Steph Jarvis’s Description of Lycoperdon perlatum Pers.

Title: Monograph Of The Lycoperdaceae Of California By Steph Jarvis (Public)
Name: Lycoperdon perlatum Pers.
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 Monograph Of The Lycoperdaceae Of California By Steph Jarvis (Public)

Description status: Unreviewed
 (Latest review: 2019-10-31 11:01:30 CDT (-0400) by jason)

General Description:

Lycoperdon perlatum Pers., Observ. Mycol. (Lipsiae) 1: 4. 1796.
FIGURE 30, 41, 70
Reported synonyms:
   = Lycoperdon lacunosum Bull., Herb. Fr. 2: tab. 52. 1782.
   = Lycoperdon gemmatum Batsch, Elench. fung., cont. prim. (Halle): 147. 1783.
   = Lycoperdon perlatum var. albidum Alb. & Schwein., Consp. fung. (Leipzig): 80. 1805.
   = Lycoperdon gemmatum var. perlatum (Pers.) Fr., Syst. mycol. (Lundae) 3(1): 37. 1829.
   = Lycoperdon perlatum var. lacunosum (Bull.) Rea, Brit. basidiomyc. (Cambridge): 34. 1922.
   = Lycoperdon perlatum var. bonordenii (Massee) Perdeck, Blumea 6: 505. 1950.

TYPE—None of the collections in European herbaria have material labeled as type. According to Kreisel (1962), the type location is in Europe. A neotype specimen has not been designated. Lycoperdon gemmatum Batsch, recognized as a later synonym of Lycoperdon perlatum Pers., is the type species of the genus Lycoperdon.

Diagnostic Description:

GASTEROCARP (11) 25-32 (55) mm tall x (7) 11-25 (70) mm broad, tapering to 5-10 (15) mm at the base; obpyriform; rhizomorph root-like, white cream colored and incrusted with soil and sand particles, up to 6-7 mm long, and up to 0.5 mm broad, thin and branching; ostiole slow to develop at the apex, opening via a round hole initially, developing into a round opening, or tearing linearly, fibers along the ostiole edge radiating inwards into the gleba, exposing the gleba. Exoperidium cream white when young (4A3-4A2), turning tawny brown, beige brown to olive brown green (5B3-4, 5E5-8, 6E4-7), sometimes tan mottled with chocolate brown (5D3) in coastal climates, or mottled with red brown beige (6D6) in elevated dry desert climates, pale around the base with darker shades around the ostiole and at the apex; exoperidium up to 4 mm thick; heavily ornamented with conical tapering pyramidal warts, some warts fused at the tips, 2-4 warts fusing, warts not typically fusing along the neck or base of the subgleba, some warts taper and bend at the tips with age, warts darkening in color with age; warts along the apex larger and with a collared attachment, warts smaller and more fragile along the margin and sides of the exoperidium; granular particles seen around the base of warts using a hand lens; warts and granular particles eventually sloughing off, leaving a circular reticulate scar-like pattern and exposing the endoperidium; irregular-shaped lateral cracks appearing with drying and desiccation, cracked portions caving in to reveal the endoperidium in old specimens. Endoperidium white when young, turning beige buff (4B3), to brownish orange to grayish brown (6C3-4, 6D3), beige grey yellow (6E3), metallic olive yellow (4F6) when mature; smooth, persistent and parchment-like, with a reticulate pattern or completely worn smooth with age and weathering. Gleba white and firm when young, cream to tan, becoming shades of yellow to green brown (4A4, 5B3, 5E4-6, 6E4), gleba spores yellow-brown to olive-brown to dark brown (5D5-6, 5E4-7) in deposit, color not uniform until mature; maturing from the center outward, elastic when young, becoming cottony and flexuous with age. Subgleba white when young, cream buff beige (4A3-4B3), to grey (5E2), slightly darker brown in the center when mature, remaining cream around the outer perimeter; with a prominent pseudostipe, medium to large in size, composed of compact chambers, comprising lower 1/4-1/2 of fruitbody. Diaphragm absent.

BASIDIOSPORES globose; (2.5) 3.2-4.8 X (2.5) 3.2-4.8 µm [xmr = 4.1-4.5 X 3.8-4.6 µm, xmm = 4.3 ± 0.1 X 4.2 ± 0.4 µm, Q = 0.8-1.4, Qmr = 1.0-1.1, Qmm = 1.0. ± 0.1, n = 25, s = 5]; spores golden brown in wet mounts, spores lightly pigmented tan brown in KOH mounts; spores echinulate with prominent ornamentation easily seen at 40x with a light microscope, under SEM spores have well-spaced echinulate pyramidal warts; oil drop present; spores thick-walled; spores with pedicel up to 0.8-1.8 μm long, and up to 1 μm broad, thick-walled, often broken off with a clean edge seen in SEM; free-floating sterigmata remnants present to scarce in young to mid-mature fruitbodies, absent in very young and very old specimens, up to 22 μm long; spores of equal size in light microscope. Eucapillitium Lycoperdon-type; threads up to 3-7.5 μm broad, walls < 0.5 μm thick; yellow to golden brown in wet mounts; subelastic, breaks when teased, to elastic; incrusted with cellular debris, striated under SEM, dichotomous branching scarce, threads mostly straight to somewhat sinuous, knob-like projections present, attenuate to a blunt terminus or attenuate to a fine pointed terminus. Pores round, small to medium-sized, abundant. Septa absent or scarce. Paracapillitium present, rare to abundant depending on the age of the specimen, more abundant in mature fruitbodies. Subgleba composed of hyaline, pale yellow, thick-walled, aseptate hyphal threads, up to 4 μm broad, infrequently branched. Exoperidium a combination of textura globulosa and textura intricata; composed of thin-walled, irregular-shaped to globose, sphaerocyst cells containing a matrix of golden pigmented material; globose cells along the upper surface appear hyaline; intertwined with golden brown or amber densely pigmented hyphal elements. Endoperidium textura intricata; composed of compact tangled hyphal threads, thick-walled, septate; some cellular elements oval, flattened, and pigmented amber brown.


DISTRIBUTION—Known from many parts of the United States, and previously reported from Alabama, Alaska, Arizona, California, Colorado, Connecticut, Florida, Georgia, Hawaii, Idaho, Iowa, Illinois, Indiana, Kansas, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Montana, Nebraska, New Hampshire, New Jersey, New Mexico, New York, North Carolina, North Dakota, Oklahoma, Ohio, Oregon, Pennsylvania, South Carolina, Tennessee, Texas, Utah, Vermont, Virginia, Washington, Washington DC, West Virginia, Wisconsin, and Wyoming. Also known from Australia, Austria, Belgium, Brazil, Canada, China, Colombia, Costa Rica, Cuba, Czech Republic, England, Finland, France, Germany, Ghana, Hungary, India, Jamaica, Japan, Latvia, Mexico, Mongolia, Norway, Panama, Poland, Russia, Scotland, South Africa, Sweden, Switzerland, and Turkey.

MATERIAL EXAMINED—CALIFORNIA, Alameda Co.: Berkeley, December 1901, coll. unknown (UCB506549)(UC); Berkeley, on ground, December 1912, coll. unknown (UCB506542)(UC); Berkeley, University of California campus, near the police kiosk at the west gate, under shrubs near Strawberry Creek, 4 February 1969, coll. unknown (UCB1408390)(UC); Berkeley, University of California campus, under shrubs and small trees near a creek on the west side of campus, 15 January 1970, coll. unknown (UCB1408397)(UC); Berkeley, University of California campus, west of the life science building, on ground under oak and pine trees, January 1970, coll. unknown (UCB1461011)(UC); Tilden Regional Park, terrestrial, 5 December 2009, coll. unknown (SSJ 352); Alameda, terrestrial, 3 December 2011, coll. unknown (SSJ 426). Butte Co.: Jonesville, terrestrial, July 1931, E.B. Copeland (Copeland 3702)(HSU); Jonesville, terrestrial, October 1940, E.B. Copeland (Copeland 2119)(HSU). Calaveras Co.: Camp Connell, near Highway 4, solitary in soil under conifers, 23 October 1983, H.D. Thiers (HDT 46642). Contra Costa Co.: Sycamore Canyon, terrestrial, February 1934, coll. unknown (UCB521140)(UC); Oakland hills near the Redwood Bowl, 7 December 1962, coll. unknown (UCB1461015)(UC); Redwood Regional Park, near Redwood Bowl in the Oakland hills, on soil, 7 December 1962, coll. unknown (UCB1461016)(UC). Del Norte Co.: Darlingtonia, caespitose in soil under conifers, November 1937, H.E. Parks (UCB637231)(UC); Jedediah Smith Redwoods State Park, gregarious in humus along the roadside, 20 October 1965, H.D. Thiers (HDT 13640); Jedediah Smith Redwoods State Park, scattered in soil in mixed woods, 9 October 1966, coll. unknown (Sundberg856); Jedediah Smith Redwoods Park, 91 meters (300 feet) elev., terrestrial under spruce, 29 October 2009, R. Pastorino (SSJ 385). Humboldt Co.: Trinidad, spruce grove, December 1937, coll. unknown (UCB1139343)(UC); Trinidad, vicinity of Bishop Pine Lodge, in soil under Alnus rubra (red alder), 4 January 1940, coll. unknown (UCB1139359)(UC); Patrick’s Point State Park, terrestrial, 5 February 1961, D. Largent (DL 450); Patrick’s Point State Park, gregarious to caespitose in soil under Picea sitchensis (Sitka spruce), 10 October 1966, coll. unknown (Ammirati 429); Sloan Creek, caespitose in duff, 27 September 1967, D. Largent (DLL 2121)(HSU); Sloan Creek, terrestrial, 27 September 1967, coll. unknown (2768-2124)(HSU); Arcata, LC Dunes, terrestrial in the Weott Coral, 30 November 1977, S. Sweet (1167477)(HSU); Tish Tang Campground, scattered and terrestrial, 3 November 1979, coll. unknown (MB2-7022); Patrick’s Point State Park, scattered in soil under Sitka spruce, 12 November 1979, H.D. Thiers (HDT 40235); Murry Road near McKinleyville, gregarious in soil under conifers, 1 November 1981, H.D. Thiers (HDT 43839); Patrick’s Point State Park, north of the Big Lagoon, gregarious in duff, 24 October 1982, H.D. Thiers (HDT 45211); McKinleyville along Murry Road, scattered in debris under Picea sitchensis (Sitka spruce), 5 November 1984, H.D. Thiers (HDT 48094); Patrick’s Point State Park, camp 24 near the faucet, clustered in large groups fused at the base, 30 October 1986, coll. unknown (VW 83-10128)(HSU); Patrick’s Point State Park, terrestrial along the road to the alder group area, 9 January 1987, D. DeShazer (DS 432-10474)(HSU); East Fork Campground along the Trinity River, terrestrial, 15 November 2009, S.S. Jarvis (SSJ 379); Gray’s Falls Campground, terrestrial in moss and damp clay rich soil, 15 November 2009, S.S. Jarvis (SSJ 452). Marin Co.: Marin, Inverness, under oaks along a hillside, 27 January 1961, coll. unknown (UCB1408400)(UC); Mt. Tamalpais, near Alpine Lake, 16 December 1962, coll. unknown (UCB1408403)(UC); Marin, between Alpine Lake and Mt. Tamalpais, 19 December 1962, coll. unknown (UCB1408411)(UC); Mt. Tamalpais, near Alpine Lake, 19 December 1962, coll. unknown (UCB1408412)(UCB); near Alpine Dam, under Arbutus menziesii (pacific madrone) and Pseudotsuga menziesii (Douglas fir), 24 November 1963, coll. unknown (UCB1461018)(UC); Alpine Lake, gregarious in mixed conifer woods, 27 October 1972, H.D. Theirs (HDT 30392); Mt. Tamalpais, on soil, 29 October 1972, coll. unknown (UCB1457610)(UC); Mt Tamalpais State Park, clustered under oaks, 16 November 1974, H.D. Thiers (HDT 119); Fairfax, scattered in soil under oaks, 16 November 1975, H.D. Thiers (HDT 35447); Picher Canyon rookery, caespitose on well decayed wood, 7 December 1977, C. Calhoun (Calhoun 77-422); below Bolinas Ridge Road, Audubon Canyon Ranch, gregarious to caespitose on woody debris under Pseudotsuga menziesii (Douglas fir), 6 December 1979, C. Calhoun (Calhoun 79-1246); Mt. Tamalpais, terrestrial in an open field near the edge of woods, 30 November 2008, S.S. Jarvis (SSJ 271); Point Reyes, growing on some hardwood under fir trees, 7 February 2009, S.S. Jarvis (SSJ 283); Point Reyes National Seashore, Bolinas Ridge, 19 February 2009, R. Pastorino (SSJ 357); Point Reyes National Seashore, Bolinas Ridge, 4 March 2009, R. Pastorino (SSJ 358). Mendocino Co.: Hendy Woods State Park, scattered at base of Arbutus menziesii (pacific madrone), 25 November 1991, M. Seidl (UCB1598446)(UC); Mendocino city, in deep damp dark woods along the Albion River, December 1949, coll. unknown (UCB915614)(UC); Jackson State Forest, on soil and mixed gravel, 21 November 1971, H.D. Thiers (HDT 28559); Ukiah, Lake Mendocino, gregarious in humus in oak woods, 24 November 1982, H.D. Thiers (HDT 45462); Coast Range Preserve, gregarious in humus under mixed woods, 2 November 1975, H.D. Thiers (HDT 35331); Jackson State Forest near Mendocino city, in soil in mixed woods, 28 October 1962, coll. unknown (Malloch 524); Jackson State Forest, gregarious in soil in mixed woods, 2 December 1974, H.D. Thiers (HDT 33159); Jackson State Forest, caespitose in humus in dense mixed woods, 3 November 1963, H.D. Thiers (HDT 10587); Jackson State Forest near Mendocino city, in moss beds under redwoods, 8 December 1974, coll. unknown (Strick 96); Gualala, Iverson Road, in moss under redwood, November 2007, S.S. Jarvis (SSJ 233); Jackson State Forest, Albion Station, 1 February 2009, S.S. Jarvis (SSJ 286); Booneville, in damp thick woods, April 2009, J. Edmonds (SSJ 393); Gualala, Fish Rock Road, along the Pacific Lumber Company land, on moss and soil, November 2011, S.S. Jarvis (SSJ 439); Gualala, Fish Rock Road, along the Pacific Lumber Company land, on moss and soil, December 2007, S.S. Jarvis (SSJ 232). Monterey Co.: Pacific Grove, about 1/3 mile east of the toll gate to 17 Mile Drive, in open places in woods with rich sandy soil, 11 March 1940, coll. unknown (UCB638444)(UC). Napa Co.: Cleary Reserve, gregarious in humus under conifers, 27 October 1963, H.D. Thiers (HDT 10561); Cleary Reserve, scattered in duff under pines, 13 December 1964, coll. unknown (Sundberg 155); Angwin, Las Posadas Preserve at Inspiration Point, under pine, oak and for woodland, 16 February 2010, S.S. Jarvis (SSJ 430). Plumas Co.: Quincy, terrestrial, 18 October 1930, L. Bonar (UCB506540)(UC). San Bernardino Co.: Camp Osceola, between fir and oak in stands of Pinus ponderosa (ponderosa pine), 27 September 1976, coll. unknown (UCB1462466)(UC); San Bernardino National Forest, Highway 38, along the Santa Ana River at South Fork Campground towards Big Bear, 2,438 meters (8,000 feet) elev., under Pinus monophylla (pinyon pine) and Salvia pachyphylla (rose sage), caespitose in a dried up water washout in sandy rocky soil, 7 October 2008, S.S. Jarvis (SSJ 251). San Francisco Co.: San Francisco, Golden Gate Park, 5 January 1928, coll. unknown (UCB506546)(UC); San Francisco, UC Medical Center, under conifers and willows, 2 December 1960, coll. unknown (Johnson 28). San Mateo Co.: San Bruno, Junipero Serra Park, scattered in humus under oaks, 5 December 1960, H.D. Thiers (HDT 8561); San Francisco Watershed, Cahill Ridge, 18 July 1967, H.D. Thiers (HDT 20185); San Francisco Watershed, solitary under mix hardwood, 23 December 1969, H.D. Thiers (HDT 649); Daly City, Doelger Senior Center, under Pinus radiata (Monterey pine), 8 March 2009, F. Stevens (SSJ 359). Santa Cruz Co.: Santa Cruz Mountains near Boulder Creek, scattered to caespitose in humus of mixed woods, 22 November 1963, H.D. Thiers (HDT 10778); along Highway 236 between Boulder Creek and Big Basin, solitary in soil in mixed woods, 10 November 1975, coll. unknown (Halling 1059). Sierra Co.: Yuba Pass, clustered near rotten conifer log, 22 September 1975, H.D. Thiers (s.n.); Gold Lake Road near Bassetts, Sand Pond picnic area, in soil under conifers and willows, 24 October 1988, coll. unknown (Zebell 120); Yuba Pass, 2,042 meters (6,701 feet) elev., caespitose in coniferous woods, 9 June 1989, coll. unknown (UCB1574381)(UC); Wild Plum Campground near Sierra City, gregarious in soil under conifers, 5 October 1989, H.D. Thiers (HDT 52790); Green Acres, terrestrial under snow, 1,524 meters (5,000 feet) elev., June 2012, R. Pastorino (SSJ 466). Shasta Co.: Lakeshore Village north of Redding, near Lake Shasta, scattered in humus in oak and pine woods, 5 November 1979, H.D. Thiers (HDT 40287). Siskiyou Co.: Klamath National Forest, Cook and Green Pass, terrestrial beneath fir trees, 7 October 1977, D. Largent (DL 7047-6209)(HSU). Sonoma Co.: Santa Rosa, along Spring Mountain Road crossing Calistoga Road, in a narrow canyon with Sequoia sempervirens (coastal redwood), 22 November 1963, coll. unknown (UCB1408398)(UC); Occidental, Camp Meeker, Largent property, scattered to gregarious beneath humus of Pseudotsuga menziesii (Douglas fir), 7 November 1968, D. Largent (DLL 3541-2122)(HSU); Occidental, Bohemian Highway, on moss under redwood canopy, 7 December 2008, S.S. Jarvis (SSJ 278); Occidental, CYO Camp along Bohemian Highway, terrestrial, December 2009, S.S. Jarvis (SSJ 429); Occidental, CYO Camp along Bohemian Highway, terrestrial, 17 January 2010, S.S. Jarvis (SSJ 363); Occidental, CYO Camp along Bohemian Highway, terrestrial, 17 January 2010, S.S. Jarvis (SSJ 365). Tehama Co.: Mineral, along Highway 36, caespitose on rotten log, 10 October 1975, H.D. Thiers (HDT 36630); Mineral, on the ground, 2 October 1976, coll. unknown (UCB1472586)(UC); Mineral, on the ground under conifers, 4 October 1976, coll. unknown (UCB1472619)(UC); Gurnsey Creek Campground, at the intersection of Highway 36 and Highway 89, south of Lassen National park, gregarious in soil under conifers, 30 September 1989, H.D. Thiers (HDT 52781). Yuba Co.: Bullard’s Bar Recreation Area, clustered in soil in mixed woods, 10 November 1984, H.D. Thiers (HDT 58209).


HABITAT—Often growing solitarily, or in groups up to twenty-five individuals. Sometimes found as small caespitose clusters with up to four to eight individuals fused together at the base, and found in dense clusters taking up nearly of a square foot of space. The right environmental conditions for prolific growth seem to be cold winter weather and rain along the coastal regions in oak woodland, pine and fir forests, or in redwood forests. Also collected in the spring after the snow melt in higher elevations up to 2,438 meters (8,000 feet) in high desert pinyon and juniper woodland, in high desert transition forests of Southern California, and in densely forested areas where moisture seems to persist. Collected growing on very decayed wood, on duff, in grass, and on soil. Among the Geographic Subdivisions of California, this species can be found in Northwestern California, in the Cascade Range, in the Great Central Valley, in the Sierra Nevada, in Central Western California, and in Southwestern California.


The Lycoperdaceae of California, Thesis by Steph Jarvis

For images, light microscopy & Scanning Electron Microscopy:

Additional References:


COMMENTS—Lycoperdon perlatum has been deemed the most common puffball in Europe by Demoulin (Bates 2004). Lycoperdon perlatum has worldwide distribution and ecological variance, being found growing on a variety of substrates from leaf litter to decaying wood, and is often divided into separate varieties due to its morphological variation. This puffball is distinct from others due to the exaggerated pseudostipe and spinose exoperidium. Persoon (1796) did not designate a type specimen, and a neotype has not been designated. The herbarium material that Demoulin (Bates 2004) used to circumscribe Lycoperdon perlatum, from the National Herbarium Nederland, Leiden, did not have collection dates or locations with the specimens, making them unavailable as potential lectotype. These collections now have barcode numbers in addition to their accession numbers; (accession number L 910.258-671, L 910.258-676, L 910.258-681, L 910.258-761), (barcode number L 0116482, L 0116484, L 0116483, L 0116486, L0116487). The ITS sequence data nests Lycoperdon perlatum with the diaphragm-forming puffballs, as was shown by Bates (2004). As well, Calvatia sculpta is shown to be sister to the clade of Lycoperdon perlatum with 100% bootstrap and 100% PP. Being the type species for the genus Lycoperdon, Lycoperdon perlatum does not seem phylogenetically related to many of the species within Lycoperdon. In light of this evidence, this shows that ITS data is not enough to resolve taxonomic questions with the Lycoperdaceae, suggesting that multi loci gene analysis is necessary to further clarify their taxonomic positions.

Lycoperdon Pers., Ann. Bot. (Usteri) 1: 4. 1794.
TYPE—Lycoperdon perlatum Pers., Observ. mycol. (Lipsiae) 1: 145. 1796.

Currently the largest genus in the Lycoperdaceae, Lycoperdon was circumscribed by Persoon in 1794, published in Annalen der Botanik ed. Usteri, Zurich. However, the genus predates this publication to the 1753 publication, Species Plantarum, by Carl Linnaeus, when it was a mix of organisms, including anything with a globose fruitbody (Demoulin 1973a, Bates 2004). Further contributions to Lycoperdon have been made by Cunningham (1942, 1979), Kreisel (1973), Demoulin (1972b, 1983), Jeppson (1984, 2006, 2010, 2011, 2012), Larsson and Jeppson (2008), and Pegler (1995), and others.

Lycoperdon is characterized by a fruitbody having a single apical pore, hyphae without clamps, and eucapillitium that are branched without having tapered tips. The eucapillitium is categorized as Lycoperdon-type, which means that the threads are mostly elastic, with very little dichotomous branching, poroid, and with variable thickness. As with most of the puffballs, Lycoperdon is a cosmopolitan genus. Several studies available, which describe the development and histology of fruitifications of Lycoperdon, such as; Rabinowitsch 1894, Swartz 1933, Ritchie 1948, Marchant 1969, and the book by Smith et al. 1981.

The genus currently has 441 species that have been recorded in Index Fungorum, 790 including all varieties. Lycoperdon is the type genus of the Lycoperdaceae, and Lycoperdon perlatum is the type species for the genus. There are twelve species reported in California: L. dermoxanthum, L. curtisii, L. molle, L. lloydianum, L. nigrescens, L. perlatum, L. pratense, L. pyriforme, L. subcretaceum, L. umbrinum, L. utriforme, and the provisional new species L. vernimontanum.

Description author: Steph Jarvis (Request Authorship Credit)
Description editor: Jason Hollinger