umber. Exo very thin, evanescent or flaky, smooth, to felty below, areolate above by reticulations. Endo thicker, brittle, easily broken along ridged reticulations. Opens irregularly with fragments. Sterile base soft, cellular, to purplish brown with metallic luster. Under ponderosa pines, Lake County, Oregon. Rare.

16a	(14b)  Exo very thick, breaking into conspicuous pyramids to 3 cm high, separate or joined at tips, tips coiling as they dry	C. sculpta
4 to 10 cm across; obovate, turbinate or pyriform, base not radicating. White, tips of spines darkening as they dry. Exo very thick, breaking into pyramids up to 3.5 cm across at base and 3 cm high, often with longitudinal or parallel lines. In age pyramids may divide vertically with apex remaining intact; apices of earlier pyramids may be coiled. Segments eventually falling away separately, upper surface first. Endo thin, membranous, fragile, breaking up with exo along pyramid base lines, usually upper surface first, exposing gleba through irregular opening. Gleba white, turning yellow then olive brown. Sterile base chambered, often purplish in base. On duff in high conifer forests, late summer and fall, Washington, Oregon, and Idaho.

16b	Exo variously formed and marked but none so conspicuous as C. sculpta	17

17a	Base constricted and plicate	18
17b	Base smooth around perimeter	19

18a	Exo thick, cracked into low, irregular plates and falling away from endo which later opens irregularly with a wide tattered mouth	C. tatrensis
4.5 to 9 cm broad, 5 to 7 cm high (var. gruberi 13 cm or more broad); turbinate to obovate, strongly furrowed or plicate below (var. gruberi lacks plication). White to cream. Exo thick, cracking into low irregular plaques which are granular-furfuraceous and mostly deciduous, remnants small floccose patches or large plate-like areas. Endo chamois color to dull tawny or brown-stained from gleba; firm to somewhat papery in age. Peridial wall more or less persistent around the margin of the sterile base. (Var. gruberi exo is spiny on upper surface.) Gleba chocolate brown to dark olivaceous brown at maturity. Sterile base chambered and chamber walls purplish to olivaceous in age. Range lands, under sagebrush in late summer. (Var. gruberi near Mt. Hood, Oregon under conifers.)

18b	Exo thin, wrinkled and rivulose, adherent to endo and falling away together to open body irregularly	C. pallida
1.5 to 5.5 cm broad, 1 to 2.5 cm high; flattened, pulvinate or turbinate, base constricted and plicate. Pallid to pale pinkish buff overall or dusted umbrinous from spores. Exo upper surface finely but distinctly wrinkled, occasionally finely rivulose between the wrinkled areas. Two layers adherent, rather thin (less than 1 mm), brittle. Endo, when discernible, thin and papery. Gleba white to olivaceous brown (no orange stage as in C. candida) to olive brown with a purplish tinge when mature. Sterile base poorly developed (1/4 to 1/8 of body), of small cells becoming less evident by maturity. Open woods and meadows in duff and soil at high elevations, summer and fall, Washington and Idaho.

19a	(17b)  Large (to 15 cm), upper surface a thick floccose layer areolate with truncated polygons; sterile base a cup with sides tapering upward a short distance, chambers distinct	C. bovista
5 to 15 cm wide; subglobose to turbinate with large long-cylindric or short-tapered stem. Whitish to pallid yellowish to brownish. Exo a thick floccose layer, usually areolate with truncate polygons above, thinner below, with tendency to form stellate flattish areas. Endo thin, breaking up into fragments at the apex, then downward to expose the powdery gleba. Gleba bright olivaceous to brownish olive to dark olive brown. Sterile base 5 to 10 cm high, furrowed below and pinched to a point. Fields, lawns and roadsides in Idaho. REMARKS This species goes by a variety of names including Bovistella utriformis and Calvatia utriformis – Ian Gibson.

19b	Smaller (to 6 cm), upper surface areolate and plaque-like but not elevated; sterile base less evident at maturity, chambers small	C. candida
1.7 to 6 cm broad, 1.5 to 6 cm high; pulvinate to turbinate. White to chalky white or slightly grayish over upper side. Exo smooth to wrinkled or areolate; areolae plaque-like and not raised up as polygons or squamules; the under side finely granular to furfuraceous. Endo opening by breaking into large plate-like areas or simply peeling away from the gleba. Peridia thin to thick, the firm exo up to 1 mm, and the endo papery and velum-like. Gleba white becoming orange but finally olivaceous brown to umbrinous. Sterile base occupying about a quarter of body. Arid places under sagebrush and range grasses; late summer and fall in Idaho.

GLOSSARY

areolate - having a pattern of “block-like” areas similar to those formed when a mudflat dries. However, such a pattern may be formed without cracks when the “block-like” areas are separated by spreading and stretching the underlying tissue which then assumes a different color from the blocks.
avellaneous - dull grayish brown, hazel-brown, or light gray yellow brown, or gray or buff tinged with pink.
bay - red-brown approaching but lighter than chestnut.
capillitium - the mass of sterile thread-like elements which, with the spores, forms the gleba of Lycoperdaceae. A single strand of capillitium, under the microscope, may reveal distinctive characteristics of size, shape, branching ends, and ornamentation.
capitate - having an expanded head-like structure atop a stem. The gleba is located in the upper portion and the sterile base in the lower portion.
cespitose - also caespitose: growing in close proximity so as to be very crowded yet remaining mostly separate. 
chambered - cellular, lacunose, loculate: bearing pits, holes and/or small cavities, pertaining to a sterile base in which the context consists of empty “cells” surrounded by thin walls of tissue, visible to the unaided eye or by hand lens, for those species with cellular sterile bases.
chamois - a pale yellow.
clay color - resembling dull ochraceous-cinnamon-brown.
columella - sterile tissue that penetrates the gleba vertically and plainly and is enclosed by the peridia. A pseudocolumella shows a fainter but similar pattern but will contain fertile tissue.
coriaceous - like leather in texture.
cortex - an outer covering.
crenulate - an edge with tiny rounded teeth.
dehiscence - The method by which a structure opens to release its contents; in this case, the way in which a fruiting body opens to discharge spores.
drab - a dull medium or brownish gray, dark gray with shades of yellow; gray with violet overtones.
duplex - of two distinct textures.
endoperidium (endo) - the inner layer of tissue surrounding the gleba.
entire - with even edges.
evanescent - soon disappearing, fleeting.
exoperidium (exo) - the outer layer(s) of tissue surrounding the gleba.
fascicle - a little bundle. A group or more-or-less slender structures are bound together at the base or at the tips.
fibrillose - with delicate fibers which are long and evenly disposed, loosely or tightly, on the outer surface.
fimbriate - fringed with hairs or fibrils along or around the edge of a structure. The peristomes of some Geastrums may be so ornamented.
fissure - a narrow opening, cleft or crevice, implying it is split all the way through, smooth or jagged.
floccose - with easily removed cottony or woolly tufts; woolly or cottony.
flocculent - woolly or cottony.
fornicate - rays raise spore case as if on stilts still attached at their tips to a nest-like remnant in the ground formed from the outermost layer of the rays, with the fibrous and fleshy layers splitting from and becoming arched over the mycelial layer.
furfuraceous - scurfy, surface covered with branlike particles resembling scales, coarser than granular.
furfurescence - a furfuraceous covering.
fuscous - the color of a very dark storm cloud; dark gray to blackish.
fulvous - fox-colored, deep orange to reddish orange, reddish cinnamon brown.
gasterocarp - sporocarp or fruiting body of a gasteromycete.
gasteromycete - a basidiomycete that does not actively discharge its spores.
glabrous - bald.
glaucous - sea-gray (as in this case); sea-blue-green; covered with white bloom.
gleba - the whole interior mass of spores, capillitium, etc., lying above any sterile base within a peridium.
lacuna (plural lacunae) - cavity, in this case in the sterile base.
locula (plural loculae) - cavity, in this case in the sterile base.
lacerate - peridia appearing torn - a method of dehiscence.
pallid - scarcely colored but not white: may be used with any hue, e.g. pallid brown, pink or yellow.
pedicel - of spore case, a stalk.
peridium - the inner and outer covering of a puffball.
peristome - special pore structure formed by some Geastrums to release spores.
plicate - the folding of tissues underneath a sporocarp where the shape changes from spherical to a projecting point.
pore - an opening confined to the upper center of a sporocarp in which a more-or-less distinct hole results: may be ragged.
pseudorhiza - a long rootlike extension of the lower stem.
pulvinate - top surface convex but lower than spherical; like a cushion but not flat.
pyriform - pear-shaped, with stem end down.
radicating - forming a root.
rays - those portions of the exoperidium resulting from the splitting of the exoperidium down to 1/2 to 3/4 of its height and reflexing open to leave the spore sac exposed.
recurved - curved back.
reflexed - turned back.
reticulation - a network of interlacing lines or ridges.
rhizoid - root-like structure from the base of a fruiting body.
rhizomorph - a thread-like or cord-like structure composed of hyphae tying puffball to the substrate while maturing and conveying water and nutrients to it as well.
rimose - cracked, referring to surface of cap or stem.
rivulose - arranged like rivulets in the soil, marked with riverlike lines.
rudimentary - underdeveloped, not mature.
scurfy - surface covered with branlike particles resembling scales, same as furfuraceous.
sepia - a moderate brown; a brownish gray to a dark olive-brown.
sessile - lacking a stem; directly attached to substrate or another structure.
spinose - with spines.
spinulose - with fine spines.
spore case (spore sac) - structure holding spore mass together until dehiscence: usually the endoperidium alone as in Geastrums.
sporocarp - a structure in which or on which spores are produced, often used for fruiting body.
stellate - star-shaped.
sterile base (subgleba) - the part of the sporocarp in which spore formation does not take place: may be felty, cellular, small, large, fused, or delimited.
stipitate - having a stem or stipe or a stem-like column of tissue on which the head or fertile structure sits.
sub-   - nearly, somewhat, more or less; under
substrate - the material on which a fruitbody is growing.
sulcate - condition on underside of sporocarp similar to plicate. Instead of appearing folded however, the extended base is furrowed or grooved longitudinally: may extend from equator to base point.
turbinate - shaped like a top or turnip.
umber - a deep dull dark brown, smoky brown; earth brown sometimes with a very slight reddish tinge.

ACKNOWLEDGEMENTS

We wish to acknowledge use of information from the works of other investigators, especially two books on non-gilled fungi by Dr. Alexander H. Smith. However, much valuable information was gained from Dr. Daniel Stuntz, Ellen Trueblood, Kit Scates and from those who brought in specimens from time to time to help in the work. An amateur mycologist needs all the help he can get.

SELECTED REFERENCES

Arora, David. 1986. Mushrooms Demystified Second Edition. Ten Speed Press, Berkeley.  
Bates, Scott Thomas. 2004. Arizona Members of the Geastraceae and Lycoperdaceae (Basidiomycota, Fungi). MSc Thesis. Arizona State University. http://www.azfungi.org/stbates/thesis.html
Bessette, Alan E., Arleen R. Bessette, David W. Fischer. 1997. Mushrooms of Northeastern North America. Syracuse University Press. 
Bowerman, Constance A. 1961. “Lycoperdon in eastern Canada with special reference to the Ottawa district.” Can. J. Bot. 39: 353-383.
Coker, William Chambers, and Couch, John N. 1928. The Gasteromycetes of the Eastern United States and Canada. Dover Publications, New York.
Kreisel, Hanns. 1989. “Studies in the Calvatia complex (Basidiomycetes)”. Nova Hedwigia 48: 281-296.
Pegler, D.N., T. Læssøe, and B.M. Spooner. 1995. British Puffballs, Earthstars and Stinkhorns. Royal Botanic Gardens. Kew, England. 255p.
Smith, Alexander H. 1951. Puffballs and their allies in Michigan. University of Michigan.
Smith, Alexander H. 1974. “The genus Vascellum in the United States.” Travaux mycologiques dédié à R. Kühner, numéro spécial du Bulletin de la Société Linnéenne de Lyon, 15 février 1974. 
Smith, Alexander H., Smith Helen V., Weber, Nancy S. 1981. How to Know the Non-gilled Mushrooms. Second Edition. Wm. C. Brown Company, Dubuque, Iowa. 
Zeller, S.M. 1947. “More Notes on Gasteromycetes”. Mycologia 39: 282-312.
Zeller, Sandford M, and Alexander H. Smith. 1964. “The genus Calvatia in North America.” Lloydia 27(3): 148-186.

INDEX

ABSTOMA  G. Cunn.
	A. reticulatum  G. Cunn. 	C	7
	A. townei  (Lloyd) Zeller	C	7

APIOPERDON  (Kreisel & D. Krüger) Vizzini
	A. pyriforme  (Schaeff.) Vizzini 	E 6a	9

ASTRAEUS  Morgan 
	A. hygrometricus  (Pers.) Morgan	B 4a	5
	A. pteridis  (Shear) Zeller	B 4b	5

BOVISTA  Pers.
	B. leucoderma  Kreisel 	D 2b	7
	B. nigrescens  Pers.: Pers.	D 3b	8
	B. pila  Berk. & M.A. Curtis	D 3a	7
	B. plumbea  Pers: Pers.	D 2a	7

CALBOVISTA  Morse
	C. subsculpta  Morse	H	12

CALVATIA  Fr.
	C. booniana  A.H. Sm.	I 3b	12
	C. bovista  (Pers.) Kambly & Lee	I 19a	16
	C. candida  (Rostr.) Hollos	I 19b	16
	C. cyathiformis  (Bosc) Morgan	I 15a	15
	C. elata  (Massee) Morgan	I 12a	14
	C. fragilis  (Vitt.) Morgan	I 15a	15
  = C. cyathiformis forma fragilis
	C. fumosa  Zeller	I 8a	13
	C. fumosa var. idahoensis  A.H. Sm.	I 8a	13
	C. gigantea  (Batsch: Pers.) Lloyd	I 3a	12
	C. hesperia  Morgan	I 8b	13
	C. lacerata  A.H. Sm.	I 5a	13
	C. lloydii  Zeller & Coker	I 12b	14
	C. lycoperdoides  A.H. Sm.	I 5b	13
	C. ochrogleba  Zeller	I 13a	15
	C. owyheensis  A.H. Sm.	I 9b	14
	C. pallida  A.H. Sm.	I 18b	16
	C. paradoxa  A.H. Sm.	I 9a	14
	C. rubroflava  (Cragin.) Morgan	I 13b	15
	C. rubrotincta  Zeller	I 15b	15
	C. sculpta  (Harkn.) Lloyd	I 16a	15
	C. subcretacea  Zeller	I 7a	13
	C. tatrensis  Hollos	I 18a	16
	C. tatrensis var. gruberi	I 18a	16

DISCISEDA  Czern.
	D. subterranea  (Peck) Coker & Couch	C	7

GEASTRUM  Pers.
	G. campestre  Morgan	B 3b	5
	G. corollinum  (Batsch) Hollós  See G. recolligens.
	G. fimbriatum Fr.	B 8b	6
	G. floriforme  Vittad.	B 3a	4
	G. limbatum  Jungh.	B 7a	5
	G. pectinatum  Pers.	B 7c	6
	G. quadrifidum  Pers.	B 7b	5
	G. recolligens  (Woodw. ex Sowerby) Desv.	B 3b	4
	G. rufescens  Pers.	B 5a	5
	G. saccatum  Fr.	B 8b	6
	G. schmidelii  Vittad..	B 7c	6
	G. striatum  Quél..	B 7c	6
	G. triplex  Jungh.	B 8a	6
	G. umbilicatum  Fr. sensu Morgan	B 3b	5

LYCOPERDON  Pers.
	L. curtisii  Berkeley	E 3a	8
  = Vascellum curtisii  A.H. Sm.
	L. ericetorum Pers. var. cepaeforme  (Pers.) Bowerman	E 2a	8
	L. floccosum  Lloyd	E 11a	10
	L. foetidum  Bonord.	E 9a	10
  = L. nigrescens
	L. gemmatum   See L. perlatum.
	L. marginatum  Vittad.	E 5b	9
	L. nettyana R.W. Ramsey	E 5a	9
	L. nigrescens   See L. foetidum.
	L. peckii  Morgan	E 8a	9
	L. perlatum  Pers.: Pers.	E 12b	11
  = L. gemmatum
	L. pulcherrimum  Berk. & M.A. Curtis	E 12a	11
	L. pusillum  (Pers.) Schwein.	E 3b	8
	L. pyriforme  Schaeff.: Pers.	E 6a	9
	L. subincarnatum  Peck	E 6a	9
	L. umbrinum  Pers.	E 11b	10

MORGANELLA  Zeller
	M. subincarnata (Peck) Kreisel.	E 6a	9

MYCENASTRUM  Desv.
	M. corium  (Guer.) Desv.	F	11
	
MYRIOSTOMA  Desv.
	M. coliforme  (Pers.) Corda  See Key to Earthballs (Sclerodermataceae).

SPHAEROBOLUS  Tode
S. stellatus  (Tode) Pers.  See Key to Bird’s Nests (Geastraceae).

VASCELLUM  F. Smarda
	V. curtisii  A.H. Sm.  See Lycoperdon curtisii.
	V. depressum  (Bon) F. Smarda	G 1b	11
	V. lloydianum  A.H. Sm.	G 1a	11
	V. pratense  (Pers.: Pers.) Kreisel	G 1b	11

-  END  -

LYCOPERDACEAE & GEASTRACEAE  -    -