Steph Jarvis’s Description of Calvatia pachyderma (Peck) Morgan

Title: Monograph Of The Lycoperdaceae Of California By Steph Jarvis (Default)
Name: Calvatia pachyderma (Peck) Morgan
View: public
Edit: private
Version: 6
Previous Version 

Descriptions: Create
 Monograph Of The Lycoperdaceae Of California By Steph Jarvis (Default)

Description status: Unreviewed
 (Latest review: 2019-10-21 05:13:22 AEST (+1000) by jason)

Taxonomic Classification:

Domain: Eukarya
Kingdom: Fungi
Phylum: Basidiomycota
Class: Agaricomycetes
Order: Agaricales
Family: Lycoperdaceae

General Description:

Calvatia pachyderma (Peck) Morgan, J. Cincinnati Soc. Nat. Hist. 12: 167. 1890.
FIGURE 14, 54
Basionym: Lycoperdon pachydermum Peck, Bot. Gaz. 7(5): 54. 1882.
Reported synonyms:
   = Langermannia pachyderma (Peck) Kreisel, Feddes Repert. 64: 120. 1962.

TYPE—Peck’s collection of Lycoperdon pachydermum is deposited in the New York State Museum in Albany, NY; some of this material is labeled as Calvatia pachyderma and was collected from Utah and California. There is a collection of Lycoperdon pachydermum Peck, at the New York Botanical Garden, collected by CG Pringle, Arizona, 1884, (NYBG 418), determined by CH Peck, that is labeled as “possibly part of holotype” by V. Demoulin in his 1977 notes on this specimen. Kreisel (1992) says that the holotype material for Lycoperdon pachyderma Peck 1882 is from Arizona.

Diagnostic Description:

GASTEROCARP 45-120 mm tall x 30-120 mm broad; globose to depressed globose, plicate at the base with folds and ridges, pulvinate with slight swelling at the base, or slightly tapered to a point; rhizomorph typically 5-15 mm long, up to 1 mm broad at the base where attached at the fruitbody, tapering quickly to fine tangled hyphae, remaining attached through maturation, radial wrinkles of the peridium ascending from the point of rhizomorph attachment; ostiole opening via lateral to stellate cracks that reveal the gleba. Exoperidium cream white at first (4A2-3), turning light tan clay color (5C4), to darker brown (5E6-6F7-7F7) or reddish brown (8E7) in patches and where exposed to the sun, with variable patches of color through maturity; exoperidium up to 1 mm thick, remaining mostly to completely adherent to the endoperidium, sloughing off in patches to expose the endoperidium, slightly furfuraceous to compact fleshy when very young to smooth or glabrous in age, becoming scaly or warted with dry weather, brittle with age. Endoperidium cream white when young (4A2-3), turning light tan clay color (5C4), dark brown with age (7F7), sometimes with a metallic sheen and sun bleached white when very old; soft when young, becoming hard, rigid, and thick in age, up to 2-3 mm thick; remaining adherent to the exoperidium in patches, persistent, irregular cracking and falling away to reveal the gleba with the exoperidium, leaving a worn away cup-like base made of only subgleba and edges of endoperidium remnants in the very oldest fruitbodies. Gleba white to cream when young (4A2-3), becoming shades of olive green to dark olive green brown (4E7-4F7) as the fruitbody matures; separating easily from endoperidium in all stages of maturity, chunky, becoming “friable into grumous powder” (Homrich and Wright, 1973), or pulverulent, disseminating with wind. Subgleba absent. Diaphragm absent.

BASIDIOSPORES globose to broadly ellipsoidal; 4-5.6 X 3.5-4.8 µm [xmr = 4.3-5.3 X 3.7-4.1 µm, xmm = 5.0 ± 0.6 X 3.9 ± 0.2 µm, Q = 0.9-1.8, Qmr = 1.1-1.5 Qmm = 1.3 ± 0.2, n = 25, s = 3]; spores golden brown in wet mounts; smooth under light microscope, smooth under SEM with peeling flakiness; oil drop present; spores thin-walled, easily deflating with desiccation, easily damaged in SEM imaging; pedicel remaining as a long torn tail, up to 1 µm long, some pedicels short up to 0.5-0.8µm and broken at the end; free-floating sterigmata absent from mounts; spores of equal size under light microscopy. Eucapillitium Calvatia-type; 3.5-9 µm broad with walls up to 1 µm thick; very fragile, readily breaking, dichotomously branching, some walls irregular and sinuous, straight along thick threads to somewhat undulate in thinner threads, ends attenuate to a rounded terminus. Pores abundant, small to medium in size, punctate. Septate, disarticulating at the septa. Paracapillitium absent. Exoperidium textura globulosa; composed of hyaline, irregular-shaped, tightly packed, inflated when fresh, collapsed when dry, thick-walled sphaerocyst cells. Endoperidium textura intricata; composed of septate, thick-walled, tightly woven hyphal threads. Basidia densely clustered, ovoid, small, with three papilliform having short stout sterigmata, 4 spores per basidia.


DISTRIBUTION—Known only in the Western United States, and previously reported from Arizona, California, Colorado, Oregon and Utah. Also reported from Argentina, Australia, Baja Mexico, Chile, Iran, Mexico, Spain, Nepal, and Uruguay.

MATERIAL EXAMINED—CALIFORNIA, Alameda Co.: Berkeley, Fall 1899, R.E. Gibbs (UCB506655)(UC); Berkeley, hillside above the California School of Deaf, 6 March 1917, N.L. Snardner (UCB506653)(UC); Berkeley, Strawberry Canyon, 27 May 1928, E.E. Morse (UCB506649)(UC); Hayward, terrestrial, Spring 1941, (UCB506650)(UC); Berkeley, gregarious on bare spots in deep grass of a hillside, 21 April 1941, T.T. SmCabe (UCBM62556)(UC); Berkeley, Westerly spurs of hills above the California School of Deaf, gregarious in a 30-foot fairy ring, 9 June 1941, coll. unknown (UCBM62398)(UC). Calaveras Co.: Valley Springs, crest of Campo Seco Road, light shade on grassy South facing slope, 12 May 1986, T. Tang (UCB15713017)(UC). Marin Co.: Tiburon, hillside, 15 April 1924, E.E. Morse (UCB506651)(UC); Muir Woods, Dipsea Trail, West of Monument, top of the ridge, terrestrial, gregarious, October1962, Brown (Brown641); Bolinas, in grass of an open field, 27 October 1980, C. Calhoun (Calhoun 208); Pt. Reyes National Seashore, open grassy area, 24 April 2007, R. Pastorino (SSJ 391). Monterey Co.: Monterey, growing in an orchard, 27 May 1933, L. Bonar (UCB506652)(UC). Napa Co: St. Helena, Biel Family Ranch, in a vineyard, in between vine rows in disturbed soil, 20 October 2011, B.A. Cothran (SSJ 411). San Diego Co.: San Clemente Canyon, Highway 52 and Highway 5 Junction, 7 October 2008, S. Andrasko (SSJ 250). Santa Clara Co.: Palo Alto, near Boble Hall, in leaves and grass, 21 November 1926, E.E. Morse (UCB5106654)(UC). Santa Cruz Co.: Wilder Ranch, common in rings in open meadow, October 1986, Heldt (Heldt 230). San Mateo Co.: San Francisco Watershed, solitary in open grassy area, 13 November 1975, H.D. Thiers (HDT 35404). Sonoma Co.: Skaggs Springs Road, ten miles west of Lake Sonoma, open pasture in disrupted soil among Quercus lobata (valley Oak), 21 November 2008, S.S. Jarvis (SSJ 262); Knights Valley, Foote Ranch, Highway 128, East facing grassy slope, 5 December 2008, S.S. Jarvis (SSJ 275); Knights Valley, Foote Ranch, solitary under Pseudotsuga menziesii (Douglas fir), 16 October 2011, S.S. Jarvis (SSJ 410); Knights Valley, Foote Ranch, upper grassy slopes in an arched ring, terrestrial, 20 October 2011, S.S. Jarvis (SSJ 413).


HABITAT—Calvatia pachyderma is said to have an amphitropical distribution of mostly temperate regions, and being discontinuous between Northern and Southern hemispheres (Ponce de Leon, 1976). Occurring in the fall after light rain, and sometimes in spring after heavy rain. Collected in grassy lawns where grass appears scant and patchy, on open grassland, in pastures, on ranch land, among Quercus lobata (valley oak), Quercus agrifolia (coast live oak), and found in clay-rich humiferous soils. Growing in fairy rings, in groups of two or three, touching but not fused, or solitarily. Growing on ground with frequent foot traffic, or periodic vehicle traffic and somewhat shaded areas. Due to teeth makings on the gasterocarps, young fruitbodies were noticed to be nibbled on by cows and small rodents. Among the Geographic Subdivisions of California, this species is found in the southern regions of Northwestern California, in the western regions of the Sierra Nevada, in Central Western California, and in South Western California.


The Lycoperdaceae of California, Thesis by Steph Jarvis

For images, light microscopy & Scanning Electron Microscopy:

Additional References:


COMMENTS—Calvatia pachyderma is currently recognized on Index Fungorum as Langermannia pachyderma (Peck) Kreisel. However, due to morphological and molecular evidence, this species will be treated in Calvatia here. Prior to this name transfer, Calvatia pachyderma was recognized as Gastropila fragilis (Lév.) Homrich and Wright. Gastropila was erected as a genus based on characters such as a peridium with three distinct layers, smooth spores, and capillitium described as “smooth threads sparsely branched, not easily broken (elastic), much entangled” (Homrich & Wright 1973). Ponce de Leon (1976) states that C. pachyderma and Gastropila fragilis are synonymous. Although C. pachyderma does have truly smooth spores, the material from California does not exhibit a three-layered peridium as described by Homrich & Wright. Additionally, C. pachyderma has Calvatia-type eucapillitium that is frequently branched, is fragile, and disarticulates easily into small fragments through the maturation of the fruitbody. Demoulin (1993) states “whether one should retain the genus Gastropila or not is very much a matter of taste,” in relation to putting this species in Calvatia, or retaining Gastropila as a monospecific genus.

Currently Langermannia pachyderma (basionym Lycoperdon pachydermum Peck) and Gastropila fragilis (Lév.) Homrich & J.E. Wright (basionym Mycenastrum fragile Lev.) are separate species on Index Fungorum. Previous studies of holotype material recognize that Lycoperdon pachydermum Peck was synonymous with Mycenastrum fragile Lév, and with Gastropila fragilis (Lév.) Homrich and Wright (Ponce De Leon 1976, Wright 1990, Demoulin 1993). After examination of spores with a scanning electron microscope from type specimens of both Lycoperdon pachydermum and Mycenastrum fragile, Ponce de Leon (1976) recognized that these two species are conspecific. The original type material of Mycenastrum fragile Lév. is from Uruguay, whereas the original type material of Lycoperdon pachydermum Peck is from Arizona. However, Ponce de Leon (1976) discusses long distance spore dispersal starting in the deserts and mountains of western North America, migrating south in recent geological history through wind, birds, etc., gained a foothold of this species in the Andes of Chimu and in the mountainous temperate regions of South America. He argues that the name Gastropila fragilis takes precedence over Calvatia pachyderma based on misunderstanding, contradictions and correct assumptions of the species from several works (Leveille 1844, Peck 1882, de Toni 1888, Morgan 1890, Lloyd 1904 & 1916, Coker and Couch 1928, Clemens 1931, and by Homrich and Wright 1973).

In the ITS analysis by Bates (2004), Calvatia pachyderma (Arizona material) was placed in a Calvatia clade that included Calvatia gigantea & C. bicolor with 96% bootstrap support. This included direct support of a sister Calvatia clade with C. cf. leiospora, C. cyathiformis, & C. fragilis supported by 99% bootstrap support. Calvatia pachyderma, based on published ITS data, clearly falls into a Calvatia clade. Bates’ ITS data does not support recognition at this time as Langermannia pachyderma as distinct from Calvatia based on this molecular data. Further molecular analysis on South America specimens of Gastropila fragilis in comparison to North American specimens of Calvatia pachyderma is needed to confirm Ponce de Leon’s suggestions of these two species being conspecific. Calvatia pachyderma will be recognized here based on macro-characters, micro-characters, and phylogenetic evidence. The ITS data here supports Calvatia pachyderma within the Calvatia clade, sister to Calvatia booniana, but with only a little support (less than 70% bootstrap and 91% PP). This suggests that further analysis using multi gene loci is necessary to clarify the taxonomic position of Calvatia pachyderma within the Lycoperdaceae.

Calvatia Fr., Summa veg. Scand., Section Post. (Stockholm): 442. 1849.
TYPE—Calvatia craniiformis (Schwein.) Fr., Summa veg. Scand., Section Post. (Stockholm): 442. 1849.

Elias Magnus Fries contributed extensive taxonomic work to the Gasteromycetes. In 1849 he published the genus Calvatia in Summa vegatabilium Scandinaviae, Section Post. (Stockholm): 442. Andrew Price Morgan (1890) amended the genus to accommodate species whose peridium breaks apart in patches from the top downward. The original concept of Calvatia remains relatively unchanged in works published by Zeller and Smith (1964) and Kreisel (1989, 1992, 1994), however various workers have transferred some species to other genera. For example, Kreisel (1989) transferred some species of Calvatia into Handkea. These species have since been re-evaluated with molecular phylogenetic data and are being moved into Lycoperdon based on evolutionary relationships.

Most species of Calvatia can be found in xerophytic, or mesophilic grasslands, in arctic-alpine regions or with semi-desert vegetation, in gardens with cultivated soils, or in temperate forests with shade (Kreisel 1992). Calvatia has a worldwide distribution, recorded by Lange (1977, 1990, 1993) in the Antarctic zone, and studied in the moist and arid tropics by many mycologists.

The genus Calvatia currently has eighty-eight species recorded in Index Fungorum, 136 including all varieties. The type species is Calvatia craniiformis (Schw.) Fries. There are six species recorded in California: C. booniana, C. fragilis, C. fumosa, C. lloydii, C. pachyderma, and C. sculpta.

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