Napa Valley Fungi

plant pathogen diagnostic laboratory & cultivation of medicinal fungi

Key to the Lycoperdaceae

 

TAXONOMY

Artificial Key to California Puffballs (Lycoperdaceae)

1a. Fruitbodies >35 mm tall, >35 mm broad when mature; peridium layers thick .......................................................................................................................................................... 2

1b. Fruitbodies <35 mm tall, <35 mm broad when mature; peridium layers thin .......................................................................................................................................................... 8

2a. Subgleba absent ............................................................................................................................. 3

2b. Subgleba present, prominent or compact and reduced  ....................................................................... 4

3a. Eucapillitium of Mycenastrum-type: spiny; exoperidium tough; spores pitted, on average 11 x 11 µm ........................................................................................................................ 35. Mycenastrum corium

3b. Eucapillitium of Calvatia-type: fragile; exoperidium brittle; spores smooth, on average 4.5 x 4.5 µm ....................................................................................................................... 12. Calvatia pachyderma

4a. Diaphragm present; fruitbody obpyriform ....................................................... 33. Lycoperdon utriforme

4b. Diaphragm absent; fruitbody depressed globose to obpyriform ............................................................ 5

5a. Eucapillitium of Calbovista-type, elastic with numerous ramified branches; exoperidium with large polygonal warts with blunt tips ................................................................................................. 6. Calbovista subsculpta

5b. Eucapillitium of Calvatia-type, fragile and disarticulating into small fragments; exoperidium with various scales and warts ............................................................................................................................................... 6

6a. Gleba spore mass purple brown; exoperidium with fragile areolate patch-like scales and cracks ................................................................................................................................. 8. Calvatia fragilis

6b. Gleba spore mass dark brown to olive brown; exoperidium with distinct warts ...................................... 7

7a. Exoperidium warts <30 mm broad and <35 mm tall, pyramidal and pointed or hooked at the tips; spores echinulate, on average 5 x 5 µm ................................................................................................ 13. Calvatia sculpta

7a. Exoperidium warts >60 mm broad and >80 mm tall, mammiform and appressed; spores asperulate to asperate, on average 4.5 x 4.5 µm .............................................................................................. 7. Calvatia booniana

8a. Diaphragm present; paracapillitium abundant; Lycoperdon-type eucapillitium ....................................... 9

8b. Diaphragm absent; paracapillitium of various amounts; eucapillitium of various types ............................ 11

9a. Fruitbody small, <20 mm tall. <20 mm broad; exoperidium densely covered in minute sharply pointed spines; ostiole developing as a slit-like crack or a small round hole .......................................................................................................................... 23. Lycoperdon curtisii

9b. Fruitbody larger, >20 mm tall, >20 mm broad; exoperidium covered in minute furfuraceous spines and granular particles when young; ostiole opening broadly at the apex ...................................................................... 10

10a. Eucapillitium growing into the gleba from inner walls of the diaphragm only; free-floating sterigmata remnants absent in wet mounts; spores asperulate to asperate, on average 4.5 x 4.5 µm   .................................................................................................................... 25. Lycoperdon lloydianum

10b. Eucapillitium growing into the gleba from the inner walls of the endoperidium only; free-floating sterigmata remnants present in wet mounts; spores smooth to asperate, on average 4 µm x 4 µm  ....................................................................................................................... 29. Lycoperdon pratense

11a. Exoperidium forming a cup-like sand case around the fruitbody; eucapillitium of Disciseda-type ....................................................................................................................................................... 12

11b. Exoperidium of various forms, cup-like sand case absent; eucapillitium of various types ........................................................................................................................................................ 21

12a. Endoperidium rigid, not pliable; spores pitted, on average 11 x 11 µm ........................ 1. Abstoma townei

12b. Endoperidium variable, pliable; spore ornamentation and size variable ............................................... 13

13a. Endoperidium layers thick and leather-like; exoperidium thin, wearing off early in maturation ........................................................................................................................................................ 14

13b. Endoperidium layers thin and flexible, paper-like; exoperidium thick, adhering to soil and sand particles, remaining throughout maturation .......................................................................................................... 15

14a. Eucapillitium pores scarce to absent; spores broadly ellipsoidal, on average 5 µm x 4 µm ................................................................................................................................. 14. Disciseda atra

14b. Eucapillitium pores oval to round, scattered; spores globose, on average 4.5 x 4.5 µm ......................................................................................................................... 19. Disciseda levispora

15a. Exoperidium basal remnants wearing thin ....................................................................................... 16

15b. Exoperidium remaining as thick basal remnants ............................................................................... 17

16a. Endoperidium wrinkled to cracked over the entire surface; dark yellowish brown; exoperidium leaving a film-like layer between cracks; spores smooth, on average 7 x 7 µm ........................................................................................................................ 15. Disciseda brandegeei

16b. Endoperidium cracked along the outer margin, smooth at the apex; tan tawny brown; exoperidium leaving a thin crust at the base; spores echinulate to echinate, on average 4.5 x 4.5 µm .......................................................................................................................... 18. Disciseda johnstonii

17a. Exoperidium becoming patch-like with maturation .............................................................................. 18

17b. Exoperidium becoming a prominent cup-like sand case attached at the base .......................................... 19

18a. Endoperidium yellow brown; spores asperulate to asperate, on average 6 x 6 µm ............................................................................................................................... 20. Disciseda luteola

18a. Endoperidium purple gray brown; spores asperate to verrucose, on average 7 x 7 µm ............................................................................................................................ 22. Disciseda uplandii

19a. Fruitbody hypogeous to subhypogeous ........................................................... 21. Disciseda subterranea

19b. Fruitbody subhypogeous to epigeous ............................................................................................... 20

20a. Exoperidium sand case up to 0.5 mm thick; reticulate pattern on the base of the endoperidium; spores asperate to verruculose, on average 5 x 5 µm .............................................................................. 16. Disciseda candida

20b. Exoperidium sand case up to 1 mm thick; reticulate pattern absent; spores smooth to asperate, on average 5.5 x 5.5 µm ................................................................................................................... 17. Disciseda cervina

21a. Exoperidium with a smooth to cracked surface, or hexagonal scales; subgleba compact to absent  ......................................................................................................................................................... 22

21b. Exoperidium with pyramidal warts to furfuraceous spines, or a granular coated surface; subgleba prominent and pseudostipitate .................................................................................................................................... 31

22a. Exoperidium with smooth to cracked surface; eucapillitium Bovista-type or Intermediate-type; grassland species .......................................................................................................................................................... 23

22b. Exoperidium with hexagonal scales; eucapillitium Calvatia-type or Lycoperdon-type; growing in conifer duff or a meadow species in montane regions ....................................................................................................... 26

23a. Eucapillitium of Intermediate-type ................................................................................................... 24

23b. Eucapillitium of Bovista-type .......................................................................................................... 25

24a. Fruitbody >10 mm tall, >10 mm broad; common, collected from lowland or montane meadows; paracapillitium absent; spore pedicel rudimentary; spores asperulate to asperate, on average 4.5 x 4.5 µm .............................................................................................................................. 2. Bovista aestivalis

24b. Fruitbody <10 mm tall, <10 mm broad; uncommon, collected from moist montane drainage fields; paracapillitium abundant; spore pedicel long and prominent; spores verruculose to echinulate, on average 5 x 5 µm ............................................................................................................................ 3. Bovista californica

25a. Exoperidium with a metallic sheen when mature, peeling away in dry thin sheets with age; young gleba tissue staining yellow with KOH; spore pedicel rudimentary; spores asperate to verrucose, on average 4.5 x 4.5 µm ...................................................................................................................................... 4. Bovista pila

25b. Exoperidium without a metallic sheen, peeling away in patches when young and moist; young gleba without a staining reaction; spore pedicel long and prominent; spores asperulate to asperate, on average 5 x 5 µm ................................................................................................................................. 5. Bovista plumbea

26a. Eucapillitium of Lycoperdon-type; paracapillitium present ......................... 24. Lycoperdon dermoxanthum

26b. Eucapillitium of Calvatia-type; paracapillitium various, abundant to absent .......................................... 27

27a. Subgleba present, compact with chambered cellular type tissue   ..................................................... 28

27b. Subgleba absent  ........................................................................................................................ 29

28. Fruitbody rooted with a tuft of mycelium; exoperidium composed of thin brittle polygonal scales; spores verruculose to verrucose, on average 4 x 4 µm ................................................................ 10. Calvatia lloydii

28b. Fruitbody rooted with a robust rhizomorph; exoperidium furfuraceous-spinulose when young, developing stellate cracking with age; spores echinulate, on average 6 x 6 µm ............................... 34. Lycoperdon vernimontanum

29a. Exoperidium with short blunt to pointed warts; paracapillitium absent ................................................................................................................... 31. Lycoperdon subcretaceum

29b. Exoperidium with a smooth cracked surface; paracapillitium present .................................................... 30

30a. Fruitbody with musty scent when mature; developing subhypogeous under thick conifer duff; eucapillitium pores abundant, slit-like cracks abundant; spores echinulate to echinate, on average 5 x 5 µm ..................................................................................................................................... 9. Calvatia fumosa

30b. Fruitbody without musty scent; developing epigeous along montane lakes, on moss in moist sandy soil; eucapillitium pores scarce to absent, slit-like cracks absent; spores asperulate to verrucose, on average 6.5 x 6.5 µm ................................................................................................................................. 11. Bovista sierraensis

31a. Exoperidium covered in pyramidal warts, leaving prominent circular scars on the endoperidium when sloughing off ......................................................................................................................................................... 32

31b. Exoperidium covered in pyramidal warts, or fine furfuraceous spines or warts, leaving a smooth endoperidium when sloughing off .............................................................................................................................. 33

32a. Exoperidium becoming dark brown to black; free-floating sterigmata remnants present; spores verruculose, ....................................................................................................................... 27. Lycoperdon nigrescens

32b. Exoperidium remaining white to tan brown; free-floating sterigmata remnants scarce; spores echinulate, ......................................................................................................................... 28. Lycoperdon perlatum

33a. Fruitbody growing on soil; free-floating sterigmata present in wet mounts .............................................................................................................................. 26. Lycoperdon molle

33b. Fruitbody growing on wood or humus; free-floating sterigmata absent from wet mounts ........................................................................................................................................................... 34

34a. Fruitbody growing on decayed wood, wood chips, raw humus; rooted to the substrate with thick entangled rhizomorphs; growing in large gregarious clusters; exoperidium furfuraceous-spinulose; spores asperulate to asperate, on average 3 x 3 µm ............................................................................................. 30. Lycoperdon pyriforme

34b. Fruitbody growing on conifer duff, sometimes on fir cones; rooted to the substrate by a tuft of mycelium; growing solitarily or in small gregarious clusters; exoperidium finely warted to minutely spinulose; spores asperate to verruculose, on average 4.5 x 4.5 µm .......................................................................32. Lycoperdon umbrinum

 

ABSTOMA G. Cunn.:

Abstoma townei (Lloyd) Zeller, Mycologia, 39(3): 306. 1947.                                                                                                                                                                                    (fig. 3, 43)

Basionym º Catastoma townei Lloyd, Mycol. Writ. 7(Letter 67): 1168. 1922.

            Type: The holotype collection is located in the U.S. National Fungus Collections (BPI), collection # 706815. A fragmented isotype collection is located at the New York Botanical Garden, catalog # 00291990. The type location is from San Bernardino, San Bernardino County, with the substrate and exact location unknown.

Gasterocarp 25 mm tall x 31 mm broad; globose to depressed globose; rhizomorph composed of thin hyphal threads; ostiole opening via a crack with age. Exoperidium white at first, turning orange white (5A2), then hazel to light brown (6D5-6) with maturity; heavily incrusted with sand and soil debris, persistent and remaining attached to the endoperidium, not leaving a cup-like sand case. Endoperidium white when young, turning orange gray (5A2-5B2); thin and smooth, tough and persistent. Gleba white to yellowish white to cream (4A3-2), turning orange to greenish yellow (4B7-4C7), eventually becoming a grayish yellow (5D3) then dark brown (5F5); being cottony and firm at first, becoming powdering with maturity. Subgleba absent. Diaphragm absent. Macromorphological description derived from the dried type specimen.

            Basidiospores globose; (9) 10-12.5 (13.5) X (8.5) 9.5-12.5 µm [xmr = 11.2 X 10.6  µm, xmm = 11.2  ± 0.0 X 10.6 ± 0.0 µm, Q = 1.0-1.2, n = 25, s = 1]; deep brown in wet mounts; roughened under light microscope, with SEM spores heavily pitted, round to oblong; oil drop absent; spores thick-walled; with a short hyaline pedicel ± 0.8 µm long, pedicel pinched where broken from the sterigma seen with SEM; free-floating sterigmata not present in wet mounts; spores of various size in wet mounts. Capillitium Calvatia-type; hyphal threads up to 4 µm broad, with walls < 0.8 µm thick; hyaline in water mounts, slightly yellow in KOH mounts; fragile and breaking easily glabrous, dichotomous branching present and not abundant, threads short, some segments curled and simple; tips with a blunt round terminus. Pores absent. Septa present and abundant. Paracapillitium absent. Exoperidium textura globulosa; composed of irregular-shaped sphaerocyst cells, collapsed and difficult to discern. Endoperidium textura intricata; composed of intertwined, thin-walled, non-septate, short hyphal elements.

Habitat: Terrestrial. Among the Geographic Subdivisions of California, this species has been found in South Western California, in the San Bernardino Mountains. It has also been reported, but not confirmed, to grow near the Greek Theater on the University of California, Berkeley, campus.

Distribution: Known from the western United States, and previously reported from Arizona, California, Colorado, Idaho, and Oregon. Also reported from La Paz, lower Baja California.

Material Examined: CALIFORNIA, San Bernardino Co.: exact location unknown, 1922, S.S. Towne (Lloyd Mycological Collection, No. 31027; BPI706815; TYPE). 

            Comments:  This is a North American species, described from material collected in Southern California. CG Lloyd (1923) states that this fungus is “resembling in a general way Bovista pila,” and writes that he received a collection from Claremont, California, from DL Crawford, but it was very immature so he ignored it at first. When he received the material from SS Towne, CG Lloyd noted that based on spore and capillitium structure, it was in between Catastoma and Bovista and should be a new genus, and tentatively placed it in Catastoma because it was very close in morphology to Catastoma luteolum. Twenty years later, Zeller transferred this species to Abstoma. Abstoma townei remains unique amongst the other true puffballs due to small gasterocarp size and pitted spore characteristics. The ITS sequence data suggests that this species is outside of the Lycoperdaceae clade, and closer related to Mycenastrum corium, which is another puffball with pitted spores. Without further collection of material at all ranges of maturity and from various reported locations in North America, or comparing additional herbarium material identified as this species, Abstoma townei will remain incompletely understood.

 

BOVISTA Pers.:

Bovista aestivalis (Bonord.) Demoulin, Beih. Sydowia 8: 143. 1979.                                                                                                                                                            (fig. 4, 40, 41, 44)

Basionym: º Lycoperdon aestivale Bonord., Handb. Allgem. mykol. (Stuttgart): 251. 1851.

Reported synonyms:

            = Lycoperdon furfuraceum Schaeff., Fung. bavar. palat. nasc. (Ratisbonae) 3: tab. 294. 1770.

            = Lycoperdon lacerum Batsch, Elench. fung., cont. prim. (Halle): 145. 1783.

            = Lycoperdon cepiforme Bull., Annales de l'Institut agronomique de Moscou: 156. 1791. var. cepiforme.

            = Globaria furfuraceum (Schaeff.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 2 5: 370. 1873.

            = Utraria furfuracea (Schaeff.) Quél., Enchir. fung. (Paris): 241. 1886.

            = Bovista cepiformis (Bull.) Massee [as 'cepaeformis'], Ann. Bot., Lond. 2(5): 65. 1888.

            = Lycoperdon furfuraceum var. ellipsosporum Lloyd, in Torrend, Brotéria, sér. bot. 11: 92. 1913.

            = Lycoperdon furfuraceum var. elongatum Torrend, Brotéria, sér. bot. 11: 91. 1913.

            = Globaria cepiformis (Bull.) Cout., Eubasid. Lusitan.: 166. 1919.

            = Lycoperdon furfuraceum f. sulphureum Hruby, 1930.

            = Lycoperdon ericetorum var. cepiforme (Bull.) Bowerman [as 'cepaeforme'], Can. J. Bot. 39: 364. 1961.

            = Bovista aestivalis (Bonord.) Demoulin, Beih. Sydowia 8: 143. 1979. var. aestivalis.

            = Bovista aestivalis var. perverrucispora A. Ortega & Buendía, Cryptog. Mycol. 10(1): 16. 1989.

            = Lycoperdon pusillum sensu auct. NZ; fide NZ fungi 2008.

            Type: According to Kreisel (1967b), a lectotype is located in the National Herbarium Nederland at Leiden in the Netherlands (L 910.262-767). The type locality is in Germany.

            Gasterocarp 10-35 mm tall x 12-40 mm broad; slightly obpyriform to compressed globose, often plicate at the base; rhizomorph up to 5 mm thick at the base where attached to the gasterocarp x 10 mm long, incrusted with vegetal debris and soil, rooting the base of the fruitbody to the soil, usually as a single thick strand with fine branches, or a tuft of compact mycelium made with tightly woven thin hyphal hairs, sometimes attaching several fruitbodies in a caespitose manner, eventually breaking off at the soil level to roll about the landscape; ostiole up to 4-5 mm broad at maturity, developing at the apex with the sloughing off of the exoperidium, irregularly torn to slit-like or circular, persistent, sometimes slightly upturned at the rim. Exoperidium cream white when young, becoming pale yellow white (3A2) to buff pink (5A3), often with pink grey tones near the apex (7A2), sometimes a greenish yellow (4D4), becoming buff beige (5C5) to brown gray (5C2) with maturity and sun exposure, often colored in a patchy or scale-like pattern; thin, seemingly glabrous at the apex, furfuraceous-verrucose to subflocculose-granulose at the base and in between folds, readily rubbing off, ornamentation obvious under a dissecting scope but not to the naked eye, wrinkled to folded near the base where plicate, sometimes with an areolate pattern when sun-exposed, cracking with desiccation to form a scale checkered-like pattern and slowly sloughing off in flakes, wearing away to expose the endoperidium with maturity. Endoperidium yellowish grey when young (3B2) to dull yellow (3B3), light gray (5B2), turning grayish orange (5B3), to brownish yellow (5C4-7) to olive brown when dry (4E5) and often a darker red brown (8F6) at the base with maturity; thin, glabrous to smooth, parchment-like, persistent, dull or with a metallic sheen when very mature. Gleba white when young (5A1), turning grayish yellow (4B3), to brownish orange (5C4), then olive green (4F7), to dark chocolate brown (6F4) at maturity; firm and solid when young, cottony in texture with age, maturing from the center outward with the lower portions maturing last. Subgleba yellow cream buff (5A3) to brownish orange (5C3); cellular and composed of compact cells in smaller specimens, greatly reduced or absent in larger specimens as if stretched thin from growth, comprising only the lower portions of the gasterocarp when present. Diaphragm absent.

            Basidiospores globose to subglobose; 3.5-6 X 3.5-5 µm [xmr = 4.5-4.9 X 4.4-4.5 µm, xmm = 4.8 ± 0.1 X 4.4 ± 0.1 µm, Q = 0.8-1.2, Qmr = 1.0-1.1, Qmm = 1.1 ± 0.0, n = 25, s = 8]; spores hyaline to golden brown in water mounts, turning bright green in KOH; seemingly smooth or slightly asperate under light microscope, ornamentation can be seen under the light microscope using the lactic acid cotton blue reaction, with SEM spores mid to large asperate warts, spaced apart and connected by thin rail-like bridges; central oil drop present; spores thick-walled; pedicel short and rudimentary, up to 0.8 mm long; free-floating sterigmata not present in wet mounts; spores mostly of equal size under the light microscope. Eucapillitium Intermediate-type, threads 2-12 mm broad with walls 0.5-1.5 mm thick; hyaline to dark brown in water mounts, golden yellow to brown-green yellow in KOH, thin threads hyaline, thicker threads pigmented; elastic, breaking evenly where severed; glabrous to finely incrusted on thicker strands, sparse to frequent dichotomous branching, threads mostly straight, rarely undulate, sometimes subundulate in thinner threads, with knob-like projections; tips attenuate to fine rounded ends; Lycoperdon-type sometimes found near the inner endoperidium wall in mature specimens. Small to medium-sized punctate pores abundant and up to 1 µm wide, pores more abundant on larger capillitial threads. Septa present and rare, pseudosepta present. Paracapillitium absent. Exoperidium textura angularis; composed of hyaline, irregular-shaped, tightly packed, inflated, thin-walled cells. Endoperidium textura intricata; composed of septate, thick-walled, tightly woven hyphal threads.

Habitat: A common puffball that is terrestrial and dispersed in open grassy terrain. Rarely found solitary, growing in numerous scattered clusters to gregarious or caespitose, and sometimes growing in ferry rings in wet years. Often fruiting in sandy, well-drained soil. Bovista aestivalis will fruit in mixed conifer forest, under Aspen (Populus tremuloides), in the Sonoran Desert Scrub, and with coastal oak species; suggesting a non-mycorrhizal ecology. Among the Geographic Subdivisions of California, this species can be found in the Cascade Range, the Modoc Plateau, in the Sierra Nevada, and in Central Western California.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, Colorado, Idaho, Indiana, Iowa, Kansas, Massachusetts, Michigan, Mississippi, Nebraska, New Jersey, North Dakota, Ohio, Oregon, Pennsylvania, Utah, Washington, Wisconsin, and Wyoming. Also reported from Germany, Italy, Mexico, Papua New Guinea, Portugal, and Sweden.

            Material Examined: CALIFORNIA, Alameda Co.: Oakland, Knowland Park, terrestrial in open field, 30 November 2011, S.S. Jarvis (SSJ 420); Oakland, Knowland Park, dispersed in grass, 19 April 2012, S.S. Jarvis (SSJ 437). El Dorado Co.: El Dorado, Lime Mine Road and Hwy 50, terrestrial, 30 April 2009, C. Kjeldsen (SSJ 348). Humboldt Co.: Samoa Peninsula, gregarious under juniper, 7 November 1986, HD Thiers (HDT 49496); East Fork Campground, parking lot area, terrestrial in grass, 15 November 2009, S.S. Jarvis (SSJ 366). Lassen Co.: Summit Lake, Big Juniper SR-44, terrestrial, 28 September 1977, W.B. Cooke (UCB1472615)(UC). Marin Co.: Pt. Reyes, terrestrial, 7 February 2009, MycoBlitz (SSJ 285); Pt. Reyes, Five Brooks parking lot, terrestrial, 20 October 2009, MycoBlitz (SSJ 384); Tomales Bay, Shell Beach, 5 March 2010, R. Pastorino (SSJ 403). Mariposa Co.: Yosemite National Park, Carlon Meadow Road, on road, 10 June 2012, R. Pastorino (SSJ 416). Napa Co.: Knights Valley, Foote Ranch, open rolling grass land, 15 November 2010, S.S. Jarvis (SSJ 361). San Mateo Co.: San Francisco Water Shed, scattered in soil under hardwoods, 16 December 1920, HD Thiers (HDT 726). Tehama Co.: Mineral, terrestrial, 1 October 1976, W.B. Cooke (UCB1472589)(UC). Tuolumne Co.: Yosemite National Park, Tuolumne River, on ground with mixed conifer, 10 June 2010, R. Pastorino (SSJ 415).

            Comments: Bovista aestivalis can be overlooked due to its small size and common growth in tall grass. Having a similar morphology and somewhat overlapping in habitat with Lycoperdon dermoxanthum (aka Bovista dermoxantha), these two puffballs can be confused easily. Mature gleba tissue near the endoperidium wall will have Lycoperdon-type eucapillitium in Bovista aestivalis, with Intermediate-type in the center of the gleba. The gleba of Lycoperdon dermoxanthum has both Lycoperdon-type and Intermediate-type in the center of the gleba. Mature gleba tissue from both the center and outer extremities needs to be examined in order to reduce the possibilities of misidentification. In comparison to Bovista aestivalis, Lycoperdon dermoxanthum has a darker green gleba, a longer pedicel on the spores, more coarsely verrucose-ornamented spores, and a subgleba that is almost always absent. Both of these species have similar exoperidium and endoperidium characteristics, both have abundant pores of various size, and spores that are about 3.5-4.5 mm. Misidentification in older specimens is common when gleba tissue is absent. The habitat for both of these species overlaps in California, yet Lycoperdon dermoxanthum seems to be more abundant along the north coast. These two puffballs were recently shown to be unrelated in phylogenetic analysis (Bates 2004, Larsson 2008), regardless of their similar morphology and habitat. As well, the ML tree containing the Lycoperdaceae of California supports this theory that Lycoperdon dermoxanthum and Bovista aestivalis are not closely related. ITS sequence data confirms that the puffballs deemed as Bovista aestivalis are the same as those identified as such from Arizona, and sits within the Bovista clade on the ML tree.

 

Bovista californica Kreisel, Nova Hedwigia, Beih. 25: 224. 1967.                                                                                                                                                                        (fig. 5, 38, 45)

            Type: Three collections labeled as isotypes are at the New York Botanical Gardens Mycological Herbarium, collected by Lee Bonar in 1934 (Bonar 467: NY 64802, 64803, 64804), from the shoreline of Hat Lake, N Slope of Mount Lassen, Lassen National Park, Lassen Co., California. Kreisel reports the holotype as deposited in NCU (Univ. of North Carolina, Chapel Hill). 

            Gasterocarp 4-8 mm tall x 5-15 (20) mm broad; globose, subglobose to depressed globose; more or less flat at the very bottom with a tuft of thin branching mycelia threads, or a mycelia pad, incrusted with particles of soil, remaining sessile and without a cord-like rhizomorph; ostiole slowly developing through cracking or tearing at the apex, slit-like to stellate, or round to ovoid, with upturned edges, slightly raised, fimbriate margin, laceration with pubescent hairs on end, persistent. Exoperidium whitish at first (5A1), turning cream buff (5A3), to grey brown (5C2) at maturity; very thin, tomentose to minutely floccose, to furfuraceous and powdery, fragile ornamentation, wearing off easily, leaving a patch-like mottled pattern with age, fragmenting into flocculose scales, revealing the endoperidium, exoperidium mostly adherent to the endoperidium, and persistent. Endoperidium cream white at first (5A1), turning buff brown to clay color (5C5), then turning dark brown (6F6) at maturity; internally villose with soft tiny hairs, externally glabrous, papery, dull, and persistent. Gleba cream white (4A3), grayish yellow (4B5), to brown buff, Isabella color (5E6), dark brown (6F6) when mature; powdery, releasing spores until a thin, flattened, empty casing remains attached to the soil. Subgleba remaining cream colored (4A2), composed of compact cellular cells, greatly reduced to absent. Diaphragm absent.

            Basidiospores globose, subglobose, to broadly ellipsoidal; (3-) 4-6.5 X (-3.5) 4-6 µm [xmr = 5.1-5.5 X 5.0-5.3 µm, xmm = 5.1 ± 0.4 X 5.1 ± 0.16 µm, Q = 0.9-1.2, Qmr = 1.0, Qmm = 1.01 ± 0.0, n = 20, s = 4]; spores golden to rusty brown in wet mounts; roughened to finely verrucose with flattened warts under light microscopy, roughened irregular pyramidal to round or flattened verrucose warts, with fused tips under SEM; central oil drop present; spore thick-walled, up to 0.8 µm thick; pedicel short, 2-6 µm long, ends roughly broken; free-floating sterigmata not present in wet mounts; spores mostly of equal size under light microscope. Eucapillitium Intermediate-type, 2.0-7.5 µm broad, walls up to 0.8 µm thick; eucapillitium threads golden yellow brown in KOH mounts, non-elastic and readily breaking; glabrous, frequent dichotomous branching, threads straight, with occasional knob-like projections; tips attenuate, often subundulate, and narrow; all or most eucapillitium radiating inwards from the endoperidium and subgleba. Pores rare, round and very small when present. Septa rare and found near the center of the gleba, or absent, with abundant false septa. Paracapillitium abundant, incrusted with cellular debris, thin, hyaline, septate. Exoperidium textura globulosa; composed of densely layered, thin-walled, swollen, globose sphaerocyst cells. Endoperidium textura intricata; composed of thin-walled, septate, tightly intertwined hyphal threads.

            Habitat: Found terrestrial along the climax prairie, on south facing ridge tops with perennial grasses (Festuca and Agropyron), along the fescue-wheatgrass zone of the Sierra Nevada, in humus rich and sandy soil, around granite rock, beneath conifers, and along moist shorelines of alpine lakes. Also collected in riparian zones, in sparse grass with other herbs and moss of moist meadows, especially those near lakes and streams. Among the Geographic Subdivisions of California, this species can be found in the southern parts of the Cascade Range, in the Sierra Nevada, in the Eastern Sierra Nevada, and in the Desert Province.

Distribution: Known from parts of the United States, and previously reported from California, Idaho, Washington, and Wyoming.

            Material Examined: CALIFORNIA, Alpine Co.: Alpine Lake, State Highway 4, gregarious on moist soil near pines, 8 September 1992, H.D. Thiers (HDT 5422). Mono Co.: Elery Lake, below the dam, 2,895 meters (9,500 feet) elev., terrestrial, 5 August 1946, M.N. Hood (UCB733943)(UC). Plumas Co.: Davis Lake, Jenkins cove, on soil in native vegetation, near shoreline, 5 July 2009, F. Stevens (FS 070509); Davis Lake, Jenkins cove, on soil in native vegetation, near shoreline, 6 June 2012, S.S. Jarvis (SSJ 459). Shasta Co.: Lassen Park, Hat Lake, N. slope of Mt. Lassen, along the shoreline, 1 July 1934, L. Bonar (Bonar 467: 64802, 64803, 64804)(NY)(TYPE). San Bernardino Co.: Camp Osceola, terrestrial under conifer, 26 September 1976, J.H. Bruha (UCB1568431)(UC).

            Comments: Bovista californica is a very small and easily missed puffball found in seepage zones of alpine lakes, streams, and meadows of the Sierra Nevada range. It is characteristic of growing in large gregarious clusters, has a slightly raised ostiole with a fimbriate upturned edge that is not quite as defined as a peristome, and a mottled to polka-dot overall look as the exoperidium sloughs off throughout maturation. Macroscopically it has very small round pores on the Intermediate-type eucapillitium walls, abundant paracapillitium, and an exoperidium composed of swollen globose sphaerocyst cells. It is reminiscent of Bovista pusilla (Batsch) Pers., from Europe. This is a very small speckled puffball that does not exceed 15 mm in diameter, and has been described to have a mixed-type gleba; having both Intermediate-type and Lycoperdon-type eucapillitium. Microscopically it does not have pores on the eucapillitium walls, with globose pedicellate spores, and exoperidium cells with club-shaped to spherical terminal cells (Larsson 2009). Equivalent to B. californica, Bovista pusilla has a flat apical stoma that protrudes slightly with lobed recurving edges. To the untrained eye, these two tiny puffballs may look identical.            

            Bovista limosa, described from Greenland, is a small puffball (5-6 mm in diameter), characterized by a prominent, delimited peristome with a fimbriate stoma (Larsson 2009). This species is described to have a peristomal depression, which does not seem to be present in collections of Bovista californica. Bovista limosa has Lycoperdon-type eucapillitium, and spores (3-8 µm broad) with a very short pedicel. Larsson & Jeppson (2009) showed that in parts of Europe, Bovista limosa consists of two species with different ITS sequences, slightly contrasting morphology, variation in ecology, but with overlapping distribution ranges. Consequently it was separated into two species; Bovista limosa and Bovista pusilla. Phylogenetic analysis with the puffballs from California shows that Bovista californica and Bovista limosa are sister to each other in ITS analysis with 99% bootstrap support, suggesting that they may be synonymous. Further studies with additional genes should be conducted on these three species of puffball to help decipher their taxonomic relationships.

 

Bovista pila Berk. & M.A. Curtis, Grevillea 2(no. 16): 49. 1873.                                                                                                                                                                                    (fig. 6, 39, 46)

            Type: The holotype is at the Royal Botanic Gardens, Kew (K-M000054928). There is isotype material at the UC Berkeley Jepson Herbarium (M46700)(UC). The type locality is indicated as United States of America, without a specific location mentioned.

Gasterocarp (18) 35-40 (50) mm tall x 20-40 (45) mm broad; globose to slightly depressed globose, slightly plicate at the base to a pinched point or entirely globose; rhizomorph a single mycelium thread up to 1 mm thick by 2-10 mm long, breaking off at the soil level, often

mycelium threads remain, becoming completely free and detached from the soil to roll about the landscape; ostiole developing through thinning at the apex with fruitbody expansion as it matures, becoming torn open unevenly as a hole at first, becoming a horizontal or stellate laceration with maturity, older specimens developing radial splitting; sometimes difficult to discern the top from the bottom in very old specimens with multiple holes. Exoperidium up to 5 mm thick; white when young (5A1), becoming cream to orange grey (5B2), yellowish grey (4A2, 4B2-3) when mature, mottled and unevenly colored; thin, sloughing off early in maturation and in patches, remaining partially adherent to the endoperidium, upturned edges of patches rolling up, inward, and peeling away in small sheets, not present in very old specimens; powdery furfuraceous granules found within plicate folds at the base when young. Endoperidium up to 1 mm thick; cream white to yellowish white when young (4A2), metal grey brown (6D3), becoming dark brown (6F5, 7F7), to dark grey brown or black brown (6F3-2), unevenly colored, distinguishable by the metallic copper to bronze hues of the mature fruitbody; parchment-like, rigid, persistent, glabrous, no obvious ornamentation other than plicate folds and dimpled depressions in some specimens. Gleba pale cream (4A2-3) when young, to yellow buff beige (4B3), becoming grayish green yellow (4C7), golden to almost orange yellow (4B6), olive green (4E6), becoming grayish brown

(6E3) to dark brown (6F7) when mature; young tissue turning bright yellow in KOH, mature tissue turning green with KOH; cottony and flexible when mature, spores falling away from the capillitium readily; solid and firm when young, having small compact irregular-shaped chambers under a dissecting scope, maturing from the base upwards. Subgleba absent. Diaphragm absent.

            Basidiospores globose to broadly ellipsoidal; (3.2-) 4-5.6 X (-3.2) 4-5.5 µm [xmr = 4.4-4.6 X 4.4-4.7 µm, xmm = 4.6 ± 0.2 X 4.5 ± 0.2 µm, Q = 0.8-1.2, Qmr = 1.0, Qmm = 1.01 ± 0.01, n = 20, s = 10]; dark brown to golden amber brown in water mounts and KOH mounts; glabrous, smooth when immature to roughened verruculose when mature under light microscope, roughened with appressed irregular-shaped knob-like verrucose bumps under SEM; oil drop present; spores thick-walled; pedicel remnant up to 0.8-1 µm long, with clean broken ends, larger spores missing a pedicel; free-floating sterigmata sometimes present water mounts; spores mostly of equal size under the light microscope. Eucapillitium Bovista-type, can tease apart individual threads with the naked eye using tweezers; having thin and thick threads up to (6.4) 8-13 (21.5) µm broad with walls up to 2-4.5 (5.5) µm thick, with thin-walled segments out at the tips, thinner walls incrusted with cellular debris; dark brown in water mounts, golden brown in KOH mounts; elastic when young, individual hyphal threads with abundant ramified dichotomous branching from a central thread; older specimens becoming brittle, short fragile branches breaking apart easily when mounted; knob-like projections present; threads mostly straight along thick strands, somewhat sinuous at the tips along thinner walls, heavily incrusted with cellular debris, tips attenuate radically to a short rounded terminus, some ends tapering to a fine point. Pores clustered, very small, round punctate, found on lighter colored threads with thinner walls. Septa rare or absent. Paracapillitium sometimes present, not abundant, and often difficult to find. Exoperidium textura globulosa; composed of thin-walled globose swollen sphaerocyst cells. Endoperidium a combination of textura intricata and textura epidermoidea; composed of tightly woven thick-walled hyphal units, difficult to discern the cellular composition from the irregular-shaped cells.

            Habitat: Often found under Hesperocyparis macrocarpa (Monterey Cypress), on thick duff, fruiting after rains through spring and summer. Growing on soil in and around herbaceous plants, in corrals and ranch fields thick with manure near the coast, also found growing with Bromus tectorum (cheat grass). In higher elevations, this puffball can be found growing in association with Pinus contorta (Lodgepole pine) duff. Among the Geographic Subdivisions of California, this species is found in Northwestern California, in the Cascade Range, in the Modoc Plateau, in the Great Central Valley, in the Sierra Nevada, and in Central Western California.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, Colorado, Connecticut, Florida, Idaho, Iowa, Illinois, Indiana, Louisiana, Maine, Massachusetts, Michigan, Minnesota, New Hampshire, New York, North Dakota, Ohio, Oregon, Pennsylvania, Tennessee, Utah, Virginia, Washington, Wisconsin, West Virginia, and Wyoming. Reported from Canada: Labrador, New Brunswick, Nova Scotia, Ontario, Prince Edward Island, and Quebec. Also reported from Jalisco, Mexico.

            Material Examined: CALIFORNIA, Alameda Co.: Berkeley, University of California, Berkeley campus, November and December 1899, R.E. Gibbs (UCB506667)(UC). Amador Co.: Silver Lake, solitary in soil under conifers, October 1975, H.D. Thiers (HDT 35045). Del Norte Co.: Crescent City, Mill Creek Horse Trail, 12 October 2009, R. Pastorino (SS J390). Fresno Co.: Fresno, Huntington Lake, Hwy 168 near Mono Hot Springs, seepage area in coniferous woods, September 1965, H.D. Thiers (HDT 13389); Shaner Lake, solitary on soil in conifer forest, June 1967, H.D. Thiers (HDT 19416); Fresno, Huntington Lake, Hwy 168 near Mono Hot Springs, in humus under conifers, September 1975, H.D. Thiers (HDT 34936). Humboldt Co.: Samra Sand Dunes, across from pulp mills, 30 March 1973, D. Largent (DL 5754-2139)(HSU); Samra Sand Dunes, across from pulp mills, 30 March 1973, Baroni (Baroni 1183-5392)(HSU). Lassen Co.: Lassen National Park, Manzanita Lake Campground, under willows (Salix spp.), July 1965, H.D. Thiers (HDT 12918). Marin Co.: Muir Woods National Monument, on forest humus, 21 July 1972, E.E. Morse (UCB638889)(UC); Audubon Canyon Ranch, Galloway Canyon, lodged in dry grass, October 1979, C. Calhoun (Calhoun 79-897); Audubon Canyon Ranch, Galloway Canyon, in mixed woods, on duff, December 1980, C. Calhoun (Calhoun 80-1704); Audubon Canyon Ranch, Galloway Canyon, in mixed woods, on duff, December 1980, C. Calhoun (Calhoun 80-

1719); Mt. Tamalpais, Hwy 1, Muir Beach, terrestrial on soil, 4 January 2008, S.S. Jarvis (SSJ 234); Tomales Bay, Radio Operations Center, on thick Monterey Cypress (Hesperocyparis macrocarpa) duff, 7 February 2009, S.S. Jarvis (SSJ 282); San Rafael, Smith Ranch Rd, 26 January 2010, R. Pastorino (SSJ 395); Point Reyes, Five Brooks Parking Lot, 20 October 2010, R. Pastorino (SSJ 382). Mendocino Co.: Albion Station, parking lot, 1 February 2009, all club foray (SSJ 290). Monterey Co.: Pacific Grove, Asilomar Conference Center, under Monterey Cypress (Hesperocyparis macrocarpa), 19 October 1995, D. Deshazer (SSJ 340). Placer Co.: Cisco, June 1909, W.A. Setchell (UCB506662)(UC). Plumas Co.: Lassen National Park, Drakesbad, Warner Valley, terrestrial in horse pasture, 11 July 1972, W.B. Cooke (UCB463213)(UC); Lassen National Park, Warner Valley to Devil’s Kitchen, terrestrial in horse pasture, 9 September 1975, W.B. Cooke (UCB463242)(UC); Lassen National Park, Devil’s Kitchen, 9 September 1975, W.B. Cooke (UCB463210)(UC). Sacramento Co.: Sacramento, American River, on sandy soil, 11 March 1930, J. Novelli (UCB507222)(UC). San Francisco Co.: San Francisco, Golden Gate Park, 15 December 1901, W.A. Setchell (UCB506665)(UC); San Francisco, Cliff House, under dense Monterey Cypress (Hesperocyparis macrocarpa), 6 July 1941, E.E. Morse & V. Miller (UCB659989)(UC); San Francisco, Harding Golf Course, on soil, October 1979, class collection (S.N.). Sierra Co.: Sierra Nevada Field Campus, Hwy 49, solitary under conifers, May 1984, H.D. Thiers (HDT 47663); Yuba Pass, Hwy 49, in meadow, 5 June 2008, S.S. Jarvis (SSJ 227); Chapman Creek Campground, Hwy 49, terrestrial, 5 June 2008, class collection (SSJ 228); Yuba Pass Campground, Hwy 49, terrestrial, 6 June 2012, S.S. Jarvis (SSJ 461). Sonoma Co.: Sebastopol, grassland, 1 March 2008, D. Deshazer (SSJ 244); Geyserville, 7 April 2009, C. Kjeldsen (SSJ 346). Petaluma, Two Rock, Magic Ranch, 20 May 2009, S.S. Jarvis (SSJ 350). Stanislaus Co.: Sonora Pass, buried in soil under conifers, June 1966, H.D. Thiers (HDT 17108). Tehama Co.: Mineral, Battle Creek Meadows Ranch, on soil in mixed conifer woods, 28 August 1974, W.B. Cooke (UCB463211)(UC); Mineral, Mr. Adam’s home, terrestrial in woodland lawn, 20 September 1982, W.B. Cooke (UCB500314)(UC). Tuolumne Co.: Pinecrest Lake, Hwy 108, solitary in humus under conifer, September 1965, H.D. Thiers (HDT 13312); Pinecrest Lake, Hwy 108, 5500 ft elevation, May 1984, H.D. Thiers (HDT 47654).

Comments:  Bovista pila is distinguishable by the metallic copper to bronze-like hues of the mature fruitbody endoperidium. The overall size, and lack of subgleba suggests a Calvatia species. However B. pila has true Bovista morphology as well as a Bovista-type of eucapillitium. Bovista pila is similar to Bovista plumbea, but the latter is smaller, has a tuft of mycelium rather than a well-defined rhizomorph, and lacks the metallic luster that is the hallmark of Bovista pila. Bovista pila is especially common along the coastal regions of San Francisco, Marin, and Sonoma counties. The ITS data here strongly supports this species within the Bovista clade with both 100% bootstrap and 100% PP support.

 

Bovista plumbea Pers., Ann. Bot. (Usteri) 15: 4. 1795.                                                                                                                                                                                                (fig. 7, 38, 47)

Reported synonyms:

            = Lycoperdon bovista Sowerby, Col. fig. Engl. Fung. Mushr. (London) 3: pl. 331. 1803.

            = Bovista suberosa Fr., Syst. mycol. (Lundae) 3(1): 26. 1829.

            = Lycoperdon suberosum (Fr.) Bonord.

            = Lycoperdon plumbeum Vittad., Monograph Lyc.: 174. 1842.

            = Endonevrum suberosum (Fr.) Czern., Bull. Soc. Imp. nat. Moscou 18(2, III): 151. 1845.

            = Globaria plumbea (Pers.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 2 5: 371. 1873.

            = Globaria plumbea var. suberosa (Fr.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 2 5: 371. 1873.

            = Bovista ovalispora Cooke & Massee, Grevillea 16(no. 78): 46. 1887.

            = Bovista brevicauda Velen., České Houby 4-5: 832. 1922.

            = Bovista plumbea var. flavescens Hruby, Hedwigia 70: 350. 1930.

            = Bovista plumbea var. brevicauda (Velen.) F. Šmarda, Fl. ČSR, B-1, Gasteromycetes: 367. 1951.

            = Bovista plumbea var. ovalispora (Cooke & Massee) F. Šmarda, .1958.

            = Bovista plumbea f. brevicauda (Velen.) F. Šmarda, Fl. ČSR, B-1, Gasteromycetes 12: 239. 1958.           

            Type: Bates (2004) states that Kreisel (1967b) selected a lectotype for this species; the Friesian type of Bovista tunicata from the University Uppsala (UPS). Kreisel (1967) lists the lectotype of Bovista plumbea as L910-262-767 (Leiden), non vidi, designated by Perdeck in 1950. Kreisel (1962, 1967) lists the type locality for this species as Germany without a specific location.

            Gasterocarp (5) 10-25 mm tall x (10) 15-40 mm broad; globose to subglobose or depressed globose, entire fruitbody becoming flat in old age, sometimes plicate at the base with folds and wrinkles; rhizomorph composed of a tuft of fine mycelium, incrusted with vegetal debris, sometimes inconspicuous or up to 6 mm wide, becoming easily detached late in maturity to roll about the landscape; ostiole forming from an apical tear, nearly circular, rim not raised and without uplifted edges, with felted hairs protruding from the torn edges, remaining small throughout maturation. Exoperidium pure white to cream colored (4A2) when young, yellow grey brown (5D3), to light brownish orange (5C4); up to 0.5 mm thick when young, becoming thin and papery with age, semi persistent; minutely tomentose to scurfy granulose with depressed scales entirely when young, scurfy granulose scales remaining within plicate folds at the base throughout maturity until very old; cracking with desiccation and sun exposure, sometimes leaving an areolate pattern with age as the exoperidium wears off, flaking off or peeling off in small sheets, often with scale remnants at the base of old specimens, sometimes incrusted with vegetal debris at the base. Endoperidium white when young (6A2), cream (4A2) to grayish brown (6E3), or mousey grey (6E2) at the apex, to dark brown (6F5) throughout, silvery red gray (7F4) at the base in old specimens, sometimes with a cream orange white (6A2) colored patch near the base where the rhizomorph was attached, mottled with uneven patchy coloration, often lighter at the apex when sun exposed; parchment-like, membranous to glabrous, sometimes with vegetal debris adhering to the bottom surface, becoming rigid and persistent with maturity. Gleba cream colored  (4A3) to light yellow (4A4), then greenish yellow (4C7), olive green (4E5), brownish beige (6E3), and dark chocolate brown (6F5) to reddish rusty brown (7F6) when mature; firm and solid when young, becoming cottony then pulverulent with maturity. Subgleba absent. Diaphragm absent.

            Basidiospores globose to subglobose, or broadly ellipsoidal; 4-6 (-9) X 4-5.6 µm [xmr = 5.3-5.8 X 4.7-5.1 µm, xmm = 5.6 ± 0.2 X 4.9 ± 0.2 µm, Q = 0.9-1.4, Qmr = 1.1-1.2, Qmm = 1.2 ± 0.1, n = 25, s = 10]; spores golden yellow in water mounts, yellow green in KOH mounts, spores remaining golden brown after heating with cotton blue, pedicels turning light blue; spores nearly smooth to verrucose under light microscope, covered in small roughened verrucose knobules under SEM; central oil drop present in most collections; spores thick-walled; pedicels long, up to 4-9 (14) µm, straight with an attenuate pointed terminus; free-floating sterigmata sometimes present in wet mounts, absent in very immature or very old material; spores of equal size under light microscopy. Eucapillitium Bovista-type; strands up to 8-16 (22.5) µm broad, walls 3-4 (5) µm thick, thinner branches 2-4 µm broad, with walls up to 0.8 µm thick; elastic and flexuous; having a central strand and branches that taper to thinner strands, thinner threads sinuous; most eucapillitium incrusted with cellular debris; abundant dichotomous branches often swollen at branch joints; rapidly tapering to a blunt end in thicker threads, some tips rounded with a swollen terminus, some tips attenuate to a long fine point in thinner threads. Pores present, not abundant, observed only with SEM; pores seen on eucapillitium, pedicels, and spores; very small and hidden under incrusted cellular debris and spore ornamentation. Septa absent, pseudosepta present and scarce. Paracapillitium absent. Exoperidium textura globulosa; composed of swollen thin-walled globose sphaerocyst cells. Endoperidium textura intricata; tissue composed of thin-walled hyphal strands, septate, fragile, tightly woven and difficult to discern.

Habitat: A common puffball often found in moist grass, in dispersed grass on soil, in meadows or open fields, and near bodies of water; along lake banks, and along stream banks. Solitary to gregarious, sometime in a ferry ring in wet years, scattered in grass, or solitary on soil. Commonly found on golf courses with regular irrigation, and year-round along the central coast of California in irrigated lawns and cemeteries. Among the Geographic Subdivisions of California, this species is found in Northwestern California, in the Cascade Range, in the Great Central Valley, along the Sierra Nevada, in Central Western California, and in Southwestern California. 

Distribution: A global puffball and known from many parts of the United States. Previously reported from Alaska, Arizona, California, Colorado, Connecticut, Georgia, Idaho, Indiana, Illinois, Iowa, Kansas, Louisiana, Maine, Massachusetts, Michigan, Minnesota, Mississippi, Nebraska, Nevada, New Jersey, New Hampshire, New Mexico, New York, North Dakota, Oklahoma, Ohio, Oregon, Pennsylvania, South Dakota, Tennessee, Utah, Vermont, Washington, West Virginia, Wisconsin, and Wyoming. Also reported from Austria, Belgium, Britain, Canada, China, Colombia, Costa Rica, Denmark, Dominican Republic, England, France, Germany, India, Italy, Mexico, Morocco, Portugal, Russia, Scotland, Solomon Islands, and Sweden.

            Material Examined: CALIFORNIA, Alameda Co.: Livermore, terrestrial, March 1929, (UCB507225)(UC); Berkeley, terrestrial, 25 February 1872, coll. unknown (UCB457621)(UC); Berkeley, University of California campus, terrestrial, November 1909, N.L. Gardner (UCB445934)(UC); Berkeley, University of California campus, in open area on grass, 7 March 1940, T.T. McCabe (UCB638439)(UC); Berkeley, University of California campus, terrestrial, 12 March 1979, I. Tavares (UCB1570958)(UC). Butte Co.: Jonesville, July 1931, E.B. Copeland, (UCB64723)(UC). Contra Costa Co.: Corral Hollow, no sandy soil in grass, M.L. Bowerman (UCB960377)(UC). Fresno Co.: Huntington Lake, 8000 ft. elev., in moist meadow, 10 August 1944, M.R. Gibbons (UCB61935)(UC). Humboldt Co.: East Fork Campground, 15 November 2009, S.S. Jarvis (SSJ 470). Los Angeles Co.: Los Angeles, in waste ground, May 1897, A.I. McClatchie (UCB152996)(UC); Los Angeles, in waste ground, May 1897, A.I. McClatchie (UCB151979)(UC). Marin Co.: San Rafael, terrestrial, Late 1920s, L. Grady (UCB507224)(UC); Marin, in grass, 27 March 1932, A. Erickson (UCB513456)(UC); Audubon Canyon Ranch, Galloway Canyon, scattered in grass, December 1977, C. Calhoun (Calhoun 77-420); Audubon Canyon Ranch, Picher Canyon, solitary on path in full sun, October 1979, C. Calhoun (Calhoun 79-874); Audubon Canyon Ranch, Volunteer Canyon, scattered in grass, February 1981, C. Calhoun (Calhoun 81-1972); Point Reyes, Bay View Trail, 26 February 2009, R. Pastorino (SSJ 336). Mariposa Co.: Bear Valley Road, J16 Road, 23 March 2009, S.S. Jarvis (SSJ 305); Yosemite, Hwy 140, 5 mile east of Mariposa, old gravel pit, 23 March 2009, S.S. Jarvis (SSJ 306). Mendocino Co.: Mendocino, in grass, 10 March 1962, D. Malloch (s.n.); Mendocino, on ground, 6 November 1967, coll. unknown (UCB499189)(UC); Paul Dimmick Campground, Hwy 128, scattered on soil under Redwoods (Sequoia sempervirens), November 1984, coll. unknown (HS 2221). Merced Co.: East of Merced Falls, Hwy J16, terrestrial, 5 April 1978, D. Baltzo (UCB1569298)(UC). Monterey Co.: San Ardo, in grass, 4 January 1940, E. Morrow (UCB629533)(UC); Hastings Reservation, terrestrial in a pasture, 16 May 1941, J.M. Linsdale (UCB652752)(UC); Hastings Reservation, Jamesburg Route, under Oak (Quercus lobata), 10 October 1942, J.M. Linsdale (UCB695700)(UC). Napa Co.: Napa, grassy slopes on the west side of Napa Valley, 8 May 2009, C. Kjeldsen (SSJ 347); Foote Ranch, on grassy slopes, October 2009, S. Jarvis (SSJ 469); Napa, Napa Valley Golf Course, 18th hole near parking lot, October 2009, S.S. Jarvis (SSJ 399). Orange Co.: N. Santa Ana Mountains, Claymine Canon, 6 February 1929, J.T. Howell (UCB507201)(UC). Placer Co.: Auburn, 1365 ft elevation, terrestrial, Spring 1934, R. Wall (UCB528325)(UC). Plumas Co.: Davis Lake, Jenkins Point, lake side on moist soil, 6 June 2012, S. Jarvis (SSJ 460); Graeagle Lake, 7 June 2012, F. Stevens (SSJ 467). San Bernardino Co.: San Bernardino Mountains, Big Bear Lake, 8000 ft. elev., in damp grassy soil, 5 September 1941, E.P. Krauth (UCB659982)(UC). San Francisco Co.: San Francisco, Golden Gate Park, in grass, 26 March 1905, coll. unknown (UCB439567)(UC); San Francisco, San Francisco State University, on soil under Monterey Pines (Pinus radiata), September 1986,  (HS 3300). San Mateo Co.: Brisbane, Sierra Point Land Fill, on sandy soil, 7 June 2008, MSSF collection (SSJ 230). Santa Clara Co.: San Jose, Guadalupe Mines Road, spring 1918, H.E. Parks (UCB297709)(UC). Shasta Co.: Lassen National Park, Manzanita Creek, 6000 ft elev., terrestrial, 5 July 1963, W.B. Cooke (UCB1318168)(UC). Sierra Co.: Yuba Pass, Hwy 49, solitary in open field of grass, September 1984, Parsoon (Parsoon 75). Siskiyou Co.: Mt. Shasta, 5000 ft. elev., in grass, October 1990, H.D. Thiers (HDT 53359). Sonoma Co.: Cloverdale, 10 miles north of town, 14 March 1963, coll. unknown (UCB457624)(UC); Skaggs Springs Road, solitary on open grassland, 28 November 2008, S.S. Jarvis (SSJ 265); Gunner, Hwy 1, 17 January 2010, SOMA collection (SSJ 373); Petaluma, Two Rock, Magic Ranch, 11 June 2011, S.S. Jarvis (SSJ 458). Stanislaus Co.: Empire, Davison Ranch, in soil in a pasture, 29 March 1940, E.E. Morse (UCB637388)(UC); Oakdale, Kemper Road, 25 December 1969, coll. unknown (UCB466106)(UC); Oakdale, Kemper Road, on ridges of cow pasture, 3 December 1978, E.C. Akers (UCB1500541)(UC). Tehama Co.: Mineral, Battle Creek Meadows Ranch, on the ground, 28 August 1974, W.B. Cooke (UCB474830)(UC); Lassen National Park, Mineral Ranger Station, terrestrial, 2 October 1977, W.B. Cooke (UCB1474837)(UC); South of Lassen National Park, Guernsey Creek Campground, intersection Hwy 36 and Hwy 89, in soil under mixed conifer, September 1989, H.D. Thiers (HDT 52760). Tulare Co.: Pixley, 5 miles east, near vernal pool on soil, 30 April 1967, J.T. Howell (UCB1319940)(UC); Sequoia National Park, Buckeye Flat, on mowed grass, 24 March 2009, S.S. Jarvis (SSJ 311); South Fork Kaweah River, 10 miles south of Three Rivers, on old paved road in grass, 26 March 2009, S.S. Jarvis (SSJ 354). Tuolumne Co.: Yosemite National Park, Tuolumne Meadows, in a moist meadow, 15 August 1952, coll. unknown (UCB164707)(UC); Mariposa, Road J59, 22 March 2009, S.S. Jarvis (SSJ 302).

            Comments:  Bovista plumbea is a common puffball found in urbanized grasslands around the globe, suggesting a non-mycorrhizal ecology. Bovista plumbea is the type species for the genus Bovista, characterized by a small, globose fruitbody, with dark spores, a single apical circular ostiole, Bovista-type capillitium, and is common in grasslands or urban park settings. This species could easily become confused with Lycoperdon dermoxanthum, which resembles a small white to grey Bovista. However, Lycoperdon dermoxanthum differs in having a single rhizomorph chord, an irregularly torn ostiole, shorter pedicels on the spores, and Lycoperdon-type capillitium mixed with Intermediate-type capillitium in the center of the gleba. Bovista plumbea is much more common than Lycoperdon dermoxanthum in California. Bovista aestivalis can also be confused with Bovista plumbea, the former being a common urban grassland species with an overlapping range and some similar macroscopic features. However, it is easy to tell Bovista aestivalis apart from Bovista plumbea using a microscope. Bovista aestivalis has very short reduced pedicels on the spores, and Intermediate-type capillitium, whereas Bovista plumbea has very long pedicels and Bovista-type capillitium. The ITS data here supports this species within the Bovista clade with both 96% bootstrap and 100% PP support.

 

Bovista sierraensis S.S. Jarvis and F. Stevens, sp. prov.

                                                                                                                        (fig. 8, 38, 48)

            Type: The holotype material is at the Harry D. Thiers Herbarium at San Francisco State University, S Jarvis (SSJ 462). The type locality is Plumas Co.: Davis Lake, along Cow Creek, collected by S. Jarvis and F. Stephens, June 2012. 

            Gasterocarp 11-20 mm tall x 14-35 mm broad; globose to depressed globose, mostly flattened at the base to slightly tapered at the very basal attachment point; rhizomorph composed of very fine hyphal threads, incrusted with soil and vegetal debris, remaining attached until very mature, then detaching and rolling about the landscape, sometimes leaving a small hole at the base where once attached; ostiole at first developing at the apex, a small hole with frayed fimbriose upturned torn edges, becoming a stellate-like laceration, expanding through maturation until a frayed stellate-like cavity, top half diminishes and sloughs away until only a frayed bottom half to two-thirds of the gasterocarp remains. Exoperidium white at first (4A2), to yellowish cream (4A3), becoming pink buff (5A3) to orange white (5A2) at maturity, staying fairly light in color; thin, tomentose to furfuraceous with a granular covering when fresh that will rub off readily when handled, incrusted with vegetal debris at the base, cracking to reveal the endoperidium, edges of the cracks lifting and peeling away in patches to expose the endoperidium over the entire gasterocarp, becoming wrinkled and creased when dry. Endoperidium white when young (2A1-4A2), orange white (5A2), becoming yellowish gray (4B2), browning with maturity to an orange gray at the apex and bleaching in the sun (5A3-5B2-5C4), the base turning dark yellowish brown (5F5) or pale brownish gray (6C2), overall grayish brown (6D3) then brown (6E6) or partially dark brown (6E7) when mature, darker at the apex and around the edges of the opening, becoming dull gray with a metallic sheen with sun exposure (5C3-5D3); glabrous, smooth, parchment-like, with uneven color, persistent, eventually torn and split when dry, a thick layer of eucapillitium remains attached to the inner lining of the endoperidium. Gleba white at first, turning cream (4A3) or orange gray (4A2), to a slight greenish yellow (4C7), then light brown (6D5), then dark brown (6F7) to dark brown (6F4-7F4) when mature, remaining a dark chocolate brown at maturity (5F4-7); maturing from the center outwards; firm and chambered when young, becoming crusty or rocky, to partially powdery, to pulverulent when mature. Subgleba absent. Diaphragm absent.

            Basidiospores globose; 4-8.8 (9.6) X 4.8-8 (9.6) µm [xmr = 6.8 X 6.6 µm, xmm = 6.8  ± 0.0 X 6.5 ± 0.1 µm, Q = 0.9-1.2, Qmr = 1.0, Qmm = 1.0. ± 0.0, n = 30, s = 2]; spores golden amber brown in water mounts, turning olive green in KOH mounts; finely verruculose under light microscopy with ornamentation up to 0.8-1.6 µm long with truncate verrucae, ornamentation densely packed with appressed round verruculose bumps connected by ridges seen with SEM; oil drop present in some spores in water mounts; spores thick-walled; pedicel broken, ± 0.8-1.6 mm long, irregularly torn pedicel confirmed with SEM; free-floating sterigmata not present in wet mounts; spores of equal size under light microscope. Capillitium Calvatia-type; up to 4-6 mm broad with walls 0.8-2.5 mm thick; light yellow brown to somewhat hyaline in KOH and water mounts; non elastic to fragile; eucapillitium slightly incrusted with cellular debris, dichotomously branched, mostly straight, some thinner threads sinuous towards the tips, attenuate in thinner hyphal threads, tips rounded, appearing swollen at the terminus in some mounts. Pores absent or scarce, not seen under light microscope or SEM. Septa abundant, breaking up regularly and disarticulating at the septa or cracking and breaking at cracks; pseudosepta present and scarce. Paracapillitium present; turning light blue with heating in lactophenol cotton blue reagent, not easily spotted without the reaction; thin-walled, septate, hyaline, dichotomously branching, somewhat tapered to a blunt pointed terminus. Exoperidium a combination of textura angularis and textura globulosa; composed of inflated hexagonal thick-walled cells, and globose to irregular-shaped sphaerocysts, thin-walled and hyaline, up to 12 µm broad, distinctly a different layer from the endoperidium layer. Endoperidium textura intricata; composed of long intertwined, thin-walled, pigmented, hyphal elements, up to 4 µm broad. Young gleba tissue composed of a matrix of basidia and indescribable cellular tissue; basidia inflated club-shaped cells with long sterigmata attached to young spores; sterigma up to 4 µm long; 4 spores per basidia; easiest seen with lactophenol cotton blue reaction in water mounts.

            Habitat: Collected from the banks of Virginia Lake and Rock Creek in Mono County, and Davis Lake in Plumas County. Found in moist sandy soil along with sparse grass and moss, often growing out of moss. Fruiting in the spring after the snow melts, or after spring rains, and through summer as long as there is plenty of moisture. Staying attached to the soil until very mature, then detaching when dry and rolling about until only a thin empty endoperidium cup remains. Growing in gregarious groups and clusters, but not caespitose. Among the Geographic Subdivisions of California, this species has been found in the Sierra Nevada and in the Eastern Sierra Nevada.

            Distribution: Known only from banks of alpine lakes and streams of the Sierra Nevada and Eastern Sierra Nevada.

            Material Examined: Mono Co.: Virginia Lakes, Virginia Creek Lodge, in a moist meadow, 2,682 meters (8,800 feet) elev., 8 August 2010, F. Stevens (FS 08-08-2010); South of Mammoth, Rock Creek Canyon, moist grass meadow, along trail to Corral Ponds, 2,682 meters (8,800 feet) elev., 8 September 2011, F. Stevens (SSJ 489). Plumas Co.: Clio, Davis Lake, Cow Creek, lake side wetland, in wet sandy soil with sparse grass, 6 June 2012, S.S. Jarvis (SSJ 462)(TYPE).

            Comments: Bovista sierraensis was first collected by Dr. Fred Stevens, in similar habitat to what Bovista californica seems to prefer, although these two puffballs are quite distinct from one another. Bovista sierraensis grows in the Sierra Nevada mountain range around an elevation of 1,737 meters (5,700 feet), and is consistently found at Lake Davis, which is one of the highest elevation lakes in the California Water Project for residential drinking water. This puffball has been collected along the shores of alpine lakes in the Sierra Nevada and further searches around similar lakes in this region should turn up additional collections.

            DNA sequence BLASTn in GenBank shows that Bovista sierraensis could be closely related to the European taxon Bovista graveolens Schwalb., which is a nomen illegitimate, according to Index Fungorum. In comparison, the morphology of Bovista graveolens describes Bovista-type eucapillitium that is thick-walled, lacking pores, and presumably elastic. Kreisel (1967) describes the eucapillitium walls as "centripetal," referring to thickenings, often uneven, on the inside of the eucapillitial wall. The spores of Bovista graveolens have long curving to u-shaped pedicels on smooth to punctate ornamented spores that are 4-5.5 µm broad. Bovista sierraensis has verrucose spores that are about 7 x 7 µm on average, with a very short, straight pedicel. This small Sierra Nevada puffball has Calvatia-type eucapillitium that is fragile with a gleba that becomes pulverulent with maturation, although it does detach from the soil to roll around tumbleweed style, like that of other Bovista taxa. In phylogenetic analysis, Bovista graveolens and Bovista sierraensis are near each other on the tree, without any bootstrap support, and do not seem related. The morphology of Bovista sierraensis is suggestive of a Calvatia species. However, the placement of this it in phylogenetic analysis is within the Bovista clade. This puffball seems unique among other taxa and therefore is suggested here as a new species, Bovista sierraensis.

 

CALBOVISTA Morse:

Calbovista subsculpta Morse ex M.T. Seidl, Mycotaxon 54: 390. 1995.                                                                                                                                                                        (fig. 9, 38, 49)

Basionym º Calbovista subsculpta Morse, Mycologia 27(2): 97. 1935.

Reported synonym

            = Calbovista subsculpta var. fumosa A.H. Sm., Mycopath. Mycol. appl. 26: 396. 1965.

            Type: The holotype specimen, no. 525436, deposited at UC, was collected by E.E. Morse at Soda Springs, Nevada Co., California, at elevation 2,062 meters (6,767 feet), May 7-May 23, 1934. Two isotype collections are deposited at the New York Botanical Garden (NY 123800, 123801).

            Gasterocarp (15) 40-100 mm tall x (15) 51-130 mm broad; obpyriform, globose to depressed globose at the apex and turbinate at the base; rhizomorph incrusted with soil and sand particles, 33-35 (45) mm broad x 20-25 mm long from the base of the fruitbody, remaining attached and sub-rooted to the soil throughout maturation, sometimes with a mat of white mycelium at the base in addition to the rhizomorph, often with two rhizomorphs growing from the base; ostiole becoming open via cracks along the margins of scales, eventually becoming a gaping hole the entire width of the gasterocarp, wearing away with maturity as the peridium layers slough off. Exoperidium yellowish white (4A2) to cream buff (5A3) when young, becoming brownish yellow (5C7), darkening brown with age (5F5), color becoming patchy and partially remaining light in color through maturation; ornamentation composed of robust polygonal warts with a octagonal or hexagonal basal shape, irregular and angular, with blunted tips; (7) 12-22 mm broad x (6) 12-15 mm wide, up to 1 mm thick; scales made of fibrils radiating upward to a raised and pointed or depressed apex, becoming further depressed or flattened with maturity, cracking apart as the gasterocarp swells with growth, staining bright yellow (4A5) in between cracks; scales mostly concentrated at the apex of the fruitbody, erupting irregularly and falling away top downward; scales coriaceous when moist, becoming brittle when extremely dry; exoperidium remaining adherent to the endoperidium throughout maturation, all pyramidal warts eventually sloughing off, exposing the gleba, completely falling apart until only a small remnant-like cup of subgleba remains. Endoperidium white to cream buff (5A3) when young, becoming bright yellow to orange yellow between exoperidium scales (4A6-4B7), dark chocolate brown when mature (5F8-7F8); 1-2 mm thick at the sides and up to 6-8 mm thick at the top; with a floccose to felt-like ornamentation when young, becoming tough and coriaceous with age; remaining adherent to the exoperidium throughout maturation, falling away with the exoperidium scales and exposing the gleba. Gleba white when young (7A1), immediately oxidizing to orange white (5A2) or cream yellow (4A3) when cut open, becoming light yellowish brown (5D6) along the outer edge, tan green (4D8), to olive brown (5F4), to dark brown (5F8); becoming powdery to completely pulverized with maturity. Subgleba broad at the apex tapering to a plicate subhypogeous rooted base, chambered with large locules. Diaphragm absent.

Basidiospores globose to subglobose; 4-5.6 X 4-5.6 µm [xmr = 4.5-4.7 X 4.5-4.6 µm, xmm = 4.6 ± 0.1 X 4.6 ± 0.1 µm, Q = 0.8–1.2, Qmr = 1.0, Qmm = 1.0 ± 0.0, n = 20, s = 4]; amber golden brown in water mounts, turning bright yellow in KOH mounts; verruculose with rounded bumps, finely roughened, with small erecting bumps under compound light microscope, covered in flattened stellate-shaped bumps with roughened verruculose granules under SEM; oil drop present; spores thick-walled; pedicel (0.8-) 1.6-4.8 µm long, unevenly broken to tapered, broken ends with long ripped strips seen with SEM; free-floating sterigmata not present in wet mounts; spores of equal size under light microscopy. Eucapillitium Calbovista-type; 10-12 µm broad, with walls up to 3 µm thick; dark brown in KOH mounts; elastic to subelastic; composed of individual threads having a thick main stem, branching abundant with numerous ramified branching appendages creating an antler-like appearance; threads incrusted, with a coarse or roughened appearance, knob-like projections absent; ends tapering to a short rounded or blunt terminus, some tips rounded to a pointed terminus, some thinner ends attenuate, tips sometimes curved, hooked, or in-rolled. Pores present, but scarce and small, punctate, sinuous slits present due to cracking of the eucapillitial wall from desiccation; slits as described by Kreisel (1997) absent. Septa present and scarce, disarticulating at the septum, pseudosepta abundant. Paracapillitium absent. Exoperidium textura globulosa; composed of large and swollen sphaerocysts, hyaline, some cells with an irregular-shaped or club-shaped, intermixed shaped cells. Endoperidium textura intricata; composed of a palisade of irregular-shaped, long hyphal threads that easily stain blue in a lacto-phenol cotton blue heating reaction. Subgleba robust and chambered with large locules; reduced and compact in small fruitbodies, composed with pseudoseptate cells. Diaphragm absent.

            Habitat: Fruiting in spring after the snow melts in subalpine to alpine areas of the Sierra Nevada. Growing in areas where deep snowdrifts accumulate, on rocky soil, moist soils, or in xeric soil types. Found terrestrial under conifers, in conifer duff, and in full sun with dispersed grass. Growing solitarily or gregariously. Reported as caespitose in some areas. Collected under Pinus ponderosa (ponderosa pine), Calocedrus decurrens (incense cedar), Pseudotsuga menziesii (Douglas fir), and Quercus velutina (black oak). Calbovista subsculpta is well known among these mountain ranges: Cascades, northern Idaho mountains, Olympics, Rocky Mountains, and in the Sierra Nevada (Seidl 1995). Often seen covered with ants, which have been observed drinking morning dewdrops on fruitbodies along the margin of open peridium, then departing covered with spores, aiding in spore dispersal. Among the Geographic Subdivisions of California, this species is found in the Cascade Range, in the Great Central Valley, in the Sierra Nevada, in the Eastern Sierra Nevada, and in the mountains of Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Alaska, California, Colorado, Idaho, Nevada, Oregon, Utah, Washington. Also known from British Colombia, Canada.

            Material Examined: CALIFORNIA, Alpine Co.: Mono National Forest boundary, Carson Pass, road from Silver Lake, 8500 ft. elev., near boulders on soil, 26 July 1937, D. Nelson (UCB568888)(UC). Amador Co.: East of Jackson, Bear River Road, 6000 ft. elev., gregarious in sandy clay soil in fir and pine needles, 31 May 1939, V. Miller (UCB621489)(UC); West Point, occurring in large patches, 18 March 1972, coll. unknown (UCB1445753)(UC). Butte Co.: Merrimac, on soil in full sun, 4 June 1933, E.E. Morse (UCB531942)(UC). Calaveras Co.: Calaveras Big Tree State Park, under Ponderosa Pine (Pinus ponderosa) and Incense Cedar (Calocedrus decurrens), 25 May 1966, H.D. Thiers (HDT 16535); Hwy 4, Big Meadows Campground, gregarious in humus under conifer, 30 May 1966, H.D. Thiers (HDT 16823). El Dorado Co.: Tahoe National Forest, terrestrial, 29 April 1940, E.E. Morse & V. Miller (UCB653754)(UC); El Dorado Forest, Pollock Pines, Bassi Falls, 16 May 2009, H. Curdts (SSJ 324). Fresno Co.: Huntington Lake, 8000 ft. elev., terrestrial, 11 August 1944, L. Bonar (UCB977164)(UC). Mariposa Co.: Yosemite National Park, Yosemite Village, Camp 14, 4000 ft. elev., on sandy soil and grass, 20 April 1940, E.E. Morse & V. Miller (UCB637360)(UC). Mono Co.: Leavitt Creek, East side of Sonora Pass, 8500 ft. elev., 4 August 1941, P. Parel d Hui (UCB695856)(UC); Leavitt Creek, East side of Sonora Pass, 7500 ft. elev., 4 August 1941, P. Parel d Hui (UCB139014)(UC). Nevada Co.: Tahoe National Forest, Hwy 20 near Whitecloud Campground, solitary in soil under Pseudotsuga menziesii (Douglas Fir), 25 April 1993, D.E. Desjardin (DED 5663). Plumas Co.:  Bucks Lake, SW side of the lake, on grassy hill slopes, 11 June 1942, V. Miller (UCB671389)(UC); Lassen National Park, Warner Valley, Boiling Springs Lake, terrestrial, 10 July 1966, coll. unknown (UCB1463164)(UC); Lake Almanor, Gurnsey Creek Campground, hwy 36, terrestrial, 31 May 2008, S. Jarvis (SSJ 203); Lake Almanor, Gurnsey Creek Campground, Hwy 36, under conifers in duff, 31 May 2008, S.S. Jarvis (SSJ 204). San Bernardino Co.: San Bernardino Mountains, Buff Lake, 28 June 1920, coll. unknown (UCB472084)(UC); San Bernardino Mountains, Glass Road, Hathaway Creek, 7000 ft. elev., 7 October 2008, S.S. Jarvis (SSJ 253); San Bernardino Mountains, Glass Road, 6500 ft. elev., 7 October 2008, S. Andrasko (SSJ 256). San Mateo Co.: San Francisco Watershed, Montana Ridge Road, Chaparral Ridge, 25 April 1969, F. Stevens (Setzer 2184). Sierra Co.: Yuba Pass, Camp Leonard, gregarious in humus under mixed conifer, 9 June 1964, H.D. Thiers (HDT 11219); Yuba Pass, Camp Leonard, solitary in soil under conifers, 5 June 1965, H.D. Thiers (HDT 12592); Yuba Pass Campground, gregarious in soil under Abies magnifica (Red Fir), 29 July 1971, H.D. Thiers (HDT 27523); Yuba Pass, Hwy 49, gregarious in soil under conifer, 10 July 1986, H.D. Thiers (HDT 49916); Yuba Pass, Hwy 49, Solitary in soil under conifer, 1 September 1989, H.D. Thiers (HDT 52624); Yuba Pass, Green Acres, Gold Lake, 3 June 2008, S.S. Jarvis (SSJ 223); Yuba Pass, Hwy 49, Cal Trans parking area, terrestrial in full sun, 5 June 2008, S.S. Jarvis (SSJ 226); Sierra Nevada Field Campus, Yuba Pass, Hwy 49, 7 June 2008, S.S. Jarvis (SSJ 229); Yuba Pass Summit Campground, Hwy 49, 8 June 2008, D. Hemmes (SSJ 218); Yuba Pass Summit, Hwy 49, on open ground, 8 June 2008, S.S. Jarvis (SSJ 220). Siskiyou Co.: Mc Cloud, Mt. Shasta, 12 miles below Medicine Lake, in open pine forest, 20 July 1928, E.E. Morse (UCB531964)(UC); Mt. Shasta, Bear Springs Road, terrestrial in chaparral forest, 22 June 1946, coll. unknown (UCB139010)(UC); Mt. Shasta, on soil in full sun, Fall 1960, (UCB209301)(UC); McCloud, Hwy 89, 23 June 2009, R. Pastorino (SSJ 331). Tehama Co.:  Feather River, Battle Creek meadows, terrestrial, June 1927, E.E. Morse (UCB531931)(UC); Mineral, Bodine Place, 5000 ft. elev., on the ground, 23 September 1983, coll. unknown (UCB1514582)(UC); State Hwy 36, Gurnsey Creek Campground, 5500 ft. elev., gregarious in soil under conifers, 7 May 1992, H.D. Thiers (HDT 54156). Tulare Co.: Sequoia National Park, growing in sand on a creek bank, June 1927, E.E. Morse (UCB531962)(UC); King’s Canyon National Park, General Grant Park, in open forest, 21 June 1931, E.E. Morse (UCB531959)(UC); Sequoia National Park, High Sierra Trail, Bear paw Meadow, in sandy soil and semi-open forest, 23 May 1940, V. Miller (UCB638340)(UC). Tuolumne Co.: Eagle Meadow, under Lodgepole Pine, Pinus murrayana, 27 August 1915, L. Bonar (UCB532342)(UC); Stanislaus National Forest, Dodge Ridge Ski area, scattered on soil in open area, 14 June 1964, H.D. Thiers (HDT 11227); Pinecrest area, scattered to solitary in humus under mixed conifers, 14 June 1965, H.D. Thiers (HDT 12624); Hwy 108 near Long Barn, solitary in soil under Firs, 25 April 1986, H.D. Thiers (HDT 49593); Tuolumne County, 1 May 2009, R. Pastorino (SSJ 335).

            Comments:  Calbovista subsculpta was described by E.E. Morse based on observations and collections made over nine years of study (Morse 1935). Prompted by a collection of Rocky Mountain Calvatia sculpta with a label reading “excessively branched eucapillitium,” E.E. Morse began to re-examine all collections of Calvatia sculpta in her possession. It was discovered that one-half of the total Calvatia sculpta collections had this same type of eucapillitium character. Further investigation warranted the erection of a new genus of puffball, Calbovista subsculpta. Morse (1935) describes in detail why Calbovista subsculpta did not belong in Bovista, Bovistella, Calvatia, Mycenastrum, or Scleroderma, having a mix of characters. The original description states that “a puffball which bridges the gap between two large, distinct groups of puffballs,” (Morse 1935) contains bits and pieces from each of these genera listed above. Zeller (1948) erected the family Mycenastraceae to include Mycenastrum and Calbovista based on the nature of the highly ramified branching character of the eucapillitium. However, Seidle (1995) explains that Calbovista is accepted in the Lycoperdaceae family by many mycologists. In 1965 A.H. Smith described a variety of Calbovista subsculpta as Calbovista subsculpta var. fumosa, based on an exoperidium with gray coloration throughout maturation and reduced wart-like scales. This is in comparison to the robust wart-like scales and cream to tan brown or dark brown tones of mature fruitbodies found in Calbovista subsculpta var. subsculpta. This gray toned variety of Calbovista should be further studied with DNA sequence analysis to clarify its taxonomic position. The ITS data here supports this species within the Lycoperdon clade with only 81% bootstrap and less than 70% PP support. This suggests that multi loci gene analysis will be necessary to further clarify the taxonomic position of Calbovista subsculpta within the Lycoperdaceae. 

 

 

CALVATIA Fr.:

Calvatia booniana A.H. Smith, in Zeller and Smith, Lloydia 27: 164. 1964.                                                                                                                                                            (fig. 10, 39, 50)

            Type: The holotype is located in the University of Michigan herbarium (MICH 65191), from Prineville, Oregon, collected by AH Smith, July 1962.

            Gasterocarp 135-300 mm tall x 270-600 mm broad; depressed globose to ovate or cushion-shaped with a broad flat base; rhizomorph composed of mycelium threads forming a tuft or mat that anchors the fruitbody; ostiole lacking, cracks forming a wide opening from the sloughing off of peridial walls. Exoperidium white when young (5A1-2), turning yellowish cream to pink buff (4A2-3, 5A3); pyramidal scales large, up to 50-60 mm tall x 80-155 mm broad, polygonal to hexagonal-shaped base, with areolate rings circling each scale in very large specimens, striations along the areolate ring pattern, striations forming ridges of scurfy hair-like projections, scales covering the entire exoperidium surface with the largest on the upper most surface, tips of the scales appressed and turning brownish gray (6C3) to brownish orange (5C4) with sun exposure and age, scales eventually sloughing off with desiccation and maturation to reveal the gleba; subfloccose or somewhat delicately cottony when young, adhering to soil and vegetal debris at the base with glass-like hairs protruding from the exoperidium, entangled with crystallized web-like formations, the base cracking into a semi-random circle pattern, exoperidium remaining adherent to the endoperidium. Endoperidium cream-white (4A1-3) when young, becoming dull brownish gray with age (6C3); parchment-like and thin, 1-2 mm thick, persistent and remaining attached to the exoperidium, sloughing off with the scales throughout maturation. Gleba cream white when young (4A1-3), becoming grayish yellow (4B5), then olive-green with age (4E5); maturing from the center outwards, firm and solid when young, becoming chunky to pulverized powdery with age. Subgleba absent. Diaphragm absent.

            Basidiospores globose to broadly ellipsoidal; 3.9-5.4 X 3.9-4.7 µm [xmr = 4.9-4.9 X 4.6-4.7 µm, xmm = 4.9 ± 0.1 X 4.6 ± 0.1 µm, Q = 0.8-1.4, Qmr = 1.1-1.1, Qmm = 1.1 ± 0.0, n = 20, s = 3]; spores golden brown in water mounts, amber brown in KOH, spores do not stain blue in cotton phenol reaction; spores smooth to very finely roughened under light microscopy, under SEM spores asperate with rounded knobby bumps; no oil drop; spores thick-walled; pedicel up to 1.6 µm long, broken cleanly, walls up to 0.8 µm thick, hyaline in wet mounts; free-floating sterigmata not present in wet mounts;  spores of various shape in light microscope and SEM, some spores of uneven shape. Calvatia-type eucapillitium; capillitium threads 3-9 µm broad with walls 0.8-1.0 µm thick; threads golden brown in wet mounts; fragile, becoming short strands as threads disarticulate; threads glabrous, dichotomous branching common, knob-like projections absent, straight threads, attenuate to round and blunt terminus. Pores present, very small, round, clustered on larger capillitium threads. Septa and pseudosepta present. No paracapillitium. Exoperidium textura globulosa; composed of hyaline, thin-walled, irregular-shaped sphaerocyst cells. Endoperidium textura intricata; composed of a network of intertwined hyphal elements.

Habitat: Terrestrial. Often found growing solitary or in small groups of 2-3 individual fruitbodies. A rare species, coming up in late spring in wet years after heavy winter snowfall. Collected along mountain dirt roads in compact soil, along cow trails, in grassy meadows, at lower coastal elevations in wet fall years, and at higher Sierra Nevada elevations in spring. Collected among Bromus tectorum (cheat grass ), Salix spp (under willows), with Juniperus spp.  (Juniper), under Populus tremuloides (poplar trees), with conifers, found in Pinus longaeva (bristle pinecone) habitat, and recorded to be growing with Swietenia mahagoni (West Indies Mahogany). Among the Geographic Subdivisions of California, this species is found in the Great Central Valley, in Central Western California, and in the high elevations of the Desert Province.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, Colorado, Idaho, Kansas, Michigan, Montana, New Mexico, Oregon, and Utah. Also reported from Northern regions of Mexico.

            Material Examined: CALIFORNIA, Alameda Co.: Oakland, Castle Wood Country Club, under live oak, 14 April 1905, Det. By E.E. Morse in 1933 (UCB506612)(UC). Glenn Co.: Mendocino National Forest, 29 October 1993, D. LeFer (HDT 043). Inyo Co.: Crooked Creak Research Station, 10,000 ft. elev., Pinus longaeva (Bristlecone Pine) habitat, 19 April 2012, T. Bruns (TDB 3185)(UC). Marin Co.: Bolinas, scatted on ground in pasture, October 1981, D.E. Desjardin (DED 512); Audubon Canyon Ranch, Volunteer Canyon, solitary in grass open area, March 1980, C. Calhoun (Calhoun 80-1606). Santa Clara Co.: Henry Coe State Park, scattered in open grassy meadow, April 1986, H.D. Thiers (HDT 49529). Stanislaus Co.: Turlock, Junction of Sante Fe Road and Keyes Road, gregarious in soil in pasture, 29 May 1976, R. Halling (Halling 1382).

Comments: Calvatia booniana is the largest species recorded from California. In some wet years, Calbovista subsculpta may grow to enormous size and could be confused with Calvatia booniana due to similar exoperidium characteristics. However, checking the eucapillitium under a microscope will quickly resolve any questions associated with confusing these two species. Calbovista has ramified branching appendages on the capillitium threads, and C. booniana does not. Calvatia sculpta is another large puffball with exaggerated scales. This species is easily ruled out of a possible misidentification, since it has a prominent subgleba where C. booniana does not. Checking the spores of these species can also aid in correct identification. Calvatia booniana has asperulate spores, Calvatia sculpta has echinulate spores, and Calbovista subsculpta has verrucose spores. The ITS data here strongly supports this species within the Calvatia clade with both 100% bootstrap and 100% PP support.

 

Calvatia fragilis (Vittad.) Morgan, J. Cincinnati Soc. Nat. Hist. 12: 168. 1890.                                                                                                                                                            (fig. 11, 51)

Basionym: º Lycoperdon fragile Vittad., Mem. R, Accad. Sci. Torrino, Ser. 2 5: 180. 1843.           

            ≡ Calvatia cyathiformis f. fragilis (Vittad.) A.H. Smith, Lloydia 27: 150. 1964.

Reported synonyms:           

          = Calvatia lilacina (Mont. & Berk.) Henn., Hedwigia 43: 205. 1845, sensu some authors.

            = Lycoperdon novae-zelandiae Lév., Ann. Sci. Nat., Bot., Sér. 3, 5: 164. 1846.

            = Bovista cinerea Ellis, Bull. Washburn Coll. Lab. Nat. Hist. 1: 40. 1885.

            = Lycoperdon violascens Cooke & Massee, J. Roy. Microscop. Soc. London: 706. 1887.

            = Bovista amethystina Cooke & Massee, Grevillea 16: 69. 1888.

            = Bovista dealbata Berk. ex Massee, J. Bot. 26: 131. 1888.

            = Calvatia lilacina var. occidentalis Lloyd, Mycol. Writings 6: 1097. 1921,

nom. nud.

            Type: A collection from Vittadini is at the Royal Botanical Gardens, Kew (K), labeled as Lycoperdon fragile Vittadini. It was previously thought that this was the holotype collection (Bates 2009 & Kreisel 1992). Confirmation from the herbarium curator states that this collection has been partially annotated to be Calvatia cyathiformis and possibly Calvatia candida. Vittadini (1843) states that the original collections were from Ticino (Italy) and Zurich (Switzerland).

            Gasterocarp 20-70 (100) tall x 35-70 (110) mm broad; compressed globose or subglobose when young, growing into an enlarged obpyriform shape, or an irregular depressed obpyriform shape; with a wide flattened apex, abruptly turbinate, tapering to a pointed lower half, with a reduced plicate and wrinkled base, slightly swollen pseudostipe but not clavate; rhizomorph composed of a thick root with a tuft of mycelium at the base attaching it to the ground, remaining attached to the soil throughout maturity, base partially rooted into the soil, incrusted with soil and vegetal debris; ostiole developing through irregular stellate splits or hexagonal cracks in the peridium layers that expose the gleba, becoming cracked and eroding away until only the pseudostipe and basal attachment of the gasterocarp remains. Exoperidium white when young, turning buff white (5A3) to cream yellow (4A3) when young, gray buff (5B3), with tints of lilac gray cream (14B1), dark chocolate brown to reddish purple dark brown (6F4-8F7), to almost black in some cases when sun exposed, mottled with patches of lilac (14D2) and rusty brown (7E4) when aged; with an uneven surface, minutely furfuraceous, thin and papery, fragile, easily separating from the endoperidium, breaking up into large areolate patch-like scales as the fruitbody expands with growth, patch corners becoming slightly upturned or rolling up, sloughing off in irregular-shaped sheets by peeling away and exposing the endoperidium, sometimes forming an areolate pattern after sloughing off; or remaining adherent to the endoperidium, sloughing off together in sheet-like patches to expose the gleba. Endoperidium white to cream yellow (4A3) when young, turning grey purple violet (14D2-14E3) with maturity, deep rich purple gray (14F3) mottled with shinny coppery bronze (6E7); smooth, 1-2 mm thick, fragile, especially brittle when dry, easily sloughing off in patches; longitudinal laceration initially opens at the apex to reveal the gleba, cracks become stellate, breaking up into smaller plates or scales that slough off throughout maturity, eventually wearing down until only a bowl-shaped pseudostipe remains. Gleba white to at first, quickly turning yellow cream (4A3) then pale green (4A4-5A2), turning gray purple (18D2), then gray purple brown (18F2), dark purple brown when mature (13F3), sometimes a violet gray color (14D2); gleba can be quite stout, up to 10-45 mm tall x 70-110 mm broad; maturing from the center outwards, firm at first, becoming cottony and semi-fragile, then somewhat powdery but remaining in large compact chunks as it matures, becoming pulverulent in old specimens. Subgleba cream color (4A3) to light yellow (4A4) and remaining so throughout maturity; sometimes swollen but not clavate; variable in thickness and in size, up to 5-35 mm tall, tapered and sharply turbinate in a v-shape, up to 10-40 (55) mm broad at the base of the gleba tapering to 1 mm broad where attached to the rhizomorph; well developed but not as a pseudostipe, composed of well developed compact chambers, cellular cottony fibers, subgleba often taking up then entire bottom half of the fruitbody, occasionally absent when the fruitbody remains globose in shape with a small stature (usually due to premature maturation from lack of moisture and direct sun exposure).

            Basidiospores globose; 5.5-7.2 (-8) X 5.5-7.2 (-8) µm [xmr = 5.7-6.0 X 5.8-6.0 µm, xmm = 5.8 ± 0.1 X 5.9 ± 0.1 µm, Q = 0.9-1.1, Qmr = 0.9-1.0 Qmm = 0.9 ± 0.0, n = 25, s = 3]; pale brown to rusty brown in wet mounts; seemingly smooth with immature collections, mature spores finely echinate to verrucose under light microscope, with SEM spores densely echinate with spinose protrusions connected by ridges; ornamentation up to ± 0.8 µm from the spore wall; central oil drop absent, too ornamented for determination; spores thick-walled; pedicel absent or rudimentary and with an irregular torn end; free-floating sterigmata not present in wet mounts; spores variable in size under light microscope. Eucapillitium Calvatia-type; 4-4.5 µm wide, walls 0.5-0.8 µm thick; pigmented golden brown in wet mounts; fragile and fracturing easily; hyphal threads incrusted with cellular debris, dichotomous branching present and not abundant, threads straight to somewhat undulate but not sinuous, some walls with knob-like inward projections, outward knob-like projections present but not abundant; tips somewhat attenuate to an acute terminus, some tips round. Pores abundant, small to medium-sized, punctate to slit-like or stellate under light microscope; using SEM, the slit-like pores appear to be cracks in the eucapillitial walls due to fragility, not due to morphology. Septa common to scarce, disarticulating at the septum readily; pseudosepta present and scarce. Paracapillitium absent. Exoperidium textura epidermoidea; composed of an unorganized, irregular-shaped, tightly packed, inflated, thin-walled cellular matrix of sphaerocyst cells, hyaline or sometimes beige-red pigmented. Endoperidium textura intricata and textura epidermoidea; composed of organized club-shaped septate cells, dichotomously attached; intertwined with heavily pored, branched, thin-walled, tightly woven hyphal threads. Subgleba composed of thick-walled hyphal threads, golden yellow in wet mounts, semi-sinuous, tips with a round terminus, round pores, septate, dichotomously branching, threads up to 2.5-8.5 µm broad with walls up to 1 µm thick. Diaphragm absent.

            Habitat: Growing in fairy rings, in groups, or solitary on grassy hillsides and along hillside drainage points. Found on west facing slopes where sun and wind exposed. Sometimes associated with tan oak (Notholithocarpus densiflorus), madrone (Arbutus menziesii) and redwood (Sequoia sempervirens). Fruiting in gregarious patches the fall after a heavy downpour of rain, or in smaller groups to solitary in spring after rain. Kreisel (1962) notes that this species is cosmopolitan, tropical and subtropical. Among the Geographic Subdivisions of California, this species is found in Northwestern California, in the Great Central Valley, in the Sierra Nevada, in Central Western California, and in the San Bernardino Mountains of Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, Colorado, Georgia, Idaho, Oregon, Tennessee, and Wyoming. Also reported from Italy, Switzerland, and in the Australian Capital Territory, Australia.

            Material Examined: CALIFORNIA, Alameda Co.: Berkeley, 19 June 1894, J.B. Davy (UCB506623)(UC); Berkeley Hills, Fall 1900, R.E. Gibbs (UCB506619)(UC); Berkeley, Fall 1913, coll. unknown (UCB528273)(UC); Manzanita, Berkeley hills, Autumn 1927, E.E. Morse (UCB506675)(UC); Oakland, Sequoia Park, open hillside, caespitose & abundant, 9 November 1941, coll. unknown (UCB62404)(UC); UC Berkeley campus, South-West of Mulford Hall, December 1976, T. Tang (UCB1466089)(UC). Amador Co.: Jackson, experimental station, Grounds A, 23 November 1901, coll. unknown (UCB506674)(UC). Contra Costa Co.: Mt. Diablo, terrestrial, 2 May 1935, V. Miller (UCB532392)(UC); East Bay Regional Park, Morgan territory, 16 March 1992, M.T. Seidl (UCB1598686)(UC). El Dorado Co.: Diamond Springs, 8 miles East, red dirt hillside, 792 meters (2,600 feet), 23 October 1937, E.E. Morse (UCB589799)(UC). Lassen Co.: Susanville, Rice Canyon, Willow Creek, 1,371 meters (4,500 feet) elev., under Pinus jeffreyi (Jeffery pine), (s.n.). Marin Co.: Tiburon, grassy hillside, 13 December 1922, coll. unknown (UCB506541)(UC); Dog Town, Copper Mine Gulch, Highway 101 junction, terrestrial in grass, December 1988, (SFSU S.N.); Samuel P. Taylor State Park, terrestrial in a horse pasture, 10 June 2009, J. Hollinger (SSJ 323). Mariposa Co.: on an open slope above chaparral, S.W. facing dry slope, 30 November 1974, C. Calhoun (Calhoun 367). Merced Co.: Merced, 27 February 1934, coll. unknown (UCB521330)(UC). Monterey Co.: Hastings Natural History Reservation, East side of Carmel Valley, open sunny hillside along an old country road, 29 April 1967, T. Tang (UCB1569962)(UC). Napa Co.: St. Helena, hillside, 2 October 1899, Mrs. D.O. Huset (UCB506680)(UC); Knights Valley, Highway 128, South side of Foote Ranch, border of Sonoma County, 16 November 2008, S.S. Jarvis (SSJ 257). Placer Co.: Auburn, terrestrial, December 1934, E.E. Morse (UCB528273)(UC). Sacramento Co.: Sacramento Junior College, in lawn of Athletic field, 17 July 1938, E.E. Morse (UCB605290)(UC). San Bernardino Co.: plains near Coraona, junction of San Bernardino, Riverside, and Orange Counties, February 1909, C.M. Wilder (UCB506664)(UC); Arrowhead Hot Springs, Cold Water Canyon, 28 May 1930, N.L. Gardner (UCB661749)(UC). Santa Clara Co.: San Jose vicinity, terrestrial, 25 September 1918, H.E. Parks (UCB369444)(UC). Santa Cruz Co.: Boulder Creek, on bare sparse grass, 14 April 1940, V. Miller (UCB638433)(UC). San Mateo Co.: San Mateo, Junipera Serra County Park, scattered in field, 27 November 1980, H. Pats (DLargent 168). Sonoma Co.: Healdsburg, 4 December 1942, V. Miller (UCB695860)(UC); Atlas Peak, Milliken Reservoir, Late November 1977, T. Tang (UCB1473486)(UC); Skaggs Springs Road, open grassland, 21 November 2008, S.S. Jarvis (SSJ 264); Petaluma, Magnolia Street, terrestrial in grass, Fall 2009, D. Deshazer (SSJ 333); Knights Valley, Foote Ranch, Highway 128, open ranch land, South-West facing slopes in full sun on low lying grass, 15 November 2009, S.S. Jarvis (SSJ 386); Petaluma, cemetery in lawn, December 2009, D. Deshazer (SSJ 371); Sebastopol, Pleasant Hill Road, on lawn, 9 February 2010, D. Deshazer (SSJ 334); Knights Valley, Foote Ranch, gregarious on west facing hillside, 16 October 2011, S.S. Jarvis (SSJ 409)(EPITYPE); Knights Valley Foote Ranch, terrestrial in Douglas fir (Pseudotsuga menziesii) grove, in a fairy ring in full sun, 20 October 2011, S.S. Jarvis (SSJ 412); Petaluma, Petaluma Catholic School, Highway 128, terrestrial in back baseball field, 3 November 2011, D. Deshazer (SSJ 422); Knights Valley, Foote Ranch, Highway 128, open field on flat land with Pseudotsuga menziesii (Douglas fir), 5 November 2011, S.S. Jarvis (SSJ 387). Yuba Co.: Brownville, on soil, no date, George Haltz, (Haltz 8407).

Comments:  Calvatia cyathiformis and Calvatia fragilis are two very closely related species, possibly even the same species. Bates (2004) showed in phylogenetic analysis that they form a well-supported monophyletic sister group. They have similar morphological features, except for three main differences: 1) overall size; 2) subgleba size and shape; and 3) subgleba cellular density. Calvatia cyathiformis has a very large and broadly bulbous to clavate-shaped subgleba that is arguably a true pseudostipe, whereas C. fragilis has a narrow reduced subgleba that is more tapered at the end, rooting it into the ground, and not acting as a true pseudostipe. The former will grow up to 80-130 mm tall x 70-130 mm broad, while C. fragilis is almost half the size, 35-60 mm tall x 40-70 broad. Calvatia fragilis has subgleba tissue with smaller chambers and a more compact appearance than Calvatia cyathiformis. Historically Calvatia cyathiformis and Calvatia fragilis have been listed as a forma and as a variety of Calvatia cyathiformis; Calvatia cyathiformis forma/var. cyathiformis and as Calvatia cyathiformis forma/var. fragilis. DNA sequence analysis confirms that they are sister to each other, but with careful examination, the morphology shows that they can be recognized as distinct species (Kreisel 1992, Bates 2004). One specimen in the UC Berkeley herbarium (UCB506673, RP Brandt 1913) was labeled as “Fungi of California,” without any location or habitat data. This specimen is clearly of C. cyathiformis, with a robust pseudostipe up to three inches long. Therefore it could be possible that C. cyathiformis has been collected in California, but very rarely. Calvatia fragilis can be found growing in large annual fairy rings, sometimes up to several hundred feet in diameter. If fairy rings grow “at a few feet per year” (Parker-Rhodes 1955), then fairy rings this large could be up to one hundred years old. In the spring these rings grow very dark green nitrogen rich grass along the margin, which becomes greener than all the grass along the hills of the surrounding area. By mid-fall, when the fungus begins to develop basidiocarps, the grass along the margins of the fairy ring dies back. It looks as if Calvatia fragilis supports the grass through spring, and when it is time to fruit, this fungus could possibly be using the grass as a nutrient source. The ITS data here suggests that both Calvatia cyathiformis and Calvatia fragilis are separate species, but closely related. They are nested inside the Calvatia clade with 100% bootstrap and 100% PP support.

 

Calvatia fumosa Zeller, Mycologia 39(3): 300. 1947.                                                                                                                                                                                    (fig. 12, 42, 52)

Reported synonyms:

            = Calvatia fumosa var. idahoensis Zeller and Smith, Lloydia 27(3): 156. 1964.

            = Gastropila fumosa (Zeller) P. Ponce de León, Phytologia 33(7): 458. 1976.

= Handkea fumosa (Zeller) Kreisel, Nova Hedwigia 48(3-4): 286. 1989.

            Type: A holotype is located at the New York Botanical Garden (NYBG 65666), from Crater Lake National Park, Oregon, collected by F.P. Sipe on July 22, 1943, and determined by S. Zeller.

            Gasterocarp 25-58 mm tall x 22-55 (75) mm broad; irregular globose to subglobose, depressed globose, or obpyriform with a flattened apex; rhizomorph up to 2-5 mm thick, up to 70-130 mm long, tapered, sometimes two coming out of the base, eventually branching & tangled, root-like, white, soil incrusted hyphal threads; ostiole developing as longitudinal to stellate cracking along the edges of peridium scales at the apex. Exoperidium buff white when young, turning pale buff cream color (4A2-3), to brown grey (5C5), to whitish buff yellow (5D3) or white gray in cracks (5A1), with yellowish gray (5B1); exoperidium up to 5-7 mm thick; the top surface cracking into irregular-shaped patches or polygonal scales that eventually crack open and expose the gleba; surface covered with minute fibrillose-scurfy spines that merge at their points and resemble tiny dots or granules to the naked eye, these wear off early in development, base smooth to roughened with many creases and dimples; exoperidium remaining attached to the endoperidium throughout maturation. Endoperidium white when young, turning pale buff cream color (4A2-3), yellow gray (5B1), dark yellowish brown (5F6), then olive green (4E7-4F7) when mature; up to 1 mm thick; glabrous, persistent and dull, adhering to exoperidium throughout maturation, young exoperidium tissue easily removed from endoperidium when manipulated. Gleba chambered and dense when mature, white and firm at first, becoming cream yellow (4A3) to bright yellow (4A5), then olive green grey (4E3), to dark chocolate brown (6F7) when mature; gradually maturing from the center outward, gleba peeling away from the inner walls of the endoperidium easily, becoming pulverized and powdery when mature. Subgleba absent. Diaphragm absent. Odor noxious.

            Basidiospores globose; 4.8-6.4 X 4.8-6.4 µm [xmr = 5.1-5.2 X 5.2-5.3 µm, xmm = 5.12 ± 0.1 X 5.2 ± 0.1 µm, Q = 0.9-1.0, Qmr = 0.9-0.9, Qmm = 0.9 ± 0.0, n = 20, s = 2]; spores yellow golden to green in wet mounts; densely echinate under the light microscope, spines up to ± 0.8 µm tall; under SEM spores ornamented with echinate spines that fuse at the tips forming a tipi-like shape; no oil drop; spores thick-walled; pedicels short, ± 0.8  µm long, with a broken end: free-floating sterigmata not present in wet mounts; spores of equal size under light microscopy. Eucapillitium Calvatia-type, threads up to 6.4-7 µm broad with walls ± 0.8 µm thick; crystalline deposits encrusting the eucapillitium, yellow golden in wet mounts; fragile, cracking and disarticulating at the cracks easily; glabrous to incrusted with crystalline cellular debris, straight; capillitium threads sinuous at the tips along thin threads; tips rounded. Pores present, round and minute; slit-like to stellate-shaped cracks in the eucapillitium, but not abundant, under SEM threads torn where the pores form on the hyphal walls, cracks resemble a fracture in the hyphal wall and not derived via morphological development. Septa absent. Paracapillitium abundant, incrusted with cellular debris. Exoperidium textura globulosa; composed of large, thin-walled, inflated, oval-shaped sphaerocyst cells. Endoperidium textura intricata; composed of interwoven chains of cells, becoming more uniform or linear and longitudinal towards the apex of the fruitbody.

Habitat: Sometimes the fruitbodies of Calvatia fumosa will remain almost completely hypogeous throughout maturity in thick conifer duff, or are partially buried, subhypogeous. Found on paths, on disturbed soils, growing in burn areas, in the open with sun exposure, and on dead vegetation. Growing solitarily or in small gregarious groups, but not fused. Calvatia fumosa has an unmistakable herbaceous swamp gas aroma that is compared to the smell of rotten socks or a men’s locker room. Collected under Pseudotsuga menziesii (Douglas fir), under conifers, pines, and in thick duff. Fruiting around 1,219-2,430 meters (4,000-8,000 feet). Occurring along the higher elevations of Western North America, in conifer forests of the Cascade and Sierra Nevada Mountain ranges. Among the Geographic Subdivisions of California, this species is found in Northwestern California, the Cascade Range, in the Sierra Nevada, and in the Eastern Sierra Nevada.

Distribution: A Montane species known from parts of the United States, and previously reported from Arizona, California, Colorado, Idaho, Montana, Oregon, Washington, Wyoming.

            Material Examined: CALIFORNIA, Alpine Co.: Alpine Lake, 304 meters (1,000 feet) elev., gregarious in soil under conifers, July 1975, H.D. Thiers (HDT 34554); Alpine Lake, vicinity of lake, scattered in humus under mixed conifer woods, June 1976, coll. unknown (Halling 1412); Alpine Lake, 2,286 meters (7,500 feet) elev., solitary in soil under conifers, August 1983, coll. unknown (ET 46045). Amador Co.: Silver Lake, Plasser Resort, terrestrial at the edge of a small meadow under Abies magnifica (red fir), 26 July 1937, V. Muntzer (UCB568849)(UC); East of Silver Lake, in sandy soil, September 1979, coll. unknown (SFSU S.N.); Silver Lake Campground, Highway 88, solitary in soil under conifers, July 1982, H.D. Thiers (HDT 44641); Silver Lake, Highway 88, solitary in soil under pines, July 1983, H.D. Thiers (UCB45961)(UC); Silver Lake, Highway 88, solitary in humus in conifer duff, May 1988, H.D. Thiers (HDT 51653). Calaveras Co.: Camp Connell, State Highway 4, gregarious in soil under pine and fir woods, May 1992, H.D. Thiers (HDT 54182). Fresno Co.: Huntington Lake, scattered in soil under conifers, August 1983, H.D. Thiers (HDT 46084). Lassen Co.: Lassen National Park, gregarious under Tsuga mertensiana (mountain hemlock), July 1976, H.D. Thiers (HDT 36225). Mariposa Co.: Yosemite Park, Crane Flat, gregarious in humus under firs and pines, June 1976, coll. unknown (Halling 1397). Mendocino Co.: Mendocino Park, in soil under Abies magnifica (red fir), June 1980, H.D. Thiers (HDT 40852). Plumas Co.: Graeagle Highway, Gold Lake, gregarious in soil under conifers, June 1989, H.D. Thiers (HDT 52190). San Mateo Co.: San Francisco Watershed, Marnhall Pan Road, in soil under pine and spruce, 28 July 1970, H.D. Thiers (HDT 26495). Shasta Co.: Lassen National Park, Emigrant Pass, 1,828 meters (6,000 feet) elev., terrestrial in Abies amabilis (silver fir) woods, 31 July 1955, coll. unknown (UCB138919)(UC); Lassen National Park, between Hat Lake and Paradise Meadow, terrestrial along trail, under conifers, 2,133 meters (7,000 feet) elev., 12 July 1966, (UCB1320785)(UC); Lassen National Park, Terrace-Paradise-Hat lakes trail, on litter and duff, 30 August 1974, coll. unknown (UCB1457089)(UC); Dead Horse Summit, Highway 89, in litter under conifers, July 1982, H.D. Thiers (HDT 44594). Sierra Co.: Yuba Pass, Chapman Creek Campground, in area with Abies concolor (white fir) and Pinus lambertiana (sugar pine), 11 June 1961, D. Noack (UCB1466436)(UC); Yuba Pass, Highway 49, solitary under Pinus ponderosa (Ponderosa pine), June 1967, H.D. Thiers (HDT 19510); Yuba Pass, Highway 49, solitary on soil under conifers, May 1986, H.D. Thiers (HDT 49806); Weber Lake, 4 miles south of the Yuba Pass, solitary in soil under conifers, June 1986, H.D. Thiers (HDT 50021); Tahoe National Forest, Haskell Peak Road, off Gold Lake Road, solitary in soil under conifers, June 1989, H.D. Thiers (HDT 52163); Gold Lakes Road, Howard Creek Road, Haskell Peak Road, scattered in mixed conifer woods, 8 June 1989, M.T. Seidl (UCB1574377)(UC); Sierra Nevada Field Campus, Highway 49, Northwest side of camp at the last tent, terrestrial, 1 June 2008, D. Smith (SS J205); Chapman Creek Campground, Highway 49, terrestrial along creek trail, 2 June 2008, S.S. Jarvis (SSJ 212); Chapman Creek Campground, Highway 49, terrestrial in grassy field, 2 June 2008, S.S. Jarvis (SSJ 214); Sierra City, Wild Plum Road, Girl Scout Camp, 3 June 2008, S.S. Jarvis (SSJ 215); Yuba Pass Summit, North side of the road, 3 June 2008, S.S. Jarvis (SSJ 217). Siskiyou Co.: Mt. Shasta, Sisson Southern trail, terrestrial, 21 June 1947, coll. unknown (UCB138970)(UC); Mt. Shasta, Sisson Southern trail near Horse Camp, 2,438 meters (8,000 feet) elev., terrestrial, 30 June 1954, coll. unknown (UCB139070)(UC); Mt. Shasta, Jeep Trail to Clear Creek, 2,286 meters (7,500 feet) elev., terrestrial in fir woods, 3 August 1955, coll. unknown (UCB139067)(UC); Mt. Shasta, between Bunny Flat and Horse Camp, 2,133 meters (7,000 feet) elev., terrestrial, 14 July 1961, coll. unknown (UCB1318122)(UC); Mt. Shasta, Panther Creek, 2,133 meters (7,000 feet) elev., terrestrial in fir woods, 18 July 1961, coll. unknown (UCB1318145)(UC); McCloud, Highway 89, solitary in humus under conifers, June 1967, H.D. Thiers (HDT 19597); Mt. Shasta, Horse Camp, terrestrial, 2,346 meters (7,700 feet) elev., 10 July 1963, coll. unknown (UCB1318098)(UC); Mt. Shasta, Horse Camp, on litter and duff, 20 August 1974, coll. unknown (UCB1457174)(UC); Mt. Shasta, Horse Camp, on litter and duff, 24 July 1980, W.B. Cooke (UCB1474847)(UC); Mt. Shasta, solitary on soil under conifers, July 1982, H.D. Thiers (HDT 44586); Road # 49, 1,219-1,524 meters (4,000-5,000 feet), terrestrial, 31 May 2008, D. Smith (SSJ 207). Tehama Co.: East of Mineral, Highway 36, solitary in soil under conifers, June 1988, H.D. Thiers (HDT 51709). Tulare Co.: Sequoia National Forest, General Grant Forest, South fork of King’s River, 2,133 meters (7,000 feet) elev., terrestrial along trail, July 1911, coll. unknown (UCB793282)(UC). Tuolumne Co.: Pinecrest Campground, scattered in humus under conifers, June 1965, H.D. Thiers (HDT 12639); Stanislaus National Forest, Pinecrest, gregarious in soil under conifers, May 1969, H.D. Thiers (HDT 23368); Yosemite, Middle Fork Campground, Evergreen Road, solitary in soil under conifer woods, May 1980, H.D. Thiers (UCB40770)(UC).

            Comments:  Calvatia fumosa is distinct with a noxious odor that develops as the spores mature, this aroma will persist even in older dry material. Without much explanation, Ponce de Leon (1976) placed Calvatia fumosa into Gastropila fumosa. Gastropila is a genus based on smooth spores, a three-layered peridium, a very thick endoperidium that is not easily separable from the exoperidium, and the lack of a subgleba. Being that Calvatia fumosa has echinulate spores, and peridium tissues that will separate from each other when manipulated, it is unclear why this should remain in Gastropila. Furthermore, Calvatia fumosa does not have a three-layered peridium. There are other workers who study the Lycoperdaceae, and recognize Gastropila for unexplained reasons. Due to the macro- and micro-characters of Calvatia fumosa following a Calvatia-like morphology, it will be treated as a Calvatia species here. However, based on ITS sequence, this falls out in Lycoperdon. Before a formal transfer is made, additional genes should be sequenced to confirm this finding.

 

Calvatia lloydii Zeller & Coker, Mycologia 39(3): 301. 1947.

                                                                                                                        (fig. 13, 53)

Reported synonyms:

            = Handkea lloydii (Zeller & Coker) Kreisel, Nova Hedwigia 48(3-4): 286. 1989.

Type: The holotype is housed at the University of California, Berkeley, Jepson Herbarium (UCB698059)(UC), collected along the Halstead Creek Trail near the Halstead Meadow, Sequoia National Park, Tulare Co, California, collected by L. Bonar, 28 July, 1945.

            Gasterocarp 10-40 mm tall x 12-55 mm broad; depressed globose, round to flat at the apex; rhizomorph of a tuft of mycelium or a pad of mycelium incrusted with soil; ostiole composed of lateral to stellate cracks forming at the apex. Exoperidium white to cream when young (4A2-3), clay to dark yellowish brown (5C5-5D5), then dark red brown with age (6F7-8F7), becoming darker with sun exposure; remaining adherent to the endoperidium, having a furfuraceous surface when young, cracking into octagonal scale-like warts or patches with flattened tops as the fruitbody expands with growth, some warts having obtuse spines. Endoperidium white to cream when young (4A2-3), clay to dark yellowish brown (5C5-5D5); glabrous, becoming thin and delicate with age, sloughing away from the apex with the exoperidium to reveal the gleba. Gleba, white to cream at first (4A2-3), becoming clay colored (5C5) to olive brown (4E7), then dark brown (6F7); remaining cottony throughout maturation. Subgleba extending up into the gleba from the base, concave, making a cup around the gleba at the outer edges of the fruitbody, composed of compact thin layers of cells, having a grey purple metallic tint (13E2). Diaphragm absent.

            Basidiospores globose; (2.4-) 4-4 X (-2.4) 4-4 µm [xmr = 3.9-4.0 X 3.9-4.0 µm, xmm = 3.9 ± 0.1 X 3.9 ± 0.1 µm, Q = 1.0, Qmr = 1.0, Qmm = 1.0 ± 0.0, n = 20, s = 2]; spores golden brown in KOH mounts; punctate verrucose ornamentation under light microscope with a hyaline envelope in KOH mounts, having long verrucose spines with appressed tips under SEM; thick-walled; oil drop absent; spores thick-walled; pedicel short with a torn broken end, < 0.8 µm long; free-floating sterigmata not present in wet mounts; spores of even size under light microscope. Eucapillitium Calvatia-type; threads 2.4-7.2 µm broad, with walls up to 0.8-1 µm thick; brown in KOH mounts; fragile and easily fragmenting, breaking unevenly; long threads with dichotomous branching, larger threads straight, glabrous, thinner hyphal threads slightly sinuous, knob-like projections present; threads attenuate to blunt round ends, some ends bulbous and somewhat swollen. Pores abundant, round punctate; slit-like and sinuous cracks present under light microscope, cracks identified using SEM; hyphal threads disarticulating at the cracks seen with SEM, cracks with random orientation, not just perpendicular along the hyphal walls. Septa scarce, difficult to see in light microscope mounts and SEM images, capillitium threads have uneven edges where broken. Paracapillitium absent. Exoperidium textura globulosa; composed of tightly packed deflated cells with little descriptive formation, sphaerocyst cells deflated from desiccation. Endoperidium textura intricata; composed of entangled hyphae, with thick, branched walls and round bulbous ends.

Habitat: Associated with Abies spp. (fir) and Pinus spp. (pine) collected in dry duff in spring after the snow melts, and fruiting with summer rainstorms. Among the Geographic Subdivisions of California, this puffball can be found in the Sierra Nevada, and in the San Bernardino Mountains of Southwestern California.

Distribution: A Montane species known only from the Western United States, and previously reported from California and Idaho.

            Material Examined: CALIFORNIA, Nevada Co.: Tahoe National Forest, Nevada City, White Cloud Campground, solitary in humus under conifer, 14 May 1960, H.D. Thiers (HDT 7611). Plumas Co.: Lassen National Park, Juniper Lake, scattered in humus under conifers, 30 June 1964, H.D. Thiers (HDT 12885). San Bernardino Co.: Camp Osceola, between fir and oak stands of Ponderosa pine laws, 27 September 1976, T. Tang (UCB1462470)(UC); San Bernardino, Camp Oongo, near Running Springs, in litter under Pinus ponderosa (Ponderosa pine), 10 October 1976, T. Tang (UCB1462467)(UC). Tuolumne Co.: Yosemite, Crane Flat, 1,889 meters (6,200 feet), solitary on soil under conifers, June 1983, H. Saylor (HS 1557); Stanislaus National Forest, in deep shade along dry stream, 29 July 1931, coll. unknown (UCB506570)(UC); Stanislaus National Forest, in open forest attached to extremely rotted wood, 29 July 1931, det. S.M. Zeller (UCB564470)(UC); Sonora Pass, 10 miles up Herring Creek Road, 2,286-2,438 meters (7,500-8,000 feet), 31 July 2013, C. Shaw (SSJ 484). Tulare Co.: Sequoia National Park, Giant Grove, 11 August 1905, K. Brandege (UCB564436)(UC); Sequoia National Park, in humus in forest, July 1940, S.M. Zeller (UCB698060)(UC); Sequoia National Park, Crescent Meadow, gregarious in soil under Abies concolor (fir), 19 May 1941, S.M. Zeller (UCB698058)(UC); Sequoia National Park, Giant Forest Park headquarters area, on dry duff, under pine and fir woods, 27 July 1945, S.M. Zeller (UCB698064)(UC); Sequoia National Forest, along Halstead Creek trail, near Halstead Meadow, 1,828 meters (6,000 feet) elev., usually single, scattered in dry duff of pine and fir, 29 July 1945, TYPE det. S.M. Zeller (UCB698059)(UC)(TYPE); Sequoia National Park, Suwannee Grove, on dry duff under fir and pine woods, 29 July 1945, L. Bonar (UCB698061)(UC); Sequoia National Park, Giant Grove, Moro Rock Trail, in dry duff under fir woods, 1 August 1945, L. Bonar (UCB698062)(UC).

Comments:  Calvatia lloydii shares many features with other Calvatia species of puffballs. The cracking scales on the exoperidium are similar to what is found in Calvatia cretacea, Lycoperdon subcretaceum, Calvatia booniana, and sometimes Calbovista subsculpta. However, C. booniana has scales that are much larger in size and appearance than any of these listed above. The fragile Calvatia-type eucapillitium is common in this genus, and is shared between Calvatia lloydii, Calvatia pachyderma, and Calvatia fumosa. Many in the Calvatia genus have a subgleba that roots the fruitbody to the soil. Calvatia lloydii has this feature in common with Calvatia fragilis, Calvatia cyathiformis, and Calvatia craniiformis.

Calvatia lloydii is currently placed in Handkea based on the slit-like nature of the pits on the eucapillitium threads, and based on Kreisel’s (1992) Handkea description, which describes this pit morphology as taxonomically important. Using SEM analysis, the slit-like nature of pits does not seem morphologically derived. Slit-like and sinuous pits are indisputable cracks in SEM imaging and not morphologically derived pits as described by Kreisel (1994). The nature of these pits is due to fragile capillitium threads, desiccation, and cracking that cause the slit-like appearance. Because of this SEM analysis, slit-like pits are not recognized, and therefore Handkea is not recognized as a genus. However, based on ITS sequence, this falls out in Lycoperdon. Before a formal transfer is made, additional genes should be sequenced to confirm this finding.

 

Calvatia pachyderma (Peck) Morgan, J. Cincinnati Soc. Nat. Hist. 12: 167. 1890.                                                                                                                                                 (fig. 14, 54)

Basionym: º 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.

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.

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.

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).

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 sculpta (Harkn.) Lloyd, Mycol. Writ.: 203. 1904.                                                                                                                                                                        (fig. 15, 42, 55)

Basionym: º Lycoperdon sculptum Harkness, Bull. Calif. Acad. Sci. 1(3): 160. 1885.

            Type: It has been recorded that the original Harkness fungi collection was destroyed in the 1906 earthquake fire at the California Academy of Sciences, (Setchell, 1908). Lloyd states in his Mycological notes, (No. 18, pp. 203, 1908), that an herbarium collection exists at Kew, of ripe gleba sent by Harkness. Two isotype collections are at the U.S. National Fungus Collection in Beltsville, MD, (BPI 711153, 734315). The type location is Sierra Nevada, Placer County, California, U.S.A, between 1,828-2,590 meters (6,000-8,500 feet) elevation.

            Gasterocarp 5-30 (65) mm tall x 2-35 (65) mm broad when young, older specimens get as large as 100-150 mm tall 70-150 mm broad; globose to obpyriform or oval in shape, or turbinate tapered at the base forming a v-shape in most collections, sometimes pyriforme, plicate at the base, some fruitbodies with a slight swollen bulbous base; rhizomorph composed of thick and thin branching hyphal threads, incrusted with soil, root-like, up to 20-30 mm long with branching hair-like fibers radiating out the sides of the main stem; ostiole opening via a horizontal or stellate crack at the apex through the exoperidium, scales falling away and tearing open the endoperidium layer to reveal the gleba. Exoperidium cream white when young (4A2-3), to yellowish brown (5C5-5D8); fruitbodies cottony and spongy when young, up to 4-10 mm thick, developing into distinct pyramidal warts as the fruitbody swells and enlarges via the cracking and separating of the exoperidium tissue, developing a dense covering of warts that may have recurved or hooked tips, warts may remain fused at the tips of other warts, or they may separate as the fruitbody grown into individual pyramidal warts; pyramidal warts can be large, ranging from 5-30 mm tall x 2-35 mm broad at the base, depending on the size of the mature fruitbody; tips staining brown when touched; often with areolate rings around the warts, becoming ridged with grooved longitudinal lines; leather-like when mature, warts readily falling away with drying and maturation, revealing the endoperidium, leaving an areolate pattern or scar on the endoperidium. Endoperidium cream white when young (4A2-3), to buff beige or yellowish brown (5C5-5D8), tawny tan brown (5D5), slightly metallic and shimmery; smooth and parchment-like, persistent, opening via lateral to stellate cracks at the apex, sloughing off slowly, sometimes with exoperidium warts, falling apart until only the sterile base remains. Gleba cream white when young (4A2-3), light tan (5C5) turning olive green brown (4E7); firm when young, becoming pulverulent to completely powdery with age. Subgleba white at first, becoming tan brown gray (5D5); taking up the lower 1/4 of the entire gasterocarp, 5-35 mm tall, tapered in a v-shape to the base, chambered, cottony with cellular type tissue. Diaphragm absent.

            Basidiospores globose to subglobose; (3.2-) 4-7 X 4-7 µm [xmr = 4.4-6.6 X 4.3-6.6 µm, xmm = 5.5 ± 1.6 X 5.4 ± 1.6 µm, Q = 0.8-1.4, Qmr = 1.0-1.0 Qmm = 1.0 ± 0.0, n = 25, s = 2]; hyaline when young, mature spores golden yellow in water mounts, amber in KOH mounts; roughened to minutely echinulate with dense pin pricks under light microscopy, with SEM the ornamentation has minute appressed echinulate warts, some fused at the tips with multiple warts to form a larger protuberance, matching that of the exoperidium ornamentation; oil drop present; spores thick-walled; pedicel absent, or pedicels very rudimentary, broken with rough edges seen in SEM; free-floating sterigmata not present in wet mounts; spores of equal size in light microscopy. Eucapillitium Calvatia-type, 2-8 µm broad with walls 0.8-2 µm thick; hyaline where thin, thick walls golden brown, thinner threads have less pigmentation; semi-elastic when young becoming fragile with age; straight where thick, to sinuous, to undulate or subundulate where thinner, incrusted with cellular debris, dichotomous branching scarce, eucapillitial tips attenuate to a long slender fine pointed terminus, or a sub-acute round terminus. Pores present, punctate round, small to medium-sized, pores aggregated into clusters, sometimes hard to discern from incrusted eucapillitium. Septa frequent, disarticulating at the septum. Paracapillitium present, with hyaline thin walls and septa, turning light blue after heating reaction with LpCB reagent. Exoperidium textura angularis; composed of hyaline thin-walled random-shaped sphaerocyst cells, with angular pointed corners in fresh material, bent and folded in dried material. Endoperidium a combination of textura intricata and textura globulosa; composed of swollen club-shaped cells, aligned and attached to one another in links, gold brown pigmented, thin walls, intertwined with long hyphal threads.

Habitat: Growing among Calocedrus decurrens (incense cedar), Pinus ponderosa ponderosa pine), Pseudotsuga menziesii (Douglas fir), under Abies spp. (fir) trees, and other conifers and hardwoods. Growing on disturbed soils, poor compact soils, in full sun, in thick duff, and sometimes growing on well-decayed conifer wood. Fruiting in the higher elevations, in spring after the snow melts. Among the Geographic Subdivisions of California, this species is found in the northeastern regions of Northwestern California, in the Cascade Range, in the Modoc Plateau, in the high elevations of the Great Central Valley, in the Sierra Nevada, in the Eastern Sierra Nevada, in Central Western California when snowmelt is present, and in the high elevations of Southwestern California.

Distribution: Known only from the Western United States, and previously reported from California, Idaho, Oregon, and Washington. 

            Material Examined: CALIFORNIA, Alpine Co.: Ebbitts Pass, fifteen miles from Markleville, 1 August 1939, E.E. Morse (UCB621547)(UC); Alpine Lake, Highway 4, in humus under Pinus contorta (lodgepole pine), October 1967, Jordan (J 856); Alpine Lake, Highway 4, solitary in conifer woods, August 1974, H.D. Thiers (HDT 32852)(UC); Bear Valley, 2,286 meters (7,500 feet) elev., SE facing slope, on moist soil with six inches of humus, beneath Abies magnifica (red fir), July 1977, Bowman (SFSU no #). Amador Co.: Silver Lake, terrestrial, 24 July 1933, L. Bonar (UCB506691)(UC). Silver Lake, scattered in humus under conifer, June 1973, H.D. Thiers (HDT 31065); Highway 88, five miles west of Silver Lake, solitary in rotten log debris under Abies magnifica (red fir), June 1976, Halling (Halling 1409); Highway 88, solitary under young fir trees, July 1983, H.D. Thiers (HDT 45963). Butte Co.: Jonesville, terrestrial, July 1941, E.B. Copeland (UCB652847)(UC); Jonesville, terrestrial, July 1941, V. Miller (UCB652847)(UC); State Highway 88, solitary in duff under aspen (Populus sp.), June 1966, coll. unknown (WJS 530); Forest Ranch, Doyle, Highway 395, North of Davis Lake, 14 June 2009, B. Wattenberg (SSJ 321). El Dorado Co.: Lake Tahoe, near Emerald Bay, summer 1942, coll. unknown (UCB506689)(UC); El Dorado, Echo Summit, terrestrial, 18 June 2009, R. Pastorino (SS J327). Fresno Co.: Huntington Lake, dry places, in forest proper growing on Abies concolor (white fir) and Pinus ponderosa (ponderosa pine), 26 July 1941, V Miller (UCB655338)(UC); Huntington Lake, solitary under conifers, October 1965, HD Thiers (HDT 13360); Huntington Lake, solitary under conifers, July 1975, HD Thiers (HDT 343323); Huntington Lake, solitary in humus under conifers, 28 September 1975, HD Thiers (HDT 34946); Huntington Lake, solitary in soil in conifers, September 1975, HD Thiers (HDT 35011); Dinkey Creek, on soil under conifers, 23 June 1998, M Bowell (s.n.). Kern Co.: Salmon Creek, Kern River tributary, Brown’s meadow, headwaters, June 1903, coll. unknown (UCB506681)(UC). Lake Co.: Snow Mountain, near Highway 162, on soil under Abies magnifica (red fir), July 1975, Toren (T 1966). Lassen Co.: Susanville, December 1930, C.M. Wilder (UCB439752)(UC); Blue Lake Campground, solitary in soil under conifer, 6 June 1979, H.D. Thiers (HDT 39878). Madera Co.: Sierra National Forest, Camp Soquel, growing on road in loose dirt, 1 September 1933, D.L. Burdick (UCB506683)(UC). Mariposa Co.: Chowchilla Mountain, West of Wawona, Sierra National Forest, growing on a stump of a log, 2 August 1934, W.E. Osborn (UCB527103)(UC). Mono Co.: Mono, Devil’s Post Pile, Soda Creek Campground, September 1967, H.D. Thiers (HDT 20916). Modoc Co.: Blue Lake, County Route 510, in campground, in duff under Pinus ponderosa (ponderosa pine), June 1981, coll. unknown (SFSU No. 427). Placer Co.: Auburn, July 1912, J.C. Hawver (UCB506690)(UC). Plumas Co.: Sung Lake, July 1912, CM Wilder (UCB439670); Quincy, summer 1912, coll. unknown (UCB506688)(UC); Bucks Lake, south shore of the lake, 11 July 1942, W.A. Satchell (UCB671386)(UCB). San Bernardino Co.: San Bernardino Mountains, New Buff Lake, beneath Salix spp. (willow), Abies sp (fir) and Pinus jeffreyi (Jeffery pines), 26 July 1906, H.M. Hall (UCB126182)(UC); San Bernardino Mountains, New Buff Lake, beneath Salix spp. (willow), Abies spp. (fir) and Pinus jeffreyi (Jeffery pines), 26 July 1906, H.M. Hall (UCB126183)(UC); San Bernardino Mountains, Bluff Lake, on a damp hillside under a pine tree, July 1920, G. Nicholson (UCB474189)(UC); San Bernardino Mountains, growing from a rotten stump, March 1929, E.E. Morse (UCB506686)(UC). San Mateo Co.: San Francisco Watershed, upper, March 1969, H.D. Thiers (HDT 23189). Shasta Co.: Dead Hare Summit, solitary in soil under conifers, 2 July 1982, H.D. Thiers (HDT 44600). Sierra Co.: Sierra Valley, 16 June 1899, Mr. Fowler (UCB506685)(UC); Truckee, 7 July 1913, W.C. Hofan (UCB506684)(UC); Truckee, Saghen Station, Highway 89, solitary under Pinus contorta (lodgepole pines), June 1964, Gaetien (s.n.); Yuba Pass summit, solitary in duff of mixed forest, August 1965, coll. unknown (WJS 344); Yuba Pass, East side near the summit, 26 May 1969, S.P. Wells (UCB1408022)(UC); Sierra Nevada, Lake Tahoe area, 15 August 1970, L. Bonar (UCB1455227)(UC); Yuba Pass, Highway 49, solitary on a very rotten Abies magnifica (red fir) log, September 1980, H.D. Thiers (HDT 41529); Yuba Pass, Highway 49, scattered in soil under conifer, June 1986, H.D. Thiers (HDT 49957); Yuba Pass, Highway 49, 2,042 meters (6,700 feet) elev., gregarious in soil under Abies spp. (fir), July 1988, H.D. Thiers (HDT 51851); Yuba Pass, Forest Highway 12, solitary in moist soil near seepage under conifer, September 1989, H.D. Thiers (HDT 52694); Yuba Pass, five miles down Webber Lake Road, terrestrial, 10 June 1992, M.T. Seidl (UCB159874)(UC). Siskiyou Co.: Ponderosa, two miles east, on soil under Pinus ponderosa (ponderosa pine), 25 May 1941, E.E. Morse (UCB652831)(UC); Mount Shasta, Mud Creek Dam Road, terrestrial along squaw creek, 9 April 1947, W.M. Cooke (UCBM139014)(UC); Mount Shasta, Bear Springs, terrestrial, 19 July 1947, W.M. Cooke (UCBM138905)(UC); Mount Shasta, Sisson southern trail route, terrestrial in logged conifer woods, 5 July 1968, W.M. Cooke (UCB1457170)(UC); Mount Shasta, solitary under conifers, October 1983, H.D. Thiers (HDT 46426); Shasta National Forest, Route #49, 1,524 meters (5,000 feet) elev., terrestrial, 31 May 2008, S.S. Jarvis (SSJ 208). Trinity Co.: Salmon Mountains, 29 July1909, H. MacIlwaine (UCB506682)(UC); South of Big Flat, near Coffee Creek Road, 3 July 1937, E.E. Morse (UCB585296)(UC). Tulare Co.: Sequoia National Park, 16 June 1931, H.E. Parks (UCB472118)(UC); Sequoia National Park, General Grant National Park, in mixed forest, summer 1931, E.E. Morse (UCB506687)(UC); Miramonte, Sequoia National Park, N36.41.280 – W119.02.798, 24 March 2009, S.S. Jarvis (SSJ 310). Tuolumne Co.: Stanislaus National Forest, Cow Creek, June 1937, coll. unknown (UCB568974)(UC); Pinecrest, solitary under conifer, June 1973, H.D. Thiers (HDT 30922); Pinecrest, 1,706 meters (5,600 feet) elev., scattered in soil under conifers, May 1980, H.D. Thiers (HDT 40813).

Comments: Calvatia sculpta is the most extravagantly ornamented Calvatia species in North America. This used to be collected and mixed with Calbovista subsculpta until the later became understood and described as it’s own species in 1935 (Morse 1935). Lloyd’s (1904) illustration of the type material lacks the elaborate spines, however the spines get quite large with the right environmental conditions. This puffball is a choice edible in the Sierra Nevada Mountains where it is found, when it is young, white, and firm inside. The fruitbody will continue to ripen after collected and will often fall apart quickly and rot if not refrigerated right away, or properly dried on a fruit dehydrator for preservation. However, based on ITS sequence, this falls out in Lycoperdon. Before a formal transfer is made, additional genes should be sequenced to confirm this finding.

 

 

 

DISCISEDA Czern.:

Disciseda atra (Lloyd) Zeller, [as ‘ater’] Mycologia 39(3): 307. 1947.                                                                                                                                                            (fig. 16, 39, 56)

Basionym: º Catastoma atrum Lloyd, Mycol. Writ. 5(Letter 53): 756. 1918.

Type: The holotype material is located at the U.S National Fungus Collections (BPI), catalog #706673 (CG Lloyd Myc. Collection #33650). There is an isotype fragment at the New York Botanical Garden, catalog # 00291979. The holotype material is from the San Gabriel Mountains, Mount San Antonio, Orange Co., CA, collected by IM Johnston, 17 September 1917.

Gasterocarp 28 mm tall x 30 mm broad; globose; rhizomorph absent; ostiole slow to develop through desiccation and thinning of the peridium layers, fine cracks appearing with minute fibrils along the margin, which separate to form a round stoma. Exoperidium whitish at first becoming dark brown to dark grayish brown (6F7-3) in age, with a dull sheen; thin, black, smooth, adherent to the endoperidium, persistent and adhering to the endoperidium. Endoperidium tawny olive to yellow brown (5D5-5E5); hard, up to 2 mm thick, smooth, and persistent. Gleba white and firm when young, becoming dark brown (6F5-7), without purple tinge; firm when young, becoming powdery with maturation. Subgleba absent. Diaphragm absent. Sand case absent or missing on the type specimen. Macromorphological description derived from the dried type specimen.

            Basidiospores subglobose or broadly ellipsoidal; 5-6 X 4.5-5.5 µm [xmr = 5.4 X 4.7 µm, xmm = 5.4 ± 0.0 X 4.7 ± 0.0 µm, Q = 1.0-1.2, n = 35, s = 1]; spores golden in wet mounts; minutely asperulate roughening under light microscope, under SEM the spores have well spaced, broad asperulate bumps and a slight granular-like encrusting material; oil drop absent; spores thick-walled; a very short mucro, up to ± 0.8 µm long; free-floating sterigmata remnants absent from wet mounts; spores of equal size under light microscope. Capillitium Disciseda-type; threads up to 5-7 µm broad with walls up to  ± 1.5 µm thick; golden yellow in wet mounts; fragile, breaking into short segments; glabrous, short dichotomous branches present, mostly straight to somewhat sinuous; tips with a rounded terminus. Pores absent or scarce. Septa present and abundant, threads disarticulating at the septum. Paracapillitium absent. Exoperidium textura globosa; composed of compact inflated sphaerocyst cells. Endoperidium textura intricata; composed of septate, tightly woven hyphal threads.

Habitat: Cow Canyon, San Gabriel Watershed, on dry gravel wash, growing solitarily. Among the Geographic Regions of California, this species is only known from Southwestern California.

Distribution: Known only from the San Gabriel Mountains, Mount San Antonio, California.

Material Examined: CALIFORNIA, Orange Co.: Mount San Antonio, Cow Canyon, San Gabriel Watershed, in dry gravel wash, 17 September 1917, I.M. Johnston (CG Lloyd Mycological Collection #33650; BPI706673; TYPE).

            Comments:  Johnston (type collection notes, 1917) writes, “this impressed me at once by its exceptionally black endoperidium. It has the habits, and I thought at first was a Bovista, but the capillitium is of a Catastoma and in these intermediate species the capillitium characters decide.”  He goes on to say that this does not have the ‘mouth down’ character normal to Disciseda, so this is not ideal for the Disciseda genera. However, there are many Disciseda species that do not exhibit the rolling over behavior. Zeller (1947) describes Disciseda atra with the appearance of a black weathered oak gall, without a sand case, an exoperidium that closely adheres to the endoperidium, and being so similar to D. candida as to confuse them for one another. Disciseda atra was put into synonymy with D. candida by Cunningham (1979) for this reason. Disciseda atra, needs additional collections of material at all ranges of maturity to be further understood.             Evidence based on morphology, SEM, and ITS data, suggests that D. atra is conspecific with D. levispora, and both are nested within the Calvatia clade on the ML tree with 100% bootstrap and 100% PP. In 1972 A.H. Smith annotated the type specimen of Disciseda levispora, (seen on type specimen notes) suggesting that it was Calvatia hesperia (Morgan) 1895, which was described from Pasadena, CA, and the data here supports this contention. Both D. atra and D. levispora were originally described from Southern California, Orange County and Riverside County, respectively. When comparing the descriptions of Calvatia hesperia (both Morgan 1895, and Ponce de Leon 1976) to descriptions of Disciseda atra and Disciseda levispora, there is a great deal of similarity. None of these descriptions mention a sand case, they all have a hard or coriaceous endoperidium, a thin exoperidium that adheres to the endoperidium, the same size and ornamentation of the spores, similar size and characters in the eucapillitium, and all seem to prefer sandy soil or growing on the terra firma. One main difference is the size of the gasterocarps, however Morgan notes that this species is “quite variable in size.”  Another difference is the color of the gleba, which could be explained by differences in maturation of gasterocarps used in the descriptions. Kreisel (1989) transferred Calvatia hesperia into Handkea, probably due to Ponce de Leon (1976) stating “the pits slit-like,” and here Kreisel notes “no specimen seen.” Disciseda atra was the only species among the California taxa seen with slit-like pits (using SEM), for which these pits looked morphologically derived and not like cracks in the eucapillitium walls (see fig. 16). Disciseda levispora and Disciseda atra were both transferred from Catastoma in the same paper (Zeller 1947). If these two are to remain in Disciseda, D. atra may become the preferred name for these two taxa, based on alphabetical preference. However, since their position in the ML tree is within the Calvatia clade with good support, a transfer to Calvatia may be warranted. Calvatia hesperia is the older name of these three, this is a distinct taxon, and all of this evidence combined suggests that Calvatia hesperia may be a well-suited name for both Disciseda atra and Disciseda levispora.

 

Disciseda brandegeei (Lloyd) Zeller, Mycologia 39(3): 307. 1947.                                                                                                                                                            (fig. 17, 39, 40, 57)

Basionym: º Catastoma brandegeei Lloyd, Mycol. Writ. 5(Letter 65): 7. 1917.

            Type: The holotype material is located at the U.S National Fungus Collections (BPI), catalog #706670, (CG Lloyd Mycological collection #37840). There is an isotype fragment at the New York Botanical Gardens, collection #291980. The holotype collection is from Sinaloa, Culiacan, Mexico, 1904.

Gasterocarp 14 mm tall x 25 mm broad; subglobose to depressed globose; rhizomorph absent; ostiole slow to develop into a fimbriate torn slit at the apex. Exoperidium white at first, turning orange cream buff (5A2-4A4), then dark yellowish brown (5F5); ornamentation granulose with very minute floccose spines, remaining creamy white throughout maturity, thin and flaking off. Endoperidium white at first, turning grayish yellow (4B5), yellowish brown (4D7), to dark yellowish brown (5E5-5F5); smooth and felty, thin, persistent and fibrous. Gleba white at first, turning yellowish brown (4D7), to dark yellowish brown (5E5-5F5); firm when young, becoming clumpy and cottony through maturation. Subgleba absent. Diaphragm absent. Sand case absent. Macromorphological description derived from the dried type specimen.

            Basidiospores globose; 6.5-8.5 X 6 - 8 µm [xmr = 7.4 X 7.0 µm, xmm = 7.4  ± 0.0 X 7.0 ± 0.0 µm, Q = 1.0-1.2, n = 35, s = 1]; spores golden yellow in wet mounts; smooth under light microscope and under SEM imaging; oil drop absent in wet mounts; spores thin-walled, easily damaged with SEM; pedicel absent to very rudimentary, ± 0.8 µm long; free-floating sterigmata absent from wet mounts; spores of equal size under light microscope. Eucapillitium Disciseda-type; threads up to 5-7 µm broad with walls ± 0.8 µm thick; golden brown in water mounts; fragile; glabrous, dichotomous branching scarce, straight to slightly sinuous near the terminus along thin walls, without knob-like projections; eucapillitium tips with a rounded terminus, not attenuate. Pores absent. Septa present, disarticulating at the septum. Paracapillitium absent. Exoperidium textura globulosa; composed of compact inflated sphaerocyst cells. Endoperidium textura intricata; composed of tightly woven, septate, hyphal threads.

Habitat: In open barren soil in the summer, growing solitarily. Among the Geographic Subdivisions of California, this species is reported from, but not verified, from Central Western California.

Distribution: Reported only from California, New Mexico, and Nevada. Also known from Culiacan, Mexico.

Material Examined: Sinaloa, Culiacan, Mexico, 1904, T.S. Brandege (CG Lloyd coll. 37840; BPI 706670; TYPE).

Comments:  Zeller (1947) states that Disciseda brandegeei is found from Texas to California and Mexico, although none of the material mentioned in Zeller’s 1947 description is from California. Brandege (type collection notes: 1917) states that this type specimen is young, yet is different from all other Disciseda species due to smooth spores. Describing a new species, with only immature specimens available, seems typical of taxonomy at the turn of the century. Today, with more knowledge of the habits of these fungi, smooth spores often points to immature material, and therefore most often these specimens remain unidentifiable. Disciseda brandegeei, without further collection of material at all ranges of maturity, may have been prematurely erected as a species. The ITS data here places this species just outside the Lycoperdon clade with out any support. This suggests that additional genes should be sequenced to further clarify the taxonomic position of Disciseda brandegeei amongst the other Lycoperdaceae.

 

Disciseda candida (Schwein.) Lloyd, Mycol. Writ. 1: 100. 1902.                                                                                                                                                                        (fig. 18, 41, 58)

Basionym: º Bovista candida Schwein., Schr. naturf. Ges. Leipzig 1: 59. 1822.

Reported synonyms:

            = Disciseda candida (Schwein.) Lloyd, Mycol. Writi. 1: 100. 1902. var. candida

                        Type: According to Coker and Couch (1928), the holotype of Bovista candida is in the Schweinitz Herbarium at the Academy of Natural Sciences in Philadelphia, Pennsylvania. There is an isotype of Bovista circumscissa at the Curtis Herbarium, Alton Natural History Society, Alton, Hampshire, England. JStor Plant Science reports an isotype collection at the herbarium of the University of Montpelier, collection number MPUC03224, from Morocco in 1934. Bates (2009) states that the holotype has been misplaced and isotype material of Bovista candida is at the herbarium of the University of North Carolina, citing Carolina, U.S.A. as the type locality.

Gasterocarp 14-15 mm tall x 26-27 mm broad; subglobose to depressed globose; rhizomorph composed of a tuft of mycelium threads, aiding in the exoperidium encasing soil and vegetal debris; ostiole slow to develop through desiccation and thinning of the peridium layers, fine cracks appearing with minute fibrils along the margin, which separate to form a round stoma, raised with upturned fibers, irregular round, oval to linear in shape. Exoperidium cream white (4A2-3), to light pink (5A2), to orange white (6A2), brownish gray (5C2), to orange gray (5B2) when mature; very cottony when young, completely enclosed in a net-like casing with sand and soil debris. A gelatinous mid-layer between the exoperidium and endoperidium exists; a net-like layer when fresh, becoming cracked and wrinkled with desiccation, creating a reticulate pattern between the sand case and exoperidium, sometimes the reticulate pattern only develops on the upper surface; persistent; most often the sand case needs removal to study. Endoperidium cream white (4A2-3), to light pink (5A2), orange white (6A2), brownish gray (5C2); glabrous, persistent, tough throughout maturity, sometimes developing more than one hole that will eventually aid in spore dispersal. Gleba white to cream (4A2-3) when young, turning olivaceous tones (4A4-4E8), then dark brown (6F5-6) at maturity, sometimes tinged with dark brown purple

(14F7); firm at first, becoming powdery, developing from the center outwards. Subgleba absent. Diaphragm absent. Sand case up to .5 mm thick; firm, persistent; breaking upon drying in a circumference-like fashion from desiccation and maturation, or peeling away from the endoperidium in sheets; remaining on the lower 1/3 to 1/4 of the fruitbody throughout maturation; often taking on the same colors as the soil, which can influence coloration of the gasterocarp. Sand Case rolling-over mechanism: after the fruitbody matures it will slowly rotate in place, with the upper portion of the sand case acting as the ballast. The bottom half of the sand case remains in the soil, not as a cup, but as fractured patches. Whereas the upper half of the sand case remains attached, with it’s weight acting to rotate the fruitbody over 180º. Matured fruitbodies of Disciseda with this mechanism will be found in this rotated position. This rolling over has been observed and listed in great detail in Mitchel et al., 1975.

            Basidiospores globose to broadly ellipsoidal; 4-5.5 X 4-5.5 µm [xmr = 4.7-4.8 X 4.4-4.5 µm, xmm = 4.8 ± 0.0 X 4.4 ± 0.1 µm, Q = 1.0-1.3, Qmr = 1.1-1.1, Qmm = 1.1 ± 0.0, n = 25, s = 2]; spores golden in wet mounts; heavily verruculose with appressed verrucae under light microscope, with SEM the spores have a dense covering of flattened round verruculose bumps; oil drop present; spores thick-walled; pedicel reduced to 0.8 µm or absent, a very reduced pedicel with a clean break where disarticulated seen in SEM; free-floating sterigmata absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Disciseda-type; threads up to 5-6 µm broad with walls up to 0.8 µm thick; threads brown in water mounts; fragile; glabrous, dichotomous branching present, sinuous, with knob-like projections; eucapillitium tips with a round terminus. Pores present, small, round, clustered. Septa and pseudosepta present, disarticulating at the septum. Paracapillitium present, septate, abundant. Exoperidium a mixture of textura globulosa and textura intricata; composed of tightly interwoven hyphal threads, with cellular debris, and swollen irregular-shaped sphaerocyst cells, difficult to discern in dry and aged material. Endoperidium textura intricata; composed of thick-walled and thin-walled septate hyphal threads. Basidia composed of club-shaped cells, long pedicels, four spores per basidia, thin-walled and hyaline, very difficult to isolate in wet mounts. Gelatinous layer composed of thin, hyaline, intertwined hyphal elements, with a matrix of cellular debris of collapsed non-symmetrical angular cells; hyphal threads with thick, sinuous, and with blunt ends; filled with granular-like material, resembling the eucapillitium but larger in overall width and more globose in shape.

Habitat: Found in the higher elevations after spring rains, and in lower elevations through winter and rainy seasons. Collected growing in natural grasslands, growing gregariously. Among the Geographic Subdivision of California, this species is found in the Great Central Valley, in the Sierra Nevada, in Central Western California, and in Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Alabama, Arizona, California, Colorado, Idaho, Illinois, Indiana, Iowa, Kansas, Michigan, Missouri, Montana, Nebraska, New Jersey, North Carolina, North Dakota, Ohio, Oregon, South Dakota, Texas, Utah, Virginia. Also known in Puerto Rico and Europe.

Material Examined: CALIFORNIA, Alameda Co.: Oakland, Knowland Park, terrestrial grassy field, in a partial fairy ring, 30 November 2011, S.S. Jarvis and F. Stevens (SSJ 421). Los Angeles Co.: Los Angeles, 3623 Kelton Avenue, November 1974, T. Tang (UCB1457599)(UC). Monterey Co.: Hastings Reservation, Jamesburg route, in a field, 6 January 1943, V. Miller (UCB95701)(UC); Hastings Reservation, terrestrial, 27 October 1941, V. Miller (UCB669653)(UC). Napa Co.: Napa City Park, Terrace Road, 2 February 2014. S.S. Jarvis (SSJ 485). Orange Co.: Santa Ana Mountains, 2,133 meters (7,000 feet) elev., 7 February 1929, V. Mentzer (UCB506717)(UC); Dana Point, collection from under trees, 3 August 1933, E.E. Morse (UCB506706)(UC). San Bernardino Co.: Little Bear Valley, San Bernardino Mountains, August 1907, C.M. Wilder (UCB476121)(UC); Colton, near San Bernardino, March 1929, V. Mentzer (UCB506707)(UC). San Diego Co.: San Diego, terrestrial in sandy soil, 1902, det. Tom Tang 1975 (UCB372358)(UC). San Francisco Co.: Golden Gate Park, San Francisco, on sandy soil, partially buried, 3 March 1902, W.E. Setchell (UCB372351)(UC). Sierra Co.: Sierra Nevada Field Campus, Bassetts, Highway 49, solitary in sandy soil, 9 June 1993, D.E. Desjardin (DED 5758). Stanislaus Co.: California State University, on campus in soil, 25 November 1979, L. Mitchell (s.n.)(UC).

            Comments:  The Disciseda group is confusing due to similar morphology among the taxa; the grey endoperidium, the sand case, and small stature of the fruitbodies. However, Disciseda candida can be easily distinguished macroscopically due to the reticulate pattern along the basal portion of the exoperidium. It is commonly described to have a gleba tinged with purple, and microscopically by the presence of paracapillitium and lacking a pedicel. The markings and spore ornamentation may vary among North America, Australia, Europe and New Zealand collections. In comparison to Disciseda candida, Disciseda cervina has a fimbriate ostiole, verrucose spores, sometimes has a small pedicel, occasionally has a purplish tint to the endoperidium, and lacks the gelatinous layer and the reticulate pattern found under the sand case in D. candida. Although reported in the literature, the rolling over nature of Disciseda remains to be seen and verified by this author. According to the ITS sequence data, Disciseda candida from Arizona (EU833654) appears to be distinct from Disciseda candida California (SSJ 421). Although nested in a Disciseda clade with two other taxa and fairly good support, the distance between these two contigs suggests that additional genes should be sequenced to further clarify the taxonomic position of Disciseda candida and other Disciseda species.

 

Disciseda cervina (Berk.) Hollós, Növénytani Közlemények 1: 107. 1902.

                                                                                                            (fig. 19, 41, 59)

Basionym:  º Bovista cervina Berk., Ann. Nat. Hist., Ser. 1 9: 447. 1842.

Reported synonyms:

= Catastoma magnum Lloyd, Mycol. Writ. 5(Letter 45): 631. 1917.

Type: A holotype collection of Bovista cervina exists at the Royal Botanical Gardens, Kew, collection number K-M000064223, collected by C.R. Darwin in 1842 from Rio Negro, Argentina (confirmed by Berkeley). Cunningham (1942) states that the type locality of Disciseda cervina is Europe.

Gasterocarp 8-19 mm tall x 6-25 mm broad; subglobose to depressed globose or ovoid; rhizomorph of a single thin mycelium chord or absent; ostiole up to 1 mm broad, slow to develop through desiccation and thinning of the peridium layers, fine cracks appearing with minute fibrils along the margin, which separate to form a slit-like to round or irregular-shaped stoma. Exoperidium orange white (5A2) when young, becoming mousey grey (5C2-5D2), turning brownish grey (5D2), to brown (6D3-6E4) when mature, unevenly colored or mottled, some specimens taking on the soil color; cottony when young, composed of a minutely floccose to furfuraceous hyphae, encrusting soil, vegetal, and sand debris, persistent, remaining adherent to the endoperidium, thinning with age. Endoperidium white and first with uneven coloration ranging from orange white (5A2), to grayish orange (5B4), buff brown (6E4), grayish brown (6D3), dark grayish brown tones (7E5-7F3), sometimes gray purple tones (14D1) to dark brown or brick red brownish (8E7-8F7); smooth and glabrous to the naked eye, furfuraceous and cottony under dissecting scope, mostly unwrinkled, persistent and remaining through maturity, taking on the soil color. Gleba white and firm when young, becoming light tan brown (5D5), dark brown (7F5) to brick red brown (6E6); subelastic to cottony at first, becoming pulverulent in age, but remaining intact as powdery clumps. Subgleba absent. Diaphragm absent. Sand case with the rolling-over mechanism; up to 1 mm thick when young and fresh; becoming brittle, sloughing away in large sheets or patches as the gasterocarp dries, splitting apart transversely, flaking completely off except at the very base, remaining on the lower 1/3 to 1/4 of the fruitbody; persistent, often taking on the same colors as the soil.

Basidiospores globose; (3.2) 4-6.4 (7.2) X 4-6.4 µm [xmr = 5.2-5.6 X 5.1-5.5 µm, xmm = 5.4 ± 0.2 X 5.3 ± 0.3 µm, Q = 0.8-1.3, Qmr = 1.0-1.0, Qmm = 1.01 ± 0.0, n = 20, s = 2]; spores golden brown in water mounts, remaining unchanged in lactophenol cotton blue reaction; slightly asperulate to verrucose under light microscope, with SEM mature spores appear to have a dense coating of large pointed verrucose bumps connected with ridges; central oil drop present; spores thick-walled; pedicel ± 0.8 um in length, pedicels cleanly detached seen with SEM; free-floating sterigmata remnants absent from mounts; spores of various size in wet mounts. Eucapillitium Disciseda-type; 3.2-5.6 um broad, walls ± 0.8 um thick; pale brown in water mounts; extremely fragile; glabrous to sometimes incrusted with cellular debris, dichotomous branching scarce, straight to undulate or somewhat sinuous; knob-like projections absent; tips with a rounded terminus. Small round pits abundant or occasional. Septa with subseptal ramifications, disarticulating at the septum into irregular-sized fragments, and having clean breaks at the septum wall; false septa present and scarce. Paracapillitium absent. Subgleba absent. Diaphragm absent. Exoperidium a mixture of textura intricata and textura globulosa; composed of tightly interwoven hyphal threads, with cellular debris of swollen irregular-shaped cells difficult to discern. Endoperidium textura intricata; composed of thick-walled and thin-walled septate hyphal threads.

Habitat: Terrestrial, growing in semi arid areas. Soil types ranging from coastal sandy soil, open grassy hillsides and meadows, prairies, montane regions, to sandy serpentine soils of the Sierra Foothills. Growing in small gregarious groups, but not fused. Among the Geographic Subdivisions of California, this species is found in the of Northwestern California, the Great Central Valley, the Sierra Nevada, Central Western California, and in Southwestern California.

Distribution: Known from parts of the United States, and previously reported from Arizona, California, Idaho, New Mexico, and North Dakota. Also reported from Argentina, Australia, Europe, India, Mexico, New Zealand, Pakistan, Romania, and South America.

Material Examined: CALIFORNIA, Alameda Co.: Knowland Park, open grassy field, 04 February 2009, S.S. Jarvis (SSJ 289). Contra Costa Co.: Wildcat Canyon, in bare soil sub hypogeous, 3 April 1939, V. Miller (UCB620889)(UC). Humboldt Co.: Samoa Peninsula, across from the pulp mill, scattered in moss, 30 March 1973, D. Largent (DLL 5751-2129)(HSU). Monterey Co.: Hastings Reservation, on soil surface, 2 June 1944, L. Bonar (UCB695697)(UC). San Diego Co.: Del Mar Beach, high sand cliff, 11 November 2009, R. Reidii (SSJ 296). San Francisco Co.: San Francisco Presidio, in sandy soil under Cupressus spp. (cypress), 9 November 1941, V. Miller (UCB659991)(UC). San Bernardino Co.: San Bernardino, on open ground in sandy soil, September 1940, V. Miller (UCB507198)(UC). Sonoma Co.: Pleasant Hill Road, Sebastopol, on dirt road, 24 January 2009, D. Deshazer (SSJ 338). Stanislaus Co.: Empire, Davidson Ranch, in pasture on loose soil, 30 March 1940, V. Miller (UCB637353)(UC). Trinity Co.: Lower Trinity River, 6 April 1954, L. Bonar (UCB977183)(UC). Tulare Co.: Sequoia National Park, Horse Corral Meadow, under grass in a meadow, 31 July 1945, L. Bonar (UCB695786)(UC). Tuolumne Co.: Cow Creek guard station, south of Pinecrest, 1,920 meters (6,300 feet) elev., on open ground, 9 July 1948, C.R. Quick (UCB966422)(UC); Red Hills Road and Serpentine Loop Road, BLM land, sparse grass between shrubs, 22 March 2009, S.S. Jarvis (SSJ 299). Yolo Co.: Clarksburg, Sacramento River, in sandy soil, 11 February 1939, V. Miller (UCB620843)(UC).

            Comments:  The Disciseda group can be confusing due to similarities among the taxa; grey endoperidium, sand case, and small stature. Disciseda cervina is distinctive by the fimbriate ostiole, occasional grayish purplish tint to the endoperidium, verrucose spores, lacking paracapillitium, and sometimes having a small pedicel on the spores. In comparison, Disciseda candida can be distinguished macroscopically by the reticulate pattern along the basal portion of the endoperidium, microscopically by the presence of paracapillitium, and lacking a pedicel on the spores. In addition, D, cervina has a dark brown to reddish brown powdery gleba when mature, whereas D. candida has a dark brown to purple brown powdery gleba when mature. Disciseda anomola, a common species in North America, but not reported from California, has a tubular ostiole with smooth to asperate spores that lack pedicels, separated from Disciseda candida and Disciseda cervina by having larger spores and a more prominent ostiole. Disciseda cervina seems more common in California than Disciseda candida. Although reported in the literature, the rolling over nature of Disciseda remains to be seen and verified by this author. The ITS data here supports this species within a Disciseda clade with 84% bootstrap and 98% PP.

 

Disciseda johnstonii (Lloyd) Zeller, Mycologia 39(3): 309. 1947.                                                                                                                                                                                    (fig. 20, 60)

Basionym: º Catastoma johnstonii Lloyd, Mycological Writings, 6(Letter 61): 898. 1919.           

            Type: The holotype material is located at the U.S National Fungus Collections (BPI), catalog #706635 (Lloyd Myc. Collection, No. 33647), collected by IM Johnston, San Bernardino Co., Cucamonga Canyon, 365 meters (1,200 feet) elev., 10 November 1918. An isotype fragment is located at the New York Botanical Garden, catalog #00291984.

            Gasterocarp 3-5 mm tall x 21 mm broad; depressed globose; rhizomorph absent; ostiole slow to develop through cracking and missing from the holotype collection. Exoperidium grayish brown to dark brown (6F3-4); thin, floccose to roughened, remaining adherent to the endoperidium throughout maturation, becoming cracked when old; incrusted with sand and vegetal debris creating a sand case. Endoperidium hazel to reddish tawny brown (6D5-6E7); thin, persistent, felty, remaining adherent to the exoperidium and visible only through cracks of the exoperidium. Gleba reddish tawny to hazel brown (6C5-6D5) when mature, clumped powdery to semi cottony. Subgleba absent. Diaphragm absent. Sand case present, as mentioned above, thin and becoming rigid with age, flaking off, remaining partially present on the lower half of the fruitbody. Macromorphological description derived from the dried type specimen. 

Basidiospores globose; 4-5 X 4-5 µm [xmr = 4.8 X 4.6 µm, xmm = 4.8 ± 0.0 X 4.6 ± 0.0 µm, Q = 1.0-1.1, n = 35, s = 1]; spores pale golden brown in wet mounts; ornamented with faint echinulate pin pricks under light microscope, with SEM covered in granular echinulate protrusions that fuse at the tips; oil drop absent; spores thick-walled; pedicel up to ± 0.8 um long, seen best with lactophenol blue staining in light microscopy; free-floating sterigmata absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Disciseda-type; 4.0-5.5 um broad with walls ± 0.8 um thick; eucapillitium hyaline to pale yellow in wet mounts; fragile; glabrous, with dichotomous branching, straight, lacking knob-like projections; tips with a round terminus. Pores numerous, round, clustered. Septa present, disarticulating at the septum. Paracapillitium absent. Exoperidium a combination of textura intricata and textura globulosa; composed of tightly interwoven hyphal threads, with cellular debris of swollen irregular-shaped cells, difficult to discern. Endoperidium textura intricata; composed of thin-walled, tightly woven, septate hyphal threads.

Habitat: Trailside in a chaparral belt at elevation 3000 ft in the upper Sonoran Zone. Growing in small gregarious groups, but not fused. Among the Geographic Subdivisions of California, this species is known from Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, and Colorado.

Material Examined: CALIFORNIA, San Bernardino Co.: Cucamonga Canyon, upper Sonoran zone, in open grassy ground, 365 meters (1,200 feet) elev., 10 November 1918, I.M. Johnston (Lloyd Myc. Collection, No. 33647; BPI 706635; TYPE). 

            Comments:  Lloyd (1908) describes the spores as smooth, whereas Zeller (1947) describes the spores to be more verrucose than D. candida. Lloyd (1908) notes that this species is very similar to a common species, Catastoma circumscissum, but distinct in it’s smooth spores, growing with its ‘mouth down,’ as is character to Disciseda. However, the holotype material clearly has echinulate spores. It is possible that the holotype material is immature, and further collections need to be examined to make a complete diagnosis on this species. Disciseda johnstonii, without further collection of material at all ranges of maturity, will remain incompletely understood. The ITS data here supports this species within a Disciseda clade with 84% bootstrap and 98% PP. 

 

Disciseda levispora (Lloyd) Zeller, Mycologia 39(3): 310. 1947.                                                                                                                                                                                    (fig. 21, 61)

Basionym: º Catastoma levisporum Lloyd, Mycological Writings, 6(Letter 59): 853. 1919.

Type: The holotype material is located at the U.S National Fungus Collections (BPI) catalog #706636 (Lloyd Myc. Collection #8782), collected by LM Clancy, Los Angeles Co., Claremont, California, 9 October 1916.

Gasterocarp 29 mm tall x 27 mm broad; globose; no rhizomorph; ostiole slow do develop through the thinning of the peridium layers, lateral to stellate cracking with maturity, wearing away to eventually become cup-like in overall shape. Exoperidium orange white (5A2) to a red brown (8E7); thin, floccose with hairs clinging to soil, incrusted with soil and sand debris, exoperidium color similar to adhering soil color, persistent and adhering to the endoperidium. Endoperidium brownish gray (6C3) to brownish orange (5C4); cartilaginous, tough and firm. Gleba brown to dark grayish mummy brown (7E4-7F5); powdery. Subgleba absent. Diaphragm absent. Sand case absent, yet the entire exoperidium incrusted with a fine and thin layer of sand and soil debris. Macromorphological description derived from the dried type specimen. 

Basidiospores globose, 4.5-6 X 3.5-5 µm [xmr = 5.0 X 4.5 µm, xmm = 5.0 ± 0.0 X 4.5 ± 0.0 µm, Q = 1.0-1.3, n = 35, s = 1]; pale golden yellow in wet mounts; seemingly smooth under light microscope, under SEM the spores have stellate-shaped asperulate bumps; oil drop present; spore walls <0.5 µm thick; pedicel up to 0.8-2 µm long with a clean break from the sterigma; free-floating sterigmata remnants absent from wet mounts; spores of various size in wet mounts. Eucapillitium Disciseda-type; threads up to 6-8 um broad with walls up to 0.8 µm thick; pale golden yellow in color; fragile; glabrous, dichotomous branching present, composed of straight short threads, knob-like projections present; eucapillitium tips round and not attenuate. Pores present, oval to round-shaped. Septa abundant, hyphal threads disarticulating at the septum wall. Paracapillitium absent. Exoperidium textura globulosa; composed of swollen sphaerocyst cells, tightly packed together and incrusted with soil and vegetal debris. Endoperidium textura intricata; composed of thin-walled, septate, tightly woven hyphal threads.

Habitat: Growing solitarily on soil. Among the Geographic Subdivisions of California, this species is found in the Southwestern California province.

Distribution: Known only from Riverside County and Los Angeles County, California.

Material Examined: CALIFORNIA, Los Angeles Co.: Claremont, terrestrial in a vacant lot under pepper trees, 9 October 1916, L.M. Clancy (Lloyd Myc. Collection #8782; BPI 706636; TYPE).

Comments:  D.L. Crawford (type specimen notes, 1916) writes in his holotype specimen notes, “Similar to Catastoma pedicellatum and Catastoma pila. As to spores, there is but one other species with smooth spores; Catastoma brandegeei, which is a much smaller plant and has olive gleba.”  Evidence based on morphology, SEM, and ITS data, suggests that D. levispora is a synonym of that D. atra. In 1972 A.H. Smith annotated the type specimen of Disciseda levispora, suggesting that it was Calvatia hesperia (Morgan,) 1895, which was described from Pasadena, CA. Both D. atra and D. levispora were also described from Southern California; Orange County and Riverside County, respectively. When comparing the descriptions of Calvatia hesperia (both Morgan 1895, and Ponce de Leon 1976) to descriptions of Disciseda atra and Disciseda levispora, there is a great deal of similarity. None of these descriptions mention a sand case, they all have a hard or coriaceous endoperidium, a thin exoperidium that adheres to the endoperidium, the same size and ornamentation of the spores, similar size and characters in the eucapillitium, and they all seem to prefer sandy soil or growing on the terra firma. The original description of Calvatia hesperia states a gleba with clay color, greenish to ochraceous. A brief glance of images of the type collections at NY, show the gleba color matching that of both D. atra and D. levispora. However, this is not enough evidence to support a change in nomenclature. Since their position in the ML tree is within the Calvatia clade with good support, a transfer to Calvatia may be warranted. The type collections of Calvatia hesperia, Disciseda atra, and Disciseda levispora need further comparison before any names are transferred, especially with multi loci genes in phylogenetic analysis. If Disciseda levispora and Disciseda atra are to remain in Disciseda, D. atra may become the preferred name for these two taxa, based on alphabetical preference. However, Calvatia hesperia is the older name of the three, this is a distinct taxon, and all of this evidence combined here suggests that Calvatia hesperia may be a well-suited name for both Disciseda atra and Disciseda levispora.

 

Disciseda luteola (Lloyd) Zeller, Mycologia 39(3): 310. 1947.                                                                                                                                                                        (fig. 22, 39, 62)

Basionym: º Catastoma luteolum Lloyd, Mycological Writings 2(Letter 11): 4. 1919.

Type: The holotype material is located at the U.S National Fungus Collections (BPI) catalog #706640, (Lloyd Myc. Collection #37839). The holotype material is from El Dorado Co., Lake Tahoe, California, collected by PB Kennedy, July 1905. There is an isotype fragment at the New York Botanical Gardens, specimen ID #291985.

Gasterocarp 17-19 mm tall x 16-22 mm broad; globose to subglobose; rhizomorph absent: ostiole developing through thinning and lateral cracking of the peridium layers, edges of the opening worn and fimbriate. Exoperidium white to cream buff (5A3), color hard to discern due to incrusted soil debris; composed of a floccose layer of granulose fibers, thin and readily sloughing off in flakes; incrusted with soil, sand and vegetal debris. Endoperidium starting out as cream to light yellow (4A3-4), becoming grayish yellow, to orange yellow (4B5-4C7), turning yellowish brown with maturity (5D8-5E8); thin, smooth, flexuous and persistent, eventually forming a cup-like shape as the fruitbody matures. Gleba tawny olive to light brown (5D5-5E6); clumped cottony to powdery. Subgleba absent. Sand case absent. Macromorphological description derived from the dried type specimen

            Basidiospores globose; (2) 5-7.5 X (2.5) 5-7 µm [xmr = 5.8 X 5.5 µm, xmm = 5.8 ± 0.0 X 5.6 ± 0.0 µm, Q = 1.0-1.3, n = 35, s = 1]; golden yellow in wet mounts; seemingly smooth under the light microscope, with SEM spores covered in flattened, partially raised, and spread apart asperate warts; oil drop absent; spores thick-walled; pedicel reduced, up to 0.8 µm long, with a clean break where disarticulated from the sterigma; free-floating sterigmata remnants absent from wet mounts; spores of various size in wet mounts. Eucapillitium Disciseda-type; threads up to 3 um broad with walls up to 0.8 um thick; pale golden yellow to hyaline in wet mounts; fragile; glabrous, short and un-branched segments, slightly sinuous, knob-like projections absent; eucapillitium tips rounded and not attenuate. Pores round, abundant, and clustered. Septa present, disarticulating at the septum wall. Paracapillitium absent. Exoperidium composed of swollen sphaerocyst cells, tightly packed together and incrusted with soil and vegetal debris; textura globulosa. Endoperidium composed of thin-walled, septate, tightly woven hyphal threads; textura intricata.

Habitat: Growing on soil in small gregarious groups, but not fused. Among the Geographic Subdivisions of California, this species is found in the Sierra Nevada.

Distribution: Known only from El Dorado County, California.

Material Examined: CALIFORNIA, El Dorado Co.: Lake Tahoe, terrestrial, July 1905, P.B. Kennedy (Lloyd Myc. Collection #37839; BPI 706640; TYPE).

Comments:  P.B. Kennedy, (type collection notes, 1905), states that this specimen has the “general appearance of being a yellow specimen of Bovista plumbea, but the internal structures is that of a Catastoma.” The exoperidium is very different from that of all other Disciseda species, being thin and of the nature of a cortex. Very close in general character to Catastoma hypogeum of Australia, which however has slightly roughened spores. The material from this type collection was omitted from the ITS sequence data due to poor sequences. Disciseda luteola needs further collection of fresh material at all ranges of maturity, and multi loci gene analysis to better understand the nature of this puffball and to further clarify the taxonomic position of it within the Lycoperdaceae.

 

 

Disciseda subterranea (Peck) Coker & Couch, Gasteromycetes E. US & Canada: 141. 1928.           

                                                                                                            (fig. 23, 40, 63)

Basionym: º Bovista subterranea Peck, Botanical Gazette, Crawfordsville. 4(10): 216. 1879.

            º Catastoma subterraneum (Peck) Morgan, 1892.

            Type: The holotype material for Bovista subterranea was collected by C.W. Irish in the Dakota Territory, in July of 1879.            

            Gasterocarp 10-28 mm tall x 16-21 (28) mm broad; globose to subglobose, creasing and folding in on its self and deflating with age, flattens as it looses spore mass; rhizomorph absent; ostiole slow to develop, round to oval when young, with a fimbriate margin having upturned fibers, becoming more irregular in shape and torn in appearance with age, sometimes more than one hole develops, exoperidium worn away at the edges revealing a darker endoperidium. Exoperidium white at first, becoming orange white (5A2), to pinkish buff (5A3), to brownish orange (5C4), and darker with age to yellowish brown (5E5), to dark yellowish brown (5F5), remaining darker towards the base where attached to the sand case, with an uneven mottled coloration overall; some specimens bleaching in the sun and returning to shades of creamy white orange gray (5B2) in old age; thin, smooth, cracked, where cracked showing the endoperidium, mostly adherent to the endoperidium throughout maturation. Endoperidium white at first, becoming orange white (5A2), to pinkish buff (5A3), to brownish orange (5C4), becoming dark chocolate brown (5F5-6F5), darker towards the base where attached to the sand case; up to 1mm thick, showing through cracks and where patches of exoperidium have sloughed off; persistent throughout maturation, thin, flexible, parchment-like. Gleba white at first, becoming cream orange white (5A2), then darkening with age to light yellowish brown (5D5), then to dark brown (6E7-6F7); the endoperidium and gleba becoming more or less the same color with age; gleba firm, cottony to compressed powdery falling apart in clumps. Subgleba absent. Diaphragm absent. Sand case starting out as white, thick and cottony; up to 2 mm thick; becoming the same color as the soil; sand case leaving a round mark on the fruitbody when becoming detached, will pop off when dry; some fruitbodies with only a small disc of sand case at the base remaining in late maturation, or covering up to 1/2 of the entire fruitbody (Coker & Couch 1928); wearing away until only a small patch remains at the very bottom of the fruitbody.

Basidiospores globose to broadly ellipsoidal; (6.4) 7.2-8.6 (9.6) X (6.4) 7.2-8.6 (9.6) µm [xmr = 7.6 - 8.0 X 7.5-8.0 µm, xmm = 8.0 ± 0.3 X 8.0 ± 0.4 µm, Q = 0.9-1.3, Qmr = 1.0, Qmm = 1.0 ± 0.0, n = 35, s = 2]; spores dark amber brown in wet mounts, golden amber in KOH; roughened with obvious echinate pin pricks under light microscope, with a dense covering with tall echinate protuberances under SEM, echinae fusing at the tips and granulose-like in texture; oil drop absent; spores thick-walled; pedicel up to 1.6-2.4 mm long, a few measuring up to 5 mm in immature specimens, most spores without, pedicels hyaline in wet mounts; free-floating sterigmata absent from wet mounts; spores of various size under light microscopy. Eucapillitium Disciseda-type; threads (3) 4.8-8.8 mm broad with walls ± 0.8-1.6 mm thick, larger threads with heavy pigmentation; hyaline to bright golden yellow in wet mounts; fragile; sinuous, dichotomous branching scarce, smooth, with knob-like projections; tips blunt with a round terminus. Pores absent or scarce, very tiny and round when present. Septa present, readily disarticulating at the septum wall. Paracapillitium absent. Exoperidium a combination of textura intricata and textura globulosa; composed of tightly interwoven hyphal threads, cellular debris, and swollen irregular-shaped sphaerocyst cells difficult to discern. Endoperidium textura intricata; composed of thin-walled, septate, tightly woven hyphal threads.

            Habitat: A rare species, seemingly preferring arid climate, and the higher elevations. Found partially submerged in the soil, subhypogeous, in dry clay soil among living low grass or among dead low-laying grass, in sandy and gravel soils, in sand dunes, on humus in sandy fields, on hard packed paths with heavy foot traffic, in open prairie, under a shallow layer of packed sand, on barren soil, in tall grass, in sand washes along eroded hillsides, dry creek beds, and resembling bird scat droppings on the ground. Growing gregariously in large groups of three to ten individual fruitbodies, but not growing fused together, and covering up to three to five square feet. Among the Geographic Subdivisions of California, this species has been found in the Modoc Plateau, in Central Western California, and in the Desert Province.

Distribution: Known from many parts of the United States, and previously reported from Arizona, California, Colorado, Hawaii, Idaho, Kansas, Michigan, Minnesota, Nebraska, New Jersey, New York, North Carolina, Oregon, South Carolina, South Dakota, Texas. Also known from Canada, Baja California, Mexico, and Hungary. 

Material Examined: CALIFORNIA, Contra Costa Co.: Wildcat Canyon, in bare soil along trail, partially or wholly covered by soil, 3 June 1939, coll. unknown (UCB620889)(UC). Inyo Co.: Inyo Mountains above Deep Springs Lake, found under Pinyon pine, on a rocky slope, May 1991, S.M. Rusiniak (Rusiniak SM). Modoc Co.: Blue Lake and Jess Lake, in soil under junipers along the road, 5 June 1979, H.D. Thiers (HDT 39883).

            Comments:  Disciseda subterranea was described under a different name at least six times in the late 19th century, and does not always grow subterranean as described in the literature (Lloyd 1908). Disciseda candida is often misidentified as Disciseda subterranea due to the spores being coarsely warted, uniformly globose, apedicellate (under light microscopy), and of an overall gasterocarp size that fits within the literature describing D. subterranea (Mitchel 1975). These two species are reported as growing side by side, and often collected with each other (Mitchel 1975). Since the gasterocarps look almost identical to the untrained eye, especially young basidiocarps, close examination of each fruitbody is important for proper determination of species. Disciseda subterranea can be recognized by: echinate spores that look bumpy with clumpy globs in KOH mounts in light microscopy, spores with a short apiculus, pores on the eucapillitium will be absent or scarce, and the gasterocarps should be growing subhypogeous. If the fruitbodies are not partially submerged in the ground, check the spores carefully for echinate ornamentation. Disciseda candid has eucapillitium with pores that are abundant in clusters, and verrucose spores. Spores mounted in KOH, of D. candida and D. subterranea, for a side by side comparison, will reveal a halo around the spores. Disciseda subterranea has more prominent halo due to larger echinate ornamentation, versus the thinner halo from the verrucose spores of D. candida, (see fig. 18.3 and 23.3 for a comparison). The ITS data here supports this species outside of the Lycoperdaceae, nested with Disciseda uplandii, with 100% bootstrap and 100% PP support. The ITS data shows Disciseda as a polyphyletic assemblage, at four points in the tree. This suggests that additional genes of multiple species of Disciseda should be sequenced to further clarify the taxonomic position of the genus amongst the other Lycoperdaceae.

 

Disciseda uplandii (Lloyd) Zeller, Mycologia 39(3): 311. 1947.                                                                                                                                                                                    (fig. 24, 64)

Basionym: º Catastoma uplandii Lloyd, Mycol. Writ. 6(Letter 61): 897. 1919.

            Type: The holotype material is located at the U.S National Fungus Collections (BPI), catalog #706792, (Lloyd Mycological Collection #8785), from San Bernardino Co., Upland, California, collected by I.M. Johnston, 25 December 1918.

Gasterocarp 11-15 mm tall x 21-23 mm broad; globose to depressed globose, creasing and folding in on its self and deflating with age, flattens as it looses spore mass; rhizomorph absent; ostiole slow to develop through maturation and thinning of the peridium layers, fine cracks appearing with minute fibrils along the margin, which separate to form a lateral to stellate-shaped stoma. Exoperidium dull brownish gray (7D2, 7E2-3), some purplish gray tints (13D2-13E2) at the top and darker purplish tints at the base (14E2), with a dull sheen; thin, creased, smooth, peeling, incrusted with sand and soil debris. Endoperidium dark brown (7F7) to reddish brown (8F7); tough and persistent, smooth, and dull. Gleba white at first, turning grayish yellow (6D3), then slightly brown when ripe (6E7); firm when young, becoming pulverulent with age. Subgleba present, very reduced and compact. Diaphragm absent. Sand case up to 1mm thick, persistent, flaking off in patches, the lower 1/4 remaining attached, sometimes covering up to 1/2 of the base of the fruitbody. Macromorphological description derived from the dried type specimen. 

Basidiospores globose; 6.5-9.5 X (6) 7-8.5 (10) µm [xmr = 7.4 X 7.2 µm, xmm = 7.4 ± 0.0 X 7.2 ± 0.0 µm, Q = 1.0 - 1.1, Qmr = 1.0, Qmm = 1.0 ± 0.0, n = 35, s = 1]; golden yellow in wet mounts; roughened with very minute bumps under light microscopy, ornamented with large, round verruculose knob-like bumps under SEM imaging; oil drop absent; spores thick-walled; pedicel reduced ± 0.5 µm, broad, with clean broken ends seen in SEM, not obvious in light microscopy; free-floating sterigmata absent from wet mounts; spores of equal size under light microscopy. Eucapillitium Disciseda-type; up to 4 um broad with walls up to 0.8 µm thick; golden to hyaline in wet mounts; broken into short hyphal strands, slightly sinuous, dichotomous branches scarce, glabrous, knob-like projections absent; tips with a blunt and round terminus. Septa abundant, disarticulating at the septum wall. Pores abundant, round and punctate, clustered to spread throughout. Paracapillitium absent. Exoperidium a combination of textura intricata and textura globulosa; composed of tightly interwoven hyphal threads, cellular debris, and swollen irregular-shaped sphaerocyst cells difficult to discern. Endoperidium textura intricata; composed of thin-walled, septate, tightly woven hyphal threads.

Habitat: Terrestrial in open grass at 365 meters (1,200 feet) elevation. Growing in abundance about the margin of a marsh, in a large gregarious group, but not fused, (Johnson, type specimen notes, 1918). Among the Geographic Subdivisions of California, this species is known form Southwestern California province.

Distribution: Known only from California and previously reported from Sacramento County, San Bernardino County, and Riverside County.

Material Examined: CALIFORNIA, San Bernardino Co.: Upland, terrestrial, 25 December 1918, I.M. Johnston (Lloyd Myc. Coll., No. 8785; BPI 706792; TYPE).

            Comments:  Lloyd (1922) notes that this species is very close to Catastoma muelleri from Australia, known only by type specimen examination from KEW; spores being the main difference. Disciseda uplandii and Catastoma muelleri both have rough spores under light microscopy, however, the spines on C. muelleri spores are three to four times as long as those of D. uplandii. It could be safe to assume with this notation from Lloyd, that under SEM imaging, C. muelleri would have echinate spores, whereas D. uplandii has verrucose spores. “These are the only two species in the purplish series with apiculate spores, all others are pedicellate,” Lloyd, (1922). Disciseda uplandii differs from other Disciseda species in that it has a reduced sterile base and abundantly pitted eucapillitium. Unlike other Disciseda, which do not have a sterile base, and may have pits that are absent, scattered or clustered, but pits not as abundant as in D. uplandii. This character of having abundant pits mirrors that of a Lycoperdon species. Zeller (1947) notes that Disciseda uplandii has a gleba that corresponds to the Mesophelliaceae and the Geastraceae, without notation of what specific gleba character he is referencing. The ITS data here supports this species outside of the Lycoperdaceae, nested with Disciseda subterranea with 100% bootstrap and 100% PP support. The ITS data shows Disciseda as a polyphyletic assemblage, at four points in the tree. This suggests that additional genes of multiple species of Disciseda should be sequenced to further clarify the taxonomic position of the genus amongst the other Lycoperdaceae. Disciseda uplandii, without further collection of material at all ranges of maturity, and multi loci gene analysis, this species will remain incompletely understood.

 

LYCOPERDON Pers.:

Lycoperdon curtisii Berk., Grevillea 2(16): 50. 1873.

                                                                                                                        (fig. 25, 65)

Reported synonym

            = Vascellum curtisii (Berk.) Kreisel, Feddes Repert. Spec. Nov. Regni Veg. 68: 86. 1963.

            Type: According to the curators at Kew, there are three type collections, two being syntypes according to the protologue, Grevillea 2(16): 50 1873; distributed as part of North American Fungi Exsiccata, no. 333. The first syntype was collected in Connecticut by C. Wright, (K(M) 189271, Wright No. 5613). The second is from North Carolina, collected by M.A. Curtis, (K(M) 189272, Curtis No. 558), although Berkeley notes on the specimen sheet that 5613 = 558.. The third is collected from North Carolina by M.A. Curtis, (K (M)189273, Curtis No. 2195). Collections dates are not noted and therefore unavailable.

Gasterocarp 5-20 mm tall x 5-20 mm wide; globose, or subglobose to ovate; rhizomorph composed of a tuft of branching mycelium at the base of the gasterocarp; ostiole slow to develop through stretching of the apex as the fruitbody expands with maturation, opening as a slit-like crack, developing into a round opening that exposes the gleba. Exoperidium white to yellow white (5A1-2) when young, becoming light beige yellow to yellow grey (4A3-4B2), remaining light in color through maturation; covered in minute echinae or sharply pointed spines, crowded, fused at the tips, sharply pointed, tips becoming appressed with age, tips darkening to brown or dark brown (6E7-6F7), fine white granular material present between spines; spines sloughing away at the apex initially, completely sloughing away through maturation, not leaving scars on the endoperidium, powdery-like granular coating remaining in old age. Endoperidium white when young to yellowish grey (5A1-2), becoming light yellowish brown (5D5); persistent and parchment-like when old, remaining mostly attached to the exoperidium. Gleba white and firm when young, becoming brownish orange (5C4-7), olive brown to dark brown (4F5-6F7) when mature; mixed-type, cottony in texture when young to powdery in old age. Subgleba grayish brown (8E3) to purple grey (14E3) metallic tinted when mature; mixed-type, chambered, compact, present only at the very base of the fruitbody, umbonate to convex or slightly radiating up into the gleba in center base of the subgleba. Diaphragm well defined, composed of compressed subgleba cells, becoming tough and skin-like, separating the gleba from the chambered subgleba.

            Basidiospores globose to subglobose; 2.5-4 X 3-4 µm [xmr = 2.6-3.6 X 3.3-3.4 µm, xmm = 3.0 ± 0.6 X 3.4 ± 0.1 µm, Q = 0.7 - 1.3, Qmr = 0.8-1.1, Qmm = 0.9 ± 0.2, n = 25, s = 3]; chocolate brown in wet mounts, turning dark olive green in KOH; minutely spiny to punctate and heavily ornamented under light microscopy, SEM spores echinulate with tall pointed echinae connected by chord-like bridges; central oil droplet present; spores thick-walled; pedicels rudimentary, < 0.8 µm or absent under light microscopy, seen with SEM as short blunt pedicels with clean broken ends; free-floating sterigmata absent from wet mounts; spores of equal size under light microscopy. Eucapillitium Lycoperdon-type; threads 3-7 µm broad with walls up to 0.8 µm thick; mostly colorless in wet mounts, slight yellow tint in KOH; elastic; incrusted with cellular debris, dichotomously branching, mostly straight, without knob-like projections, not attenuate with a blunt round terminus; restricted to the periphery of the gleba along the inside of the endoperidium wall, not abundant, absent from very mature fruitbodies. Pores absent. Septa absent. Paracapillitium abundant throughout the gleba, dominant form of capillitial threads; thin-walled, hyaline, septate, pores absent, incrusted with cellular debris. Diaphragm composed of compact, globose, randomly aligned cells. Exoperidium textura globulosa; composed of sphaerocyst cells, inflated, long, thin-walled, hyaline, septate, and in linked chains. Endoperidium textura intricata; composed of interwoven hyphal threads.

Habitat: Growing in caespitose clusters and gregarious groups, often forming odd shapes as the fruitbodies grow around and into each other, often two to three fruitbodies fused together. Growing in irrigated lawns, shaded areas, on soil, and in conifer duff. Among the Geographic Subdivisions of California, this species is known from Northwestern California, the Great Central Valley, and the grassy lowlands of the Sierra Nevada.

            Distribution: Known from many parts of the United States, and previously reported from Alaska, Arizona, California, Colorado, Georgia, Hawaii, Idaho, Illinois, Indiana, Iowa, Kansas, Nebraska, Nevada, New Hampshire, New Jersey, New Mexico, New York, North Carolina, North Dakota, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Ohio, Oklahoma, Oregon, Pennsylvania, South Dakota, Tennessee, Texas, Vermont, Virginia, Washington, Washington DC, West Virginia, Wisconsin. Also known from Canada, Bahamas, Bermuda, Mexico, Philippines, Puerto Rico, Sweden, and Vieux-Fort on the island of St. Lucia.

Material Examined: CALIFORNIA, Sonoma Co.: Windsor, terrestrial in lawn near a creek, 08 June 2008, D. Deshazer (SSJ 247); Freestone, SOMA camp, CYO Catholic Charities camp, Bohemian Highway, in grass along baseball field, January 2012, D. Deshazer (SSJ 400). Yolo Co.: Sacramento, in lawn, 9 September 1937, E.E. Morse (UCB589747)(UC). Yuba Co.: Marysville, terrestrial in a field, 21 February 2009, R. Hopson (SSJ 474).

            Comments:  Lycoperdon curtisii Berk. was once distinguished as Vascellum curtisii (Berk.) Kreisel, due to Smarda’s Vascellum distinction of a paper-like diaphragm separating the gleba from the subgleba. Among the distinctive characters defining species within Vascellum, Kreisel added eucapillitium being replaced by a dominance of paracapillitium within the gleba tissue, and a reduced pseudocolumella (Kreisel 1963, Larsson 2008). This North American species, Lycoperdon curtisii, has been considered identical to European collections of Vascellum pratense (Pers.) Kreisel by Hollós, except that V. curtisii has a thinner diaphragm, a smaller subgleba, and is overall reduced in size (Kreisel 1963, Ponce de Leon 1970). However, there are many features that distinguish these two species as being different. For one, the spore ornamentation seen with SEM is remarkably different between Lycoperdon curtisii and L. pratense, not to mention the slight difference in spore size, and the striking difference in overall size. In addition, the type for the genus Vascellum, V. depressum (Bonord.) F. Šmarda, has been accepted as a synonym of Lycoperdon pratense Pers. (Larsson 2008). This transfer reduces the genus Vascellum to synonymy with Lycoperdon.

            The ML tree produced in this analysis shows Vascellum species (now in Lycoperdon) as a monophyletic clade within the Lycoperdon clade, with 92% bootstrap and 100% PP. In support of the claims above, Vascellum curtisii is placed within this Vascellum clade, and shows that it is distinct from Lycoperdon pratense. Two California specimens identified as Lycoperdon curtisii (SSJ 247, and SSJ 400) are distinct from two GenBank sequences identified as Vascellum curtisii (HQ235048, and HQ235043), representing material from both NC and HI, respectively. This suggests that additional analysis on many specimens from multiple locations coupled with multi loci gene analysis will be necessary to further clarify the taxonomic position of Lycoperdon curtisii within the Lycoperdaceae and better understand the nature of this puffball.

 

Lycoperdon dermoxanthum Vittad., Monograph Lyc.: 178. 1843.                                                                                                                                                                                    (fig. 26, 66)

Reported synonyms

            = Lycoperdon ericetorum Pers., J. Bot. (Desvaux) 2: 17. 1809.

            = Lycoperdon ericetorum Pers., J. Bot. (Desvaux) 2: 17. 1809. var. ericetorum.

            = Globaria dermoxantha (Vittad.) Quél., Compt. Rend. Assoc. Franç. Avancem. Sci. 13: 283. 1885.


            = Utraria dermoxantha Quél., Enchir. fung. (Paris): 241. 1886.           

            = Bovista dermoxantha (Vittad.) De Toni, in Berlese, De Toni & Fischer, Syll. fung. (Abellini) 7: 100. 1888.

            = Lycoperdon ericetorum var. pusillum (Pers.) F. Šmarda, Fl. ČSR, B-1, Gasteromycetes: 321. 1958.

            = Bovista pusilla sensu auct. mult. (e.g. Kreisel); fide Checklist of Basidiomycota of Great Britain and Ireland. 2005.

            = Lycoperdon pusillum sensu auct. mult.; fide Checklist of Basidiomycota of Great Britain and Ireland. 2005.

            Type: Italy. Type specimen not designated in the protologue or reported elsewhere.

            Gasterocarp (6) 12-25 mm tall x (8) 10-25 (35) mm broad; globose to depressed globose, sometimes obpyriform, tapering to a point where attached to the rhizomorph, plicate at the base; rhizomorph composed of finely entangled hyphae, persistent, soil incrusted, rooting the fruitbody; ostiole developing as a lateral torn opening, becoming stellate in age, the edges not lifting or curling, often irregularly split. Exoperidium cream when fresh to light yellow when dry (4A2-3), becoming yellowish brown to dark yellowish brown (5F5-6) with age; persistent and remaining adherent to the endoperidium initially, cracking in age as the exoperidium expands and dries, breaking up into appressed hexagonal patches that slough off with age, edges up-lifting when peeling away from the endoperidium, coated with a granular material that sloughs off early in development, incrusted with soil at the base, cracking open age and desiccation to expose the endoperidium. Endoperidium white at first, becoming brown to yellowish brown, then dark yellowish brown (5D7, 5E6-7, 5F5), remaining darker than the exoperidium; persistent, parchment-like, glabrous. Gleba white at first, turning orange white (5A2), orange yellow (4B5-6), grayish yellow (5D3), then yellow brown (5E4-5), and eventually dark olive brown (4F7-5F5) at maturity, lighter in color at the base when mature; firm when young, becoming cottony at mid-maturity, and powdery when completely mature. Subgleba absent. Diaphragm absent.

            Basidiospores globose; (3.2-) 4-7 X 4-7 µm [xmr = 4.4-6.6 X 4.3-6.6 µm, xmm = 5.46 ± 1.6 X 5.44 ± 1.6 µm, Q = 0.8-1.4, Qmr = 1.00-1.02 Qmm = 1.01 ± 0.0, n = 25, s = 2]; turning golden bright yellow to brown in KOH mounts, yellow in wet mounts; seemingly smooth with immature spores, could lead to misidentification, faintly verrucose in KOH mounts with very mature spores under light microscopy, verrucose ornamentation with well-distanced spines under SEM, verrucose spines prominent raised columns with a flattened apex; oil drop present; spores thick-walled, up to 0.8 µm thick; pedicels < 0.8 µm long, short and stubby in light microscopy, torn with uneven ends under SEM; free-floating sterigmata absent from wet mounts; spores of equal size under light microscopy. Eucapillitium Lycoperdon-type, Intermediate-type eucapillitium present in the center of the gleba; walls up to (0.8) 1.0-4.8 (5.8) µm broad, 7.2 µm broad at branching junctures, thread walls up to (0.8) 1.6-4.8 µm thick; golden brown in wet mounts; subelastic and breaking into segments; glabrous, dichotomously branching, swollen where branching mostly straight threads, somewhat sinuous towards the tips, sinuous where thin, knob-like projections present, tips with a blunt rounded terminus, some tips swollen and club-shaped. Pores abundant, round, size variable, very small to medium. Septa abundant, absent in some mounts, threads disarticulating at the septum wall. Paracapillitium present and not abundant, septate; seen with Lactic Acid Cotton Blue reaction. Exoperidium a combination of textura intricata and textura globulosa; composed of short, branched, septate, interwoven hyphal elements, intertwined with swollen globose sphaerocyst cells. Endoperidium textura intricata; composed of interwoven hyphal threads.

Habitat: Lycoperdon dermoxanthum is found in the northern parts of California. Found solitary or in small groups, on humus, and often in sun-exposed, dry, acidic soils, and grasslands. Common in coastal regions, in grass meadows surrounding heavily wooded conifer forest, and rarely in wet spring years of the Sierra Nevada and northern parts of the Great Central Valley. Among the Geographic subdivisions of California, this species will occur in Northwestern California, the Great Central Valley, the Sierra Nevada, and in Central Western California.

Distribution: Known from many parts of the United States, and previously reported from Alabama, Arizona, California, Colorado, Florida, Indiana, Iowa, Kansas, Massachusetts, Michigan, Missouri, Nebraska, New Jersey, New York, North Carolina, North Dakota, Ohio, Pennsylvania, South Carolina, Tennessee, Utah, and Wisconsin. Also reported from Brazil, Canada, Colombia, Dominican Republic, England, France, Germany, India, Peru, Philippines, Puerto Rico, Trinidad, and Uruguay.

Material Examined: CALIFORNIA, Alpine Co.: Alpine Lake, State Highway 4, gregarious on moist sandy soil near pines, September 1992, H.D. Thiers (HDT 54220). Butte Co.: Jonesville, terrestrial, October 1941, E.B. Copeland (UCB660284)(UC). Del Norte Co.: Smith River, in sand on the river bar, November 1939, H.E. Parks (UCB139378)(UC). Humboldt Co.: Ing Fems. sugar pine area, 22 November 1976, D. Largent (DL 7252-6196)(HSU); Arcata, terrestrial, 22 November 1976, D. Largent (DL 76-304-6198)(HSU). Marin Co.: Bolinas, Audubon Canyon Ranch, below Bolinas, scattered on soil in grassy area along fire road, October

1979, C. Calhoun (Calhoun 79-903); Bolinas, Audubon Canyon Ranch, Pitcher Canyon, scattered on soil, November 1981, C. Calhoun (Calhoun 81-2499b); Marin, Bolinas Ridge, terrestrial under Pseudotsuga menziesii (Douglas Fir) 20 October 2009, R. Pastorino (SSJ 383). Placer Co.: Towle, Baxter’s Camp, terrestrial, 5 May 1934, E.E. Morse (UCB521239)(UC). San Francisco Co.: San Francisco State University campus, solitary in soil under pines along sidewalk, November 1997, coll. unknown (Gregory 2). Sierra Co.: Highway 89, five miles North from junction of Highway 49, on soil in meadow in seepage area, June 1984, coll. unknown (HS 3817). Siskiyou Co.: Happy Camp, terrestrial under Pinus strobus (white pine), 29 November 1976, D. Largent (DL 3-6197)(HSU); Happy Camp, USFS Sugar Pine Outplant, 2 May 1978, coll. unknown (Niesen 76-1167880)(HSU); Happy Camp, USFS Sugar Pine Outplant, terrestrial in grass, 18 September 1978, coll. unknown (Niesen 130-1167881)(HSU); Happy Camp, USFS Sugar Pine Outplant, 9 October 1978, coll. unknown (Niesen 159)(HSU); Happy Camp, USFS Sugar Pine Outplant, terrestrial, 9 October 1978, coll. unknown (Niesen 168-1167882)(HSU); Happy Camp, USFS Sugar Pine Outplant, gregarious and terrestrial in low grass, 29 October 1979, coll. unknown (Niesen 551)(HSU). Trinity Co.: Big Flat, 1,545 meters (5,072 feet) elev., gregarious in a meadow, 3 July 1937, coll. unknown (UCB585261)(UC). Tuolumne Co.: Yosemite National Park, May Lake Trail, 2,712 meters (8,900 feet) elev., in meadow, 10 August 1933, coll. unknown (UCB506723)(UC).

            Comments:  Lycoperdon dermoxanthum is a rare and often misidentified species. Having a small size and lack of a subgleba, it resembles a Bovista species. With a similar morphology and somewhat overlapping in habitat of Bovista aestivalis and Bovista plumbea, these three puffballs can be confused easily. In comparison to Bovista aestivalis, Lycoperdon dermoxanthum has a darker green gleba, a longer pedicel on the spores, more coarsely verrucose-ornamented spores, and a subgleba that is almost always absent. Both of these species have similar exoperidium and endoperidium characteristics, both have abundant pores of various size, and spores that are about 3.5-4.5 mm. In Bovista aestivalis, mature gleba tissue near the endoperidium wall will have Lycoperdon-type eucapillitium, with Intermediate-type in the center of the gleba. The gleba of Lycoperdon dermoxanthum has both Lycoperdon-type and Intermediate-type in the center of the gleba. Mature gleba tissue from both the center and outer extremities needs to be examined in order to reduce the possibilities of misidentification. Older specimens can easily be misidentified when gleba tissue is absent. The habitat for both of these species overlaps in California, yet Lycoperdon dermoxanthum seems to be more abundant along the north coast. These two puffballs were recently shown to be unrelated in phylogenetic analysis (Bates 2004, Larsson 2008), regardless of their similar morphology and habitat. Lycoperdon dermoxanthum can appear similar to Bovista plumbea in the field, with a cracked to smooth exoperidium ornamentation and similar coloration. However, older Bovista plumbea specimens have a metallic grey tinted endoperidium, and very long pedicels on the spores, sometimes up to 11 µm long. A simple check under the microscope will rule out B. plumbea specimens by looking for small pedicels versus very long ones. Lycoperdon dermoxanthum is placed in thee different locations in this ITS sequence analysis. This suggests that additional genes should be sequenced to further clarify the taxonomic position of Lycoperdon dermoxanthum amongst the other Lycoperdaceae.

 

Lycoperdon lloydianum (A.H. Sm.) Jarvis, comb. prov.

                                                                                                                        (fig. 27, 67)

Basionym º Vascellum lloydianum A.H. Sm., Bulletin Mensuel de la Société Linnéenne de Lyon: 410. 1974.           

Type: There is a holotype collection at the University of Michigan (MICH 5847, AH Smith 83267). Collected from the Cispus Environmental Center, Lewis, Washington, by AH Smith, 29 October 1972.

            Gasterocarp 2.5-38 mm tall x 2-35 mm broad; globose to depressed globose or obpyriform, oblong, tapered to the bottom, wrinkled with ridges along all sides; rhizomorph composed of thin hyphal threads clinging to soil and vegetal debris; ostiole 0.5-0.2 mm broad, peridium cracking, then tearing horizontally, or forming a stellate-shaped broad opening in the center of the apex, creating a single hole, stretch marks surrounding the tear, fragile rim, sloughing away slowly until forming a vase-shape. Exoperidium pure cream white when fresh, becoming cream buff grey to brown (4A3-4B3, 5B3-4, 5D3), mottled with uneven shades of color; covered entirely with furfuraceous spines or with minute pyramidal echinae-like scales when young, echinae erect, some scale tips fusing with others to form larger units, individual scales > 0.01 mm tall x 0.01 mm broad, sloughing off and leaving a smooth surface, scarring at the bottom in folds, scars round or oval in shape, becoming depressed or rubbing off easily, remaining in the folds and crevices, overall roughened exoperidium surface. Endoperidium brownish orange clay to tan beige (5C4, 5D5) color, shiny metallic and reflective in old age, with sections of tan brick red (4B2, 6E5), the base with lighter color than the apex; fragile when dry, glabrous, parchment-like, sloughing away with the exoperidium around the ostiole until the entire apex has sloughed off, until a vase-shaped structure remains. Gleba color white to cream buff at first (4A3), interior becoming grayish yellow (4B5) when spore maturation has initiated, to olive green (4E7) in the very center, fading to cream white at the outer extremities, then shades of dark brown (5C3, 5E7, 5F7) when mature; mixed-type, starting out firm, cottony and subelastic, hyphal threads radiating up through the center, columella-like, outer hyphal threads radiating inwards from the endoperidium wall, the two layers pull away from each other easily, becoming pulverulent in age, gleba completely blowing away once the ostiole opens and the gleba becomes powdery. Subgleba lilac grey (18C2), with a grey lilac brown, and mottled with pale yellow (5A3, 5B4), metallic tint overall, with tones of brown when looked at under a dissecting scope; prominent, up to 9-20 mm tall, taking up to 2/3 of the entire gasterocarp, a chambered honeycomb composed of thin layers of hyphal threads woven together creating thin, glossy, transparent sheets. Diaphragm composed of compact subgleba tissue, up to 1 mm thick, a skin-like border, becoming papery thin and parchment-like in old age, clearly visible inside mature sporocarps when cut vertically, not well developed in young sporocarps.

            Basidiospores globose; 3.2-6 X 4-6.2 µm [xmr = 3.5-5.1 X 3.7-5.1 µm, xmm = 4.3 ± 1.1 X 4.4 ± 1.0 µm, Q = 0.8-1.3, Qmr = 1.0-0.9, Qmm = 1.0 ± 0.1, n = 25, s = 3]; amber brown in water mounts, golden brown in KOH; roughened with rounded bumps covering entire surface, finely warted in KOH, appearing spinulose in cotton blue, spores echinulate at 1000x under oil emersion, echinulate with conical protrusions seen under SEM; oil droplet present; spores moderately thin-walled, easily deflating with desiccation; with a minute pedicel up to 0.8-2.4 um, very few with pedicel remnant broken at the base of spore, some pedicels long with a broken end, pedicels short and often recurved into the spore under SEM, some have a clean broken end or a torn tissue fragment; free-floating sterigmata absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Lycoperdon-type; hyphal threads 4-8 um broad with walls up to 0.8 um thick; amber to slightly green flecked in water mounts, golden brown in KOH mounts; elasticity semi-fragile, disarticulate readily into individual threads; heavily incrusted, non-branching or dichotomous branching scarce, hyphal threads slightly sinuous wavy, knob-like projections present; tips with a round blunt terminus; dominant tissue type around the extremities of the gleba and at the diaphragm, longer elastic threads occurring in the outer portions of the gleba, with shorter more fragile hyphal strands occurring in the inner portions of the gleba, eucapillitium growing out from the diaphragm into the gleba and found along the endoperidium walls. Pores absent and not seen with SEM imaging. Septa abundant. Paracapillitium present throughout the gleba, dominant form of capillitial threads; abundant in center of the gleba, dominant in young and old specimens; incrusted with cellular debris that has non-distinguishing shape or features, with abundant septa, rounded blunt tips, poroid; turns brilliant cobalt blue in the lactic acid cotton blue reaction, abundant septa show after 20 minutes of staining, paracapillitium turns hyaline and eucapillitium turns blue green after 30 minutes from initial staining. Exoperidium a combination of textura globulosa and texture intricata; composed of large, thin-walled, globose to subglobose sphaerocyst cells, intertwined with hyphal threads, roughened with rounded bumps, with true septa walls, branching, distinct hyphal threads with a blunt rounded terminus. Endoperidium textura intricata; composed of tightly interwoven hyphal threads, septate, dichotomously branching. Diaphragm composed of very tightly interwoven capillitium threads, some areas more thickened and tightly woven to develop vein-like threads; branching, thick-walled, golden amber brown in wet mounts. Subgleba tissue similar to diaphragm tissue, some dichotomous branching, non-pored, septum scarce, hyphae 1-layer thick, except where they converge to create a tightly woven dense sections of vein-like threads.

Habitat: A grassland species, fruiting in fall after light rain and in spring after heavy rain. Lycoperdon lloydianum will detach from the soil when mature and roll around in the environment dispersing spores. The gasterocarps will persist in the environment for up to a year. Growing in rings, caespitose, in groups, or solitary. Among the Geographic Subdivisions of California, this species is found in the Northwestern California, and in the Central Western California.

Distribution: Known only from the Western United States, and previously reported from Arizona, California, Colorado, Oregon, and Washington.

            Material Examined: CALIFORNIA, Fresno Co.: Alameda Co.: Oakland, Knowland Park, above the Oakland Zoo, terrestrial in grass, gregarious in rings, 19 April 2012, S.S. Jarvis (SSJ 438). Humboldt Co.: Arcata, coastal, 17 November 1977, T. Baroni (Baroni 772-5239)(HSU). Marin Co.: Audubon Canyon Ranch, Galloway Canyon, gregarious in disturbed grassy area, 05 October 1977, C. Calhoun (Calhoun 77-73); Audubon Canyon Ranch, Pitcher Canyon, scattered in grass, 9 November 1979, C. Calhoun (Calhoun 79-914); Audubon Canyon Ranch, Volunteer Canyon, scattered in open area, 11 December 1980, C. Calhoun (Calhoun 80-1722); Point Reyes National Seashore, Limantour Ridge, solitary, terrestrial at edge of Pseudotsuga menziesii (Douglas fir) grove, 18 October 1981, coll. unknown (PDB 46); Mt. Tamalpais, Muir Beach overlook, in grass, 08 January 2008, S.S. Jarvis (SSJ 241). Napa Co.: Knights Valley, Foote Ranch, 07 November 2008, S.S. Jarvis (SSJ 238); Knights Valley, Foote Ranch, East facing grass slope, under a big Quercus lobata (valley oak), 05 November 2009, S.S. Jarvis (SSJ 388); Healdsburg, Stuhlmuller Vineyards, on upper hills overlooking the Russian River, 5 November 2012, S.S. Jarvis (SSJ 479); Knights Valley, Foote Ranch, numerous, terrestrial in grass, 15 November 2012, S.S. Jarvis (SSJ 478); Napa, Alston Park, terrestrial in grass, 27 December 2012, S.S. Jarvis (SSJ 477). San Mateo Co.: San Mateo County Park, in grass, 3 December 1986, (Burwen s.n.). Solano Co.: Fairfield, Rockville Heights Regional Park, 26 December 2009, R. Pastorino (SSJ 397). Sonoma Co.: Sebastopol Memorial Lawn Cemetery, in grass, 08 January 2003, D. Deshazer (SSJ 339); Knights Valley, Foote Ranch, North upper ranch slopes, at drip line of Quercus geminata (live oak), 16 October 2011, S.S. Jarvis (SSJ 407); Knights Valley, Foote Ranch, scattered on Northern ranch grassy slopes, 20 October 2011, S.S. Jarvis (SSJ 414).

            Comments: The diaphragm-forming Lycoperdon puffballs have similar features and it may be difficult for amateur mycologists to tell them apart. A good microscope with oil emersion capability at high power is necessary for identifying eucapillitium versus paracapillitium from various parts of the gleba. One main distinction that sets Lycoperdon lloydianum apart from other diaphragm-forming puffballs is the unique feature of eucapillitium growing out from the diaphragm into the gleba. Lycoperdon pratense and Lycoperdon lloydianum are very similar in outward appearance, which can cause confusion and misidentification among them. Lycoperdon pratense has non-poroid eucapillitium that is scarce to absent, and can be found only along the endoperidium wall in mature gasterocarps, as long as gleba tissue is still present. As well, Lycoperdon pratense has a wide and very well developed diaphragm that can be seen earlier in development than that in Lycoperdon lloydianum (Ponce de Leon 1970, Bates 2004). Both of these grassland species have been collected together in the same habitat. Close examination of the diaphragm and growth of the eucapillitium is necessary for proper treatment of these species. Lycoperdon curtisii remains small in size (under 20 mm broad) and has Lycoperdon-type eucapillitium that can be found along the periphery of the endoperidium wall. This small puffball should be easier to separate from Lycoperdon lloydianum collections due to smaller size when mature, reduced subgleba, and smaller globose spores. Based on ITS, the ML tree includes V. lloydianum and V. intermedium in Lycoperdon. Several of these species have already been transferred; Lycoperdon asperrimum, Lycoperdon curtisii, Lycoperdon bicolor, and Lycoperdon pratense. Whereas others represented in this molecular data set may need to be transferred to Lycoperdon: V. intermedium and V. lloydianum.

 

Lycoperdon molle Pers., Syn. meth. fung. (Göttingen) 1: 150. 1801.                                                                                                                                                            (fig. 28, 39, 68)

Reported synonyms

= Lycoperdon gemmatum var. furfuraceum Fr., Syst. mycol. (Lundae) 3(1): 38. 1829.


= Lycoperdon gemmatum var. molle (Pers.) De Toni, Syll. fung. (Abellini) 7: 107. 1888.

            = Lycoperdon atropurpureum sensu auct.; fide Checklist of Basidiomycota of Great Britain and Ireland. 2005.

            = Lycoperdon umbrinum sensu auct.; fide Checklist of Basidiomycota of Great Britain and Ireland. 2005.

Type: There is type material listed at the National Herbarium Nederland (L 0054569, L 00545070), without any notes regarding collection dates or location. According to Kreisel (1962), the type location is from central Europe.

Gasterocarp 25-80 (110) mm tall, 25-50 (75) cm broad; obpyriforme to turbinate, developing an umbo at the apex in young maturity; rhizomorph up to 12.5-30 mm long, rooting the fruitbody to the soil, attaching to the substrate with thin fine hyphal rhizoids; ostiole late forming, developing through a small torn hole at the apex of the umbo, becoming larger and round to stellate as the fruitbody matures, with upturned and frayed edges, sporocarp dehiscing via the apical pore. Exoperidium cream to pale pinkish buff (4A3-5A3), grayish yellow (4B5) at the base with light brown (5D5) scattered cottony coating when very young, maturing to tan clay color (5C5), light yellowish brown (5D5), grey-brown to medium-brown (6D3-6E7); exoperidium coated with a mixture of granules and short furfuraceous-spinulose hair-like spines that easily slough off with rain and handling, hair-like spines solitary or grouped with fused tips, lighter colored soft spines at the apex with darker pigmented spines towards the base, ornamentation wearing off with age to expose the endoperidium, not forming scars. Endoperidium grayish orange to clay color (5B5-5C5); thin, persistent, and parchment-like when mature. Gleba firm and white when young, turning shades of yellowish-green (4C5-4E5), becoming dark brown with maturation (6F4-7); flesh granular or brittle, becoming cottony with age. Subgleba cream colored (4A3), partially dark olive-brown (4F5) with metallic lilac tints (11E4), unevenly colored with more pigmentation at the outer edges and topmost section; taking up to half of the fruitbody, as a well developed pseudostipe, exterior pleated at the base, the interior with large alveolate chambers, most often wide and distinct, rarely narrowed or rudimentary. Diaphragm absent.

            Basidiospores globose to subglobose; 5.2-7.2 X 5.4-7.2 µm, [xmr = 5.8-6.1 X 6.8-7.0 µm, xmm = 6.1 ± 0.2 X 7.0 ± 0.2 µm, Q = 0.8-1.0, Qmr = 0.8, Qmm = 0.9 ± 0.0, n = 30, s =2]; golden brown in wet mounts; distinctly verrucose warted, under SEM with very prominent raised verrucose columns with flattened tops; with a central oil droplet; spores thick-walled; pedicel nub-like, < 0.5 µm in length; free-floating sterigmata remnants common in wet mounts; spores of equal size in light microscope. Eucapillitium Lycoperdon-type; eucapillitium size 4.0-8.0 µm in diameter with walls up to 1.0 µm broad; golden brown in wet mounts; elastic; glabrous, frequent dichotomous branching, straight to occasionally subundulate, with knob-like projections; eucapillitium tips attenuate to a narrow terminus. Pores abundant, relatively small, round to oval. Septa absent. Paracapillitium scarce to absent. Exoperidium textura globulosa; composed of globose inflated sphaerocyst cells. Endoperidium texture intricata; composed of tightly woven hyphal threads with abundant septa.

Habitat: Found widely distributed in North America. Collected on soil or humus in deciduous or coniferous forests, and in live oak coastal communities. Often found growing terrestrially on sandy soil or in semi-grassy areas near Oak tree species, in disturbed soil and along trails. Frequent but not abundant, found in small groups or singularly. Fruiting in the late fall and early spring with heavy rains. Among the Geographic Subdivisions of California, this species is found in Northwestern California, the Cascade Range, Central Western California, and in Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Alabama, Arizona, California, Colorado, Connecticut, Florida, Indiana, Iowa, Kansas, Kentucky, Maine, Maryland, Massachusetts, Michigan, Missouri, New Jersey, New Mexico, New York, Ohio, Oregon, Pennsylvania, South Dakota, Texas, Vermont, Virginia, Washington, West Virginia, Wisconsin, and Wyoming. Also known from Baja California, Canada, Czech Republic, France, Germany, and Mexico.

            Material Examined: CALIFORNIA, Alameda Co.: Oakland, terrestrial in woods in the Oakland Hills, 2 March 1934, coll. unknown (UCB521367)(UC); Berkeley, Strawberry Canyon, terrestrial, 22 November 1937, coll. unknown (UCB585370)(UC); Oakland, Sequoia Park, terrestrial, 22 November 1937, V. Mentzer (UCB585708)(UC); College Pines, above Oaklane, under edge of a patch of Rubus occidentalis (blackberry), 9 March 1938, V. Miller (UCB585840)(UC); Piedmont, Oakland Hills, in deep duff of pine forest, 12 November 1944, T.T. McCabe (UCB977180)(UC); Berkeley, University of California campus, Astronomy Hill, 7 December 1972, T. Tang (UCB1445925)(UC). Del Norte Co.: Darlingtonia, terrestrial under conifers, November 1937, H.E. Parks (UCB637234)(UC); Darlingtonia, Smith River, under Libocedrus decurrent (incense cedar) and Pseudotsuga menziesii (Douglas fir), 2 November 1939, H.E. Parks (UCB1139348)(UC). Humboldt Co.: Grays Falls Campground, terrestrial in mixed woods, 15 November 2009, S.S. Jarvis (SSJ 454). Marin Co.: Phoenix Lake, terrestrial, 4 February 1935, V. Mentzer (UCB528870)(UC); Inverness, under Quercus agrifolia (live oak) and Umbellularia californica (California bay laurel), 20 February 1940, coll. unknown (UCB62534)(UC); Audubon Canyon Ranch, Pitcher Canyon, solitary on soil under Quercus agrifolia (coast live oak) and mixed evergreen forest, 23 November 1977, C. Calhoun (Calhoun 77-281); Audubon Canyon Ranch, Pitcher Canyon, scattered in soil under mixed evergreen forest, 7 December 1977, C. Calhoun (Calhoun 77-423); Mount Tamalpais, Marin watershed district, Simmons Trail, solitary near Arctostaphylos hookeri ver montana (Mt. Tam Manzanita), 22 Jan 1997, J.R. Blair (JRB 393). Mendocino Co.: Little River, in thick Sequoiadendron giganteum (redwood) forest, 5 November 1950, P. McMillan (UCB977201)(UC). Monterey Co.: Fort Ord, on ground in grassy area under Quercus agrifolia (live oak), 11 February 2010, R. Pastorino (SSJ 402); Fort Ord, on the side of the road, terrestrial under Quercus agrifolia (live oak), 18 February 2010, R. Pastorino (SSJ 401). Napa Co.: Cleary Reserve, solitary in humus under pine, 23 November 1963, H.D. Thiers (HDT 10808); Cleary Reserve, scattered to gregarious in humus in dense mixed woods, 24 November 1963, H.D. Theirs (HDT 10831). Placer Co.: Folsom Lake, along a hard packed trail, in grass in full sun, March 2012, L. Maddison (SSJ 433). San Diego Co.: Marian Bear Memorial Park, near the beach in dark sandy soil near decomposing wood, 7 February 2009, S.S. Jarvis (SSJ 318). San Francisco Co.: Golden Gate park, terrestrial, December 1934, M. Mentzer (UCB528277)(UC). San Mateo Co.: Crystal Springs Lake, in Umbellularia californica (California bay laurel tree) woods, 2 March 1938, V. Miller (UCB585871)(UC); La Honda, YMCA Camp, on soil, 9 January 1961, D. Largent (Largent 380); San Francisco Watershed, gregarious in soil under hardwoods, 30 December 1969, H.D. Thiers (HDT 24676); San Francisco Water Shed, solitary in humus under hardwood, 18 March 1970, H.D. Thiers (HDT 25303); San Francisco Watershed, solitary in humus under hardwoods, 19 March 1970, H.D. Thiers (HDT 25305); San Francisco Watershed, gregarious in soil under hardwoods, 20 March 1970, H.D. Thiers (HDT 25292); Pescadero, Gazos Creek Campground, along a stream on a wet bank, 17 March 2009, D. Viess (SSJ 447). Santa Cruz Co.: San Lorenzo River, one mile East of Boulder Creek, in dripping wet Sequoiadendron giganteum (redwood) forest on steep riverbank, 6 December 1937, V. Miller (UCB585827)(UC); Boulder Creek, terrestrial, 20 December 1937, L. Bonar (UCB585363) (UC); Boulder Creek, solitary in Quercus agrifolia (live Oak) woods, on sandy loam soil and madrone duff (Arbutus menziesii), 10 April 1938, T.T. McCabe (UCB585874)(UC); Boulder Creek, in deep soft soil in Arbutus menziesii (madrone duff), 20 November 1938, coll. unknown (UCB1138802)(UC). Shasta Co.: Lassen National Park, Paradise Meadow trail to Hat Lake, terrestrial at 2,133 meters (7,000 feet) elev., 12 July 1966, V.G. Cooke (UCB1320689)(UC). Sonoma Co.: Occidental, Camp Meeker, solitary in soil in manzanita thicket, 2 April 1961, D. Largent (Largent 524); Salt Point State Point, solitary in humus in mixed woods, 25 November 1972, H.D. Thiers (HDT 30806); Sebastopol, Pleasant Hills Memorial Park & Mortuary, Pleasant Hill Road, in grass in shade, December 2007, D. Deshazer (SSJ 246). Tehama Co.: Mineral, home of EL Adams, terrestrial, 7 September 1975, V.G. Cooke (UCB1464670)(UC); Mineral, home of EL Adams, terrestrial, 7 September 1975, V.G. Cooke (UCB1464676)(UC); Mineral, home of EL Adams, terrestrial, 7 September 1975, V.G. Cooke (UCB1464678)(UC). Tuolumne Co.: Pinecrest, North of Yosemite, gregarious in soil under conifer, 22 October 1967, H.D. Thiers (HDT 21303).

            Comments:  There are a few medium-sized puffballs in California that can be collected together and misidentified, if not carefully considered. Lycoperdon molle can be distinguished by the granular and spinose exoperidium that does not leave scars on the endoperidium. Under light microscopy, locating the free-floating sterigmata remnants sets this puffball apart from other look-alikes. Lycoperdon umbrinum can resemble Lycoperdon molle at mid-maturity, especially when there is heavy rain and the exoperidium has been wearing away. Dissimilar to Lycoperdon molle, Lycoperdon umbrinum has an exoperidium with darker greenish brown hues, smaller spines that are less dense in arrangement, and a shiny metallic tint to the endoperidium upon reaching maturity. Microscopically, L. umbrinum does not have free-floating sterigmata in wet mounts, and verrucose spores that are smaller than what is found in L. umbrinum (4.0-5.6 µm). Lycoperdon nigrescens is another medium-sized puffball that often becomes mixed up with collections of L. molle and L. umbrinum. The spinose exoperidium of L. nigrescens will leave scars on the endoperidium when they slough off through maturation, a feature parallel to Lycoperdon perlatum. This dark black puffball does not have the shiny metallic tint of L. umbrinum, and does not have the combined granular and spinose furfuraceous exoperidium characters found in L. molle. Microscopically, L. nigrescens has free-floating sterigmata remnants in wet mounts, paracapillitium, small pedicellate spores with verruculose ornamentation. The presence of abundant paracapillitium and the scars on the exoperidium help to separate L. nigrescens from L. umbrinum and L. molle. The results of the ITS ML show that Lycoperdon molle is within the Lycoperdon clade, but distant from each other, and neither have strong support. This suggests that multi loci gene analysis will be necessary to further clarify the taxonomic position of Lycoperdon molle within the Lycoperdaceae.

 

Lycoperdon nigrescens Pers., Neues Mag. Bot. 1:87. 1794.                                                                                                                                                                        (fig. 29, 38, 41, 69)           

Reported synonyms

             = Lycoperdon album var. nigrescens (Pers.) Pers., Syn. meth. fung. (Göttingen) 1: 146. 1801.

            = Lycoperdon perlatum ß nigrescens Pers., Syn. meth. fung. (Göttingen) 1:146. 1801.

            = Lycoperdon foetidum Bonord., Handb. Allgem. mykol. (Stuttgart): 253. 1851.

            = Lycoperdon peckii J.B. Morgan

            = Lycoperdon perlatum var. peckii (Morgan) Bowerman, Can. J. Bot. 39(2): 375. 1961.

            = Lycoperdon foetidum var. peckii (J.B. Morgan) Demoulin, Beih. Sydowia 8: 150. 1979.

            Type: There is a collection labeled ‘typus’ at The Botanical Conservatory and Garden at Geneva; SIB identifier 299167/1, Barcode number G00126979, located in herbarium G. The location of this collection is from Herford, Nordrein-Westfalen, Germany, stored under the name Lycoperdon foetidum Bonard., but validated as Lycoperdon nigrescens Pers.. The collector name and date are unknown.

            Gasterocarp (15) 30-43 mm tall x 34-60 mm broad; obpyriform to subglobose; rhizomorph up to 10-20 mm long, rooting the fruitbody to the soil, with fine hyphal strands entangled with soil and vegetal debris; ostiole formed by a stellate or lateral tear, up to 5-25 mm broad, persistent, frayed edges without an upturned margin. Exoperidium white to cream buff when young (5A1), turning orange white (5A2), brown to dark yellowish brown (5F6-5F5), becoming dark brown (6F4) with maturity; conical spinulose and granular, having spines with convergent tips, spines falling away to leave a reticulate scar pattern on the endoperidium; spines persistent if the fruitbody dries up to quickly, in this case the spines may not leave a scarred appearance which may lead to misidentification. Endoperidium light yellow brown (5D4-5D5); parchment-like, thin, persistent. Gleba Isabella color (5E6) to light yellowish brown at maturity (5D5); cottony, flexuous, remaining cottony when mature. Subgleba cream when young, tan brownish (5C4) with age; pseudostipe reduced, 15 mm tall x 4 mm wide. Diaphragm absent. Macromorphology derived from dried herbarium specimens.

            Basidiospores globose; 3.2-4.8 X 3.2-4.8 µm [xmr = 4.2 X 4.3 µm, Q = 0.8-1.0, Qmr = 0.97, n = 35, s = 1]; yellowish brown to dark brown amber in KOH mounts; verruculose spores with small bumpy truncate warts in light microscope, short verruculose columns with a flattened to cone-shaped apex seen under SEM; oil drop absent; spores thick-walled; very short remnant pedicel present, < 0.8 µm; free-floating sterigmata present in wet mounts, not abundant, seen in SEM images; spores of equal size under light microscopy. Eucapillitium Lycoperdon-type; threads 1.6-6.5 µm broad, walls up to 0.8 µm thick; yellow brown in wet mounts; elastic; incrusted with cellular debris, dichotomous branching present and not abundant, knob-like projections present, tips attenuate to a fine pointed terminus, thinner sections with a wavy sinuous appearance toward the tips. Pores abundant, small and round. Septa scarce to absent. Paracapillitium abundant, with septate, thin walls. Exoperidium textura globulosa; composed of inflated, globose, sphaerocyst cells. Endoperidium texture intricata; composed of intertwined, septate, hyphal threads.

            Habitat: A rare species, mostly confined to the North coast of California. Most often growing solitarily on litter in damp dark redwood and fir forests. Collected growing under alder and maple on the East Coast of the United States. Among the Geographic Subdivision of California, this species is only found in Northwestern California.

            Distribution: Known only from United States, and previously reported from Alaska, Arizona, California, Colorado, Idaho, Michigan, New Jersey, Oregon, Pennsylvania, Tennessee, Washington, West Virginia, and Wisconsin. Previously reported from Austria, Belgium, British

Isles, Czech Republic, Denmark, England, France, Germany, Holland, Mexico, Portugal, Spain, Sweden.

            Material Examined: CALIFORNIA, Del Norte Co.: Highway 199 near the Oregon boarder, solitary in humus in mixed woods, 29 October 1971, H.D. Thiers (HDT 28480); Highway 199 near the Oregon boarder, solitary in humus in mixed woods, 29 October 1971, H.D. Thiers (HDT 28480). Humboldt Co.: Trinidad, Spruce Grove, on old wood, March 1946, coll. unknown (UCB64713)(UC); Old Highway 101 to Trinidad, 15 October 1968, D. Toll (DT 3666-2145)(HSU); Old Highway 101 to Trinidad, 15 October 1968, D. Toll (DT 3657-2144)(HSU); Trinidad, Patrick’s Point State park, gregarious in soil under conifers, 16 October 1971, H.D. Thiers (HDT 28364); Picea sitchensis (Sitka spruce) forest, 13 October 1975, D. Largent (DL 7866)(HSU); Big lagoon spruce forest, 15 October 1977, D. Largent (DL 7461-5207)(HSU); Humboldt area, 17 November 1977, D. Largent (DL 7754-6195)(HSU); LC Dunes, 1979-1981, S. Sweet (259-1167447)(HSU); Patrick’s Point State Park, 6 November 1986, D. Deshazer (VW 104-10180)(HSU); Humboldt area, 26 November 1988, coll. unknown (SM 645Stout)(HSU); Patrick’s Point State Park, date unknown, H.D. Thiers (HDT 22013). Mendocino Co.: Eel River, lower Walker Meadow, 2 November 1975, S. Sweet (Sweet 69)(HSU). IDAHO, Bonner Co.: Kaniksu National Forest, Binarch Creek Road, solitary in duff of mixed conifer forest, 6 September 1966, W.J. Sundberg (Sundberg 704). WASHINGTON, Clallam Co.: Olympic National Park, Boulder Creek drainage, 10 October 1991, K. Shanks (KMS 139). Lewis Co.: Cispus Environmental Center, gregarious to clustered in soil in mixed woods, 10 October 1986, H. Saylor (HS 3386).

            Comments: Lycoperdon nigrescens is very similar to Lycoperdon perlatum, but can be distinguished by the dark pigmented conical spines covering the exoperidium. When the spines slough off, they leave scar markings on the endoperidium just as exhibited on Lycoperdon perlatum. The spines on both species can be persistent if the fruitbody dries up too quickly. In this case the spines may not leave a scar appearance and may lead to misidentification. However the exoperidium of Lycoperdon perlatum remains in the cream to brown color tones through maturation, whereas Lycoperdon nigrescens becomes dark brown to almost black. Lycoperdon nigrescens Pers. should not be confused with Lycoperdon nigrescens (Pers.) Vittad., Monogr. Lycoperd. 2: 176 (1843) [nom. illegit.], which is currently named Bovista nigrescens Pers., Neues Mag. Bot. 1: 86 (1794). Bovista nigrescens Pers. is a completely different puffball, which could be confused with Bovista pila. Bovista nigrescens Pers. has a glabrous dark metallic endoperidium when mature, lacking a sterile base, and spores become dark purple brown. Bovista nigrescens is a European species, and has not been reported from California, whereas Lycoperdon nigrescens is a puffball with a large sterile base and is found along the coastal forests of Northern California. The results in ITS analysis shows strong support of a Lycoperdon nigrescens clade nested within Lycoperdon with 100% bootstrap and 100% PP.

 

Lycoperdon perlatum Pers., Observ. Mycol. (Lipsiae) 1: 4. 1796.                                                                                                                                                                        (fig. 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.

            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.

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.

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).

            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 pratense Pers., Tent. disp. meth. fung. (Lipsiae): 7. 1797.

                                                                                                                        (fig. 31, 71)

Reported synonyms

            = Lycoperdon hiemale Bull., Herb. Fr. 2: 148. 1782. 1781-1782.


= Lycoperdon depressum Bonord., Bot. Ztg. 15: 611. 1857.


= Utraria pratensis (Pers.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 25: 368. 1873.


= Lycoperdon subpratense Lloyd, Mycol. Writ. 2(Letter 20): 231. 1905.


= Vascellum depressum (Bonord.) F. Šmarda, Bull. int. Acad. pol. Sci. Lett. 1: 305.             1958.

= Calvatia subpratensis (Lloyd) Coker & Zeller, Mycologia 39(3): 305. 1947.


            = Vascellum pratense (Pers.) Kreisel, Feddes Repert. 64: 159. 1962.

= Vascellum pratense subsp. subpratense (Lloyd) Kreisel, Feddes Repert. Spec. Nov.             Regni Veg. 68: 87. 1963.


= Calvatia depressa (Bonord.) Zeller & A.H. Sm., Lloydia 27: 171. 1964.


= Vascellum subpratense (Lloyd) P. Ponce de León, Fieldiana, Bot. 32(9): 113. 1970.

Type: According to Kreisel (1962), the type locality for Lycoperdon pratense is Europe. A holotype specimen was not designated in the protologue and nor a lectotype or neotype have been designated.

Gasterocarp 20-40 mm tall x 22-45 mm broad; obpyriform; rhizomorph composed of tightly woven hyphae, attached to duff and soil debris, persistent; ostiole developing from a lateral to circular tear in the peridium layers, typically circular or irregular-shaped, sometimes torn horizontally down the side of the fruitbody, fibers along the torn edges radiating inwards, developing outward until a large vase-shaped opening exists. Exoperidium white cream buff (4A3) turning tones of tan yellow brown (5C5, 5E5); heavily ornamented, covered in small hexagonal scales with granules and spines radiating outward making it minutely furfuraceous, many spines fusing at the tips to form larger spines, small spines persistent in peridial cracks and folds, larger spines forming at the apex, smaller spines forming on the sides and base, sloughing off and not leaving scars on the endoperidium. Endoperidium white when young, turning brownish orange to yellowish brown beige (5C4, 5E5); smooth, parchment-like, persistent. Gleba white and firm when young, turning dark olive brown (4F7), to dark brown (6F4); mixed-type, becoming powdery, dehiscing and leaving a vase-shape fruitbody that persists in the environment, detaching from the substrate and tumbling around, disseminating spores up to a year. Subgleba white when young, turning dark brown (6F4), to dark yellowish brown (5F5); chambered and compact, 10-15 mm tall, taking up to 1/2 of the fruitbody. Diaphragm present, clearly defined and distinct.

            Basidiospores globose to subglobose; 3.5-4.5 X 4.0-4.9 µm [xmr = 4.1-4.2 X 4.1-4.4 µm, xmm = 4.2 ± 0.1 X 4.3 ± 0.2 µm, Q = 0.8-1.1, Qmr = 0.9-1.0, Qmm = 1.0 ± 0.1, n = 25, s =2]; amber brown in wet mounts, turning dark turquoise blue in lacto-phenol cotton blue staining reaction; smooth to roughened with asperate warts in wet mounts and SEM; central oil drop present; spores thick-walled; pedicel up to 1 μm long; free-floating sterigmata present in wet mounts from mid-mature fruitbodies, not abundant; spores of equal size in wet mounts. Eucapillitium Lycoperdon-type; threads 3-7 μm broad, walls < 0.5 μm thick; light amber brown in water mounts; elastic; found connected to and growing inward from the endoperidium walls, not growing inward from the inner diaphragm walls, scarce to absent and difficult to find under light microscopy in mature fruitbodies, hyphal threads incrusted with cellular debris, dichotomous branching absent or unnoticed, knob-like projections absent, straight, tips with a round terminus. Pores absent. Septa absent. Paracapillitium abundant throughout, dominant form of capillitial threads; thin-walled, hyaline, septate and disarticulating at the septum wall, dichotomous branching scant but present, almost entire gleba composed of paracapillitium, tips with a round terminus. Subgleba composed of long, tightly woven, dichotomously branched hyphal threads; septa scarce and present; tissue readily takes in lacto-phenol cotton blue stain, turns green to dark blue with stain, stain almost takes up the entire hyphal thread. Diaphragm composed of tightly woven hyphal threads. Exoperidium textura globulosa; covered in orbicular sphaerocysts, staining purple in methylene blue chloride (MBC), cells linked in chains, occasionally pigmented. Endoperidium textura intricata; composed of tightly woven hyphal threads, stain slightly green in MBC, other non-distinguishable cellular material stains purple with the MBC stain.

            Habitat: Growing in caespitose to gregarious clusters, in fairy rings, or growing solitary. Collected on soil in pastures, in prairies, in grass-specked sand dunes, and in meadows. Found in meadows around pine and conifer forests along the coastal ranges. Considered a common lawn species, but not commonly collected in California. Lycoperdon pratense seems to prefer area where morning fog banks and morning dew are common. Among the Geographic Subdivisions of California, this species can be found in Northwestern California, and in Central Western California.

Distribution: Known from many parts of the United States, and previously reported from California, Colorado, Idaho, Illinois, Indiana, Nebraska, Oregon, Tennessee, Texas, Utah, Washington, West Virginia. Also known from Denmark, France, Germany, Italy, Mexico, Scotland, and Sweden.

Material Examined: CALIFORNIA, Alameda Co.: Oakland, Knowland Park, gregarious in rings two to ten feet in diameter, 30 November 2011, S.S. Jarvis (SSJ 419). Sonoma Co.: Petaluma, Pleasant Hill Road, terrestrial in grass under conifer, 7 July 2009, D. Deshazer (SSJ 337). Trinity Co.: South Fork Mountain, 16 July 1933, coll. unknown (UCB1162669)(UC).

            Comments: The diaphragm-forming Lycoperdon puffballs have similar features and it may be difficult to tell them apart. A good microscope with oil emersion capability at high power is necessary for identifying eucapillitium versus paracapillitium from various parts of the gleba. Lycoperdon pratense and Lycoperdon lloydianum are very similar in outward appearance, which can cause confusion and misidentification among them. Lycoperdon pratense has non-poroid eucapillitium, which is scarce to absent. Ponce de Leon (1970) argues that the eucapillitium in Lycoperdon pratense is absent, however it can be found along the periphery of the endoperidium wall in collections from California. Staining with LpCB or methylene blue chloride is helpful in this process, as well as having young to mid-mature fruitbodies that are fresh. Lycoperdon pratense has a wide and very well developed diaphragm that can be seen earlier in development than that of other diaphragm-forming puffballs. One main distinction that sets Lycoperdon pratense apart from Lycoperdon lloydianum is the unique feature of eucapillitium growing out from the inner endoperidium walls into the gleba, whereas the eucapillitium grows strictly out from the inner diaphragm wall in L. lloydianum. Both of these grassland species have been collected together in the same habitat. However, Lycoperdon pratense is a rare species in California. Lycoperdon curtisii is smaller (under 20 mm broad), has Lycoperdon-type eucapillitium that can be found along the periphery of the inner endoperidium wall, a reduced subgleba, and smaller echinulate spores. The type for the genus Vascellum, V. depressum (Bonord.) F. Šmarda, has been accepted as a synonym of Lycoperdon pratense Pers. (Larsson 2008), and this is supported with the ITS analysis. The ML tree produced in this analysis shows Vascellum species (now in Lycoperdon) as a monophyletic clade within the Lycoperdon clade, with 92% bootstrap and 100% PP. Despite the similarities in outward appearance of Lycoperdon pratense and Lycoperdon lloydianum, the ITS sequence data suggests that these are two separate species with strong bootstrap and PP support within the Vascellum clade of Lycoperdon.

           

Lycoperdon pyriforme Schaeff., Fung. Bavar. Palat. 4: 128. 1774.                                                                                                                                                                        (fig. 32, 72)

Reported synonyms

= Lycoperdon pyriforme ß tessellatum Pers., Syn. meth. fung. (Göttingen) 1: 148. 1801.

= Utraria pyriformis (Schaeff.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 25: 369. 1873.

= Morganella pyriformis (Schaeff.) Kreisel & D. Krüger [as 'pyriforme'], in Krüger & Kreisel, Mycotaxon 86: 175. 2003.

Type: According to Krüger (2003), the iconotype is a plate from J.C. Schaeffer, Fungorum Bavar II: 185m 1763 (Thurn und Taxis library in Regensburg). Krüger (2003) created an epitype: Kelheim county, Niederbayern district, Bavaria, Germany, near Regensburg, collected from under Picea on wood, collected on 18 October 1991, (GenBank sequence AJ237620) (Linzenkirchner collection Os97/91, REG1810910s). The selection was based on the type locality and morphology similar to Schaeffer’s plates. There are two type collections listed on the National Herbarium Nederland website for which the location, date, and collector are unknown (catalog number L 733595; accession number 910258341; barcode numbers L 0054559, and L 0054571).

            Gasterocarp 20-55 mm tall x 11-35 mm broad, 4-8 mm broad at the base; obpyriforme  to elongated obpyriforme, with and without a bulbous base; rhizomorphs abundant, white, persistent, elastic, branching, creating a network of thick hyphal strands attaching individuals together in large gregarious clusters, sometimes up to several hundred fruitbodies gregariously growing together; ostiole umbonate, slow to form at the apex via a thinned out worn away patch from the expansion of growth, typically torn laterally to expose the spores, torn edges uplifted and frayed. Exoperidium white when young, becoming clay color, light yellow brown, Isabella color to dark yellowish brown (5C5, 5D5-6, 5E6-7, 5F5), when dry the base ranges from light yellow to grayish yellow (4A4-3, 4B4), with patches turning reddish brown in age (8F5), overall non-uniform color, upper portions of fruitbody darker than the base, darker where hit by sunlight; exoperidium ornamentation furfuraceous-spinulose to scurfy granulose, covered in verrucose spines and granules, verrucae spines becoming appressed with age, spines persistent, heavily ornamented at the base and in between folds, sloughing off in patches to reveal the endoperidium. Endoperidium yellowish brown to light yellowish brown (4B5, 5D5), turning metallic golden yellow bronze brown (5E5) with age; endoperidium ornamentation slightly granular when young, becoming smooth with age and weathering, parchment-like, persistent. Gleba white at first, turning cream to pale yellow (4A3, 4B3), to shades of pale yellow grey (4C4, 4D4), maturing from the center outward, becoming dark yellowish brown at maturity (5F5-7); firm and chambered when young, cottony and persistent with maturation. Subgleba stipitate, a permanent sterile base, cream white when young (4A2), yellow peach tones (3A3-4), or turning pale pink (9A2) in some specimens; cellular to compact chambered, taking up the lower 1/3 to 3/4 of entire fruitbody. Diaphragm absent.

            Basidiospores globose; 2.4-3.2 X 2.4-4 µm [xmr = 3.0-3.11 X 3.1 µm, xmm = 3.0 ± 0.1 X 3.1 ± 0.0 µm, Q = 0.8-1.0, Qmr = 0.9-1.0, Qmm = 0.9 ± 0.0, n = 25, s = 2]; spores yellow golden in water mounts, staining olive green in KOH; spores roughened to smooth with a light microscope, under SEM spores asperate with rounded warts connected by ridges; oil drop present; spores thin-walled, deflating easily with desiccation; pedicel very rudimentary to absent, seen on a few spores with SEM images; free-floating sterigmata remnants absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Lycoperdon-type; up to 4 μm broad, walls 1-2 μm thick; hyaline to slight golden in wet mounts; subelastic; glabrous, dichotomously branched, branches rare, mostly straight at then main stem of capillitium, subundulate to sinuous towards the tips along thinner walls; tips attenuate to a pointed terminus, not attenuate. Pores absent. Septa abundant, disarticulating at the septum wall, with clean breaks at septa junctions or torn where broken. Paracapillitium present, abundant, septate, not pored. Exoperidium textura angularis; composed of large and inflated sphaerocyst cells, globose to hexagonal in shape, tightly compact, with walls up to 1-2 μm thick, slightly pigmented in water mounts; thick-walled finger-like cystidia protruding from the layer of globose cells, spinose to fusoid or lanceolate-shaped, cystidia pointed and with somewhat jagged edges. Endoperidium textura globulosa; composed of spherical, flat, intertwined, thick-walled, longitudinally aligned, pigmented cells. Subgleba chambered, composed of long intertwined hyphal strands hyaline, thick-walled, flexible, aseptate, non-poroid, sinuous, with knob-like projections, and rounded tips.

            Habitat: Lycoperdon pyriforme is one of the few Lycoperdaceae that is known to grow strictly lignicolous, and is a common white rot fungus in Central Western California. Most often it is collected from woody debris, very decayed wood, or buried wood of hardwood and conifer tree species. It can occasionally be found growing in soil, or seemingly without the presence of and wood debris. Typically this puffball grows in regions where no other lignicolous Lycoperdales occur (Krüger 2001). Most often Lycoperdon pyriforme is found in large gregarious clusters, with up to several hundred fruitbodies attached to one another with thick elastic rhizomorphs. It has also been noted to grow as a solitary fruitbody on occasion. The growing habit depends on environmental conditions that favor prolific growth for large clusters to form, rain and moisture being the key factor. Lycoperdon pyriforme is found in the subarctic, boreal, and temperate zones of Eurasia and North America. Surprisingly, it has been reported from high elevations of the Swiss Alps and the Himalayas of Nepal, and also occurring in subtropical zones as well. Among the Geographic Subdivisions of California, this species can be found in Northwestern California, in the Cascade Range, in Central Western California, and parts of the Sierra Nevada.

Distribution: Known from many parts of the United States, and previously reported from Alabama, Alaska, Arizona, California, Colorado, Connecticut, Delaware, Florida, Georgia, Idaho, Illinois, Iowa, Indiana, Kansas, Kentucky, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missouri, Montana, Nebraska, New Hampshire, New Mexico, New York, North Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, South Dakota, Tennessee, Texas, Utah, Vermont, Virginia, Washington, West Virginia, Wisconsin, and Wyoming. Also known from Australia, Austria, Belgium, Brazil, Canada, China, Colombia, Congo, Costa Rica, Cuba, Czech Republic, Ecuador, England, Finland, France, Germany, Hungary, India, Mexico, Nepal, Papua New Guinea, Philippines, Poland, Mongolia, Netherlands, Nepal, New Zealand, Norway, Russia, Slovakia, Spain, Sweden, Switzerland, and Uruguay.

            Material Examined: CALIFORNIA, Alameda Co.: Berkeley, behind the football stadium, hillside on a rotting oak log, gregarious to caespitose, 28 January 2009, S.S. Jarvis (SSJ 281); Berkeley, UC Berkeley Botanical Gardens, hillside on rotting oak wood, 5 December 2009, S.S. Jarvis (SSJ 351). Amador Co.: Silver Lake, caespitose on buried rotten log, 5 October 1975, coll. unknown (Halling 876); Silver Lake, caespitose on rotten conifer log, 5 October 1975, H.D. Thiers (HDT 35042). Del Norte Co.: Jedediah Smith Redwoods State Park, gregarious under pine and redwood, 9 October 1966, coll. unknown (Ammirati 415); Jedediah Smith Redwoods State Park, gregarious in duff under sequoia, 3 November 1966, H.D. Thiers (HDT 17820); Jedediah Smith Redwoods State Park, gregarious in soil under conifers, 30 December 1966, H.D. Thiers (HDT 18316). Del Norte Co.: Highway 199 at the Oregon boarder, gregarious on wood under redwood, 30 October 1971, H.D. Thiers (HDT 28508). El Dorado Co.: Crystal Basin Recreation Area, Icehouse Lake, caespitose on rotten logs, 16 October 1971, H.D. Thiers (HDT 36711); Crystal Basin Recreation Area, Icehouse Lake, solitary on rotten conifer log, 17 October 1976, H.D. Thiers (HDT 36714); Crystal Basin Recreation Area, Icehouse Lake, caespitose on dead conifer wood, 16 October 1976, H.D. Thiers (HDT 36717). Humboldt Co.: Richardson Grove State Park, gregarious in soil in mixed woods, 16 November 1965, H.D. Thiers (HDT 14209); Sloan Creek, growing on rotten conifer, 27 September 1967, coll. unknown (2777-2135)(HSU); Shores Acres Park, along coast trail, gregarious on decaying log at sea level, 9 November 1979, C. Ardrey (W327-7122)(HSU); Willow Creek, North of Highway 299, gregarious in mixed oak and fir forest, 8 November 1987, R.S. LaChance (RSL 12-10584)(HSU); Coastal, on decaying oak, 30 November 1987, R.S. LaChance (RSL 47-10606)(HSU). Marin Co.: Muir Woods National Monument, Bohemian Grove Comfort Station, growing from soil through redwood needles under redwood, 4 December 1961, coll. unknown (Brown 554); Lily Lake, gregarious in soil under oak and bay trees, 22 October 1972, M. Concarnon (HDT 141); Audubon Canyon Ranch, Volunteer Canyon Ranch, on soil near decaying wood, 20 November 1981, C. Calhoun (Calhoun 81-2625b). Mendocino Co.: Jackson State Forest, caespitose beneath fir along roadside, 12 November 1961, D. Largent (Largent 60); Mendocino, Jackson State Forest, on mulch under redwood, 12 November 1984, coll. unknown (Youst 139); Gualala, Fish Rock Road, Pacific Lumber Company logging roads, on a rotting log buried in a river bank, 09 December 2007, S.S. Jarvis (SSJ 231); Gualala, Fish Rock Road, on a rotting log buried in a river bank, 30 November 2008, S.S. Jarvis (SSJ 270); Gualala, stump beach, along path near the parking lot, December 2009, S.S. Jarvis (SSJ 431); Gualala, Fish Rock Road, Pacific Lumber Company logging roads, on soil and rotting wood chips in between planks of a bridge, 15 December 2009, S.S. Jarvis (SSJ

294); Gualala, Fish Rock Road, Pacific Lumber Company logging roads, buried wood along a stream bank, gregarious and numerous, 15 January 2010, S.S. Jarvis (SSJ 360); Fort Bragg, Jackson State Forest, along overgrown path among grass and rotten wood chips, 1 February 2011, S.S. Jarvis (SSJ 425). Santa Cruz Co.: Santa Cruz Mountains, near Boulder Creek, solitary in humus in dense mixed woods, 22 November 1963, H.D. Thiers (HDT 10779). Siskiyou Co.: Klamath National Forest, Marble Wilderness area along Haypier trail, halfway to the first meadow, D. Largent (DL 7544-6204)(HSU). Sonoma Co.: YMCA Camp, Skaggs Springs Road, scattered in duff along an overgrown abandoned road, 21 November 2008, S.S. Jarvis (SSJ 261); YMCA Camp Skaggs Springs Road, gregarious on a rotting log, 21 November 2008, S.S. Jarvis (SSJ 266); Roy’s Redwoods, growing in thick humus, 13 January 2010, S.S. Jarvis (SSJ 372). Tehama Co.: Mineral, on soil under Pinus contorta (lodgepole pine), 30 September 1975, coll. unknown (Showers 2900); Mineral, along Highway 36, gregarious to caespitose on rotten fir log, 10 October 1976, H.D. Thiers (HDT 36622). Tuolumne Co.: Highway 108, 2 miles East of Long Barn, 1,219 meters (4,000 feet) elev., gregarious on soil under conifer, H.D. Thiers (HDT 46988).

Comments:  Growing almost strictly on wood, collecting Lycoperdon pyriforme from the soil without traces of decomposing wood can make identification efforts confusing. There are a few other puffballs that have been found growing on wood in California: Calvatia sculpta, Lycoperdon perlatum, and Lycoperdon nigrescens. However, a lignicolous growing habit is not common for puffball species (Alfredo 2014). The use of a good compound light microscope and locating the finger-like cystidia-type cells projecting from the exoperidium tissue on Lycoperdon pyriforme is a key to proper identification.

            In 2001, Krüger revealed a distinction between lignicolous Lycoperdaceae species based on the DNA analysis results, and discussed how Lycoperdon pyriforme does not cluster with other Lycoperdon species. The results showed that 9 sequences of Lycoperdon pyriforme fell outside the Lycoperdon clade, but within the Lepiota outgroup. In 2003 Krüger transferred Lycoperdon pyriforme into Morganella based primarily on its lignicolous growing habit. Bates' (2004) ITS sequencing results also showed one sequence of Morganella pyriforme as outside of the Lycoperdon clade, sister to one sequence of Disciseda candida and distinct from two other Morganella species. ITS sequence analysis of collections from California show three sequences of Lycoperdon pyriforme forming its own clade with 100% bootstrap and 100% PP support, sister to a supported Disciseda clade containing several sequences of D. candida, D. cervina and D. johnstonii. Larsson and Jeppson (2008) showed that Morganella pyriforme was sister to the Lycoperdon clade with 70% bootstrap and 93% PP support. Based on these results, they recognized Morganella pyriformis as a species of Lycoperdon. None of these studies included the type species of Morganella, M. mexicana Zeller, so the taxonomic status of Morganella remains questionable. However, Lycoperdon pyriforme being sister to Disciseda, and basal to Bovista and Lycoperdon clades, warrants additional genes to be sequenced to further clarify the taxonomic position of it amongst the other Lycoperdaceae.

 

Lycoperdon subcretaceum (Zeller) Jeppson & E. Larss., Agarica 29: 90. 2010.                                                                                                                                                (fig. 33, 42, 73)

Basionym º Calvatia subcretacea Zeller, Mycologia 39(3): 298. 1947.

Reported synonyms

=  Gastropila subcretacea (Zeller) P. Ponce de León, Phytologia 33(7): 461. 1976.

= Handkea subcretacea (Zeller) Kreisel, Nova Hedwigia 48(3-4): 288. 1989.

            Type: A holotype is housed at the New York Botanical Gardens (NYBG65669). This material is from Mt. Hood in Hood River County, above Cloud Cap Inn, Oregon, collected by J.R. Keinholtz, determined by S.M. Zeller, July 1936. The collection is labeled as Calvatia subcretacea.

            Gasterocarp 14-62 mm tall x 15-75 mm broad; globose to depressed globose, ovate or pulvinate, sometimes flabelliform and irregular in shape, most often more broad than tall, base sharply tapering to a point; gasterocarp sometimes sub-hypogeous or growing under duff with only the top most apical portions exposed; rhizomorph compose of a single thick short strand of mycelium up to 10-90 mm long x 2-5 mm broad, incrusted with soil, protruding from the center of the gasterocarp base or off center, thin threads of mycelium present when absent of a thick rhizomorph, typically tough and persistent, remaining attached to the soil throughout maturity; ostiole developing through lateral to stellate cracks at the apex, slow to develop with maturation, swelling and desiccation. Exoperidium white at first, turning dull white to beige buff and brownish orange (5A3, 5C4), becoming tan to dark yellowish brown (5E6, 5F5) with maturity and sun exposure; up to 4 mm thick, thicker at the top and sides, thinning towards the base; furfuraceous to cottony in appearance under dissecting scope when very young, becoming entirely covered in short blunt to pointed warts, sometimes to smooth or roughened at the base; warts grey yellow brown (5D3) to dark brown or black (5F4-5, 5F1) at the apex, darker with sun exposure; warts up to 7 mm broad x 0.5-1 mm tall; warts forming with gasterocarp growth by way of cracking and separation of cracks as the gasterocarp swells in size, warts hexagonal to polygonal in shape, apical warts larger and sometimes pyramidal-shaped when young and often merging at the pyramidal points, scaling down in size towards the base, becoming flattened with maturation, larger warts marked by vertical lines that converge at the top of the wart, persistent and remaining attached to the endoperidium mostly throughout maturation, will eventually wear away in very old specimens, warts and bits of exoperidium and endoperidium eventually sloughing off together to expose the gleba. Endoperidium white when young, then bright yellow hues (4A5-4B5) in between exoperidium cracks, then a dull yellow brown Isabella color (5E5-6); up to 1mm thick, waxy and brittle when young and moist, glabrous and persistent. Gleba white when young to cream yellow to grayish yellow (4A2-3, 4B5, 4C7) when young, then turning olive (4E5), then dark chocolate brown (6F4-7) when spores mature; firm and dense when young, maturing from the center outwards, easily separating from the endoperidium when young, viscid when fresh with yellow liquid at mid-maturity, becoming crust-like when mature and dry to powdery, gleba becoming completely pulverulent, staining green in KOH. Subgleba absent. Diaphragm absent.

            Basidiospores globose to subglobose; 4-5.6 X 4-5.6 µm [xmr = 5.0-5.1 X 4.6-4.9 µm, xmm = 5.0 ± 0.1 X 4.8 ± 0.2 µm, Q = 0.9-1.2, Qmr = 1.0-1.1, Qmm = 1.0 ± 0.0, n = 25, s = 3]; golden yellow in wet mounts, golden amber brown in KOH mounts, hyaline envelope on spores in KOH mounts; under light microscope roughened when immature and echinate when mature, under SEM spores covered in granulose coating with pyramidal to conical echinate protuberances with a rounded terminus on the echinae; oil drop present; spores thick-walled; pedicel short to medium length, up to 0.8-3.2 (7) µm, with clean broken or pinched tails seen in SEM; free-floating sterigmata not present in wet mounts; spores of equal size in wet mounts. Eucapillitium Calvatia-type; threads up to 5.5-7.2 (16) µm broad, walls up to 0.5-1.6 µm thick, uneven inner hyphal walls; brown yellow in water mounts, yellowish green in KOH mounts; non-elastic and fragile; incrusted with cellular debris, dichotomous branching present but scarce, some capillitium lengths irregularly swollen, knob-like projections present, striate lateral stripes along the eucapillitium walls seen in SEM, mostly straight, sinuous only at the tips along thin threads; ends tapering to a round terminus. Pores round, medium to large in size, abundant; with abundant slit-like pits, these can be wavy, straight, or stellate in appearance, under SEM the slit-like pits resemble jagged cracks or torn capillitial threads due to the fragile nature of the hyphae. Septa rare, disarticulating clean at the septum wall, or fracturing at random locations along the hyphal wall, roughened broken hyphae and clean disarticulation at the septum seen in SEM. Paracapillitium absent. Center of mature gleba often shredded with small fragments of hyphal threads, longer threads located along the inner endoperidium wall. Exoperidium textura globulosa; composed of chains of oblong, hyaline, sphaerocyst cells. Endoperidium a combination of textura intricata and textura globulosa; composed of a network of intertwined hyphal threads and thin-walled globose sphaerocyst cells, swollen when young and fresh, collapsed when aged or dry.

Habitat: Usually growing on dry conifer duff above 1,524 meters (5,000 feet) elevation in small gregarious groups, or solitarily. Most often found with damp soil located around the rhizomorph, even in very dry soil conditions. Fruitbody often found partially submerged or subhypogeous in the duff. Growing in rocky soil and in association with pine, fir, and other conifers. Most often collected along the drip zone of nearby trees. Among the Geographic Subdivisions of California, this species is found in the Cascade Range, in the Modoc Plateau, in the Sierra Nevada, and in the Eastern Sierra Nevada.

Distribution: Known only from the Western United States, and previously reported from Arizona, California, Colorado, Idaho, Oregon, Washington, and Wyoming. Recently reported from Central Norway.

            Material Examined: CALIFORNIA, Alpine Co.: Alpine Lake, shore of the lake, in coniferous diff, 2,225 meters (7,300 feet) elev., 1 August 1952, L. Bonar (UCB960454)(UC); Alpine Lake, scattered in humus under mixed conifers, June 1967, coll. unknown (Halling 1411); Ebbitts Pass, solitary in soil under conifers, 21 August 1974, H.D. Thiers (HDT 32800); Alpine Lake, 2,286 meters (7,500 feet) elev., scattered and common under fir and pine, August 1983, H.D. Thiers (HDT 46037). Amador Co.: Highway 88, Silver Lake campground, scattered in soil under conifers, July 1967, H.D. Thiers (HDT 20145); Silver Lake, Highway 88, buried in humus under conifers, July 1967, H.D. Thiers (HDT 20057); Silver Lake, buried in soil under conifers, July 1967, H.D. Thiers (HDT 21123); Silver Lake, gregarious in humus under conifers, 28 June 1973, H.D. Thiers (HDT 31063); Silver Lake campground, Highway 88, scattered in soil under conifers, July 1982, H.D. Thiers (HDT 44650); Silver Lake vicinity, Highway 88, 2,438 meters (8,000 feet) elev., in soil under fir and pine, July 1983, H.D. Theirs (HDT 45964); Highway 88, East of Silver Lake, buried in humus under conifers, June 1985, H.D. Thiers (HDT 49069). Bute Co.: Jonesville, terrestrial, 12 May 1936, V. Mentzer (UCB554978)(UC). Calaveras Co.: Sand Flat, East of Betts Pass, on black soil, 5 June 1939, V. Miller (UCB621506)(UC). El Dorado Co.: Grass Lake, Luther Pass, buried in humus under Pinus contorta (lodgepole pine), July 1967, H.D. Thiers (HDT 20062); El Dorado, Echo Summit, 18 June 2009, R. Pastorino (SSJ 328); El Dorado, Echo Summit, 18 June 2009, R. Pastorino (SSJ 329). Fresno Co.: Kings River Canyon, Rae Lake, growing on trail, 20 July 1911, N.L. Gardner (UCB793293)(UC); Kings River Canyon, along trail from General Grant Forest to South fork of Kings River, July 1911, N.L. Gardner (UCB793298)(UC); Huntington Lake, in forest under Abies concolor (white firs) and Pinus palustris (yellow pines), 29 July 1941, coll. unknown (UCB655344)(UC); Kings Canyon National Park, Great Grave, 1,920 meters (6,300 feet) elev., in soil under white fir (Abies concolor), 30 June 1980, C. Calhoun (Calhoun 41229). Lassen Co.: Lassen Volcanic National Park, Juniper Lake, scattered in humus under conifer, 2,042 meters (6,700 feet) elev., 30 June 1965, H.D. Thiers (HDT 12885); Lassen National Park, Morgan Summit, Highway 89, June 1991, H.D. Thiers (HDT 53536). Mariposa Co.: Yosemite National Park, Crane Flat vicinity, scattered on soil under firs and pines, June 1976, coll. unknown (Halling 1396). Modoc Co.: Blue Lake, in campground, under Pinus ponderosa (ponderosa pine), June 1981, coll. unknown (Curlin 429). Placer Co.: near Cisco, under fir, 2,133 meters (7,000 feet) elev., June 1909, coll. unknown (UCB506624)(UC); Donner Lake, in forest duff under Larix laricina (tamarack larch), 1,828 meters (6,000 feet) elev., 8 May 1934, coll. unknown (UCB521308)(UC). Plumas Co.:  Bucks Lake, on slopes of South-West shore, 11 July 1942, coll. unknown (UCB671387)(UC); Gold Lake, Graeagle Highway, scattered in soil under conifers, June 1989, H.D. Thiers (HDT 52189). Shasta Co.: Lassen National Park, Hat Lake, 1,920 meters (6,300 feet) elev., gregarious in humus under aspen trees, 2 July 1965, H.D. Thiers (HDT 12926); Lassen National Park, King Creek Meadow, on litter and duff, 28 August 1974, coll. unknown (UCB1457104)(UC); Lassen Volcanic National Park, King Creek Meadows, cold boiling area, terrestrial, 30 July 1980, W.M. Cooke (UCB1473311)(UC); Dead Horse Summit, Highway 89, in soil under conifers, July 1982, H.D. Thiers (HDT 44595). Sierra Co.: Chapman Creek campground, Yuba River Canyon, on soil with Abies concolor (white fir) and Pinus lambertiana (sugar pine), 11 June 1961, D. Noack (U of Idaho Forestry Herb. 2705)(UC); Chapman Creek trail, Chapman Creek campground, gregarious in soil under conifers, June 1980, H.D. Thiers (HDT 40897); Sierra Nevada Field Campus, Highway 49, solitary in soil under conifers, May 1984, H.D. Thiers (HDT 47664); Sierra Nevada Field Campus, Highway 49, solitary in soil under conifers, May 1986, H.D. Thiers (HDT 49799); Yuba Pass, solitary on soil in mixed conifer forest, 10 June 1986, M.T. Seidl (UCB1573136)(UC); Yuba Pass, on soil under Abies spp., 9 June 1987, coll. unknown (UCB1516061)(UC); Tahoe National Forest, Haskel Peak Road, off Gold Lake Road, scattered in soil in conifer woods, June 1989, H.D. Thiers (HDT5 2173); near Bassett’s Station, Green Acres, solitary and terrestrial in mixed conifer woods, 6 June 1989, M.T. Seidl (UCB1576949)(UC); Yuba Pass, terrestrial and scattered in mixed conifer woods, 2,042 meters (6,701 feet) elev., 9 June 1989, M.T. Seidl (UCB1576955)(UC); Sierra Nevada Field Campus, Chapman Creek, Highway 49, 2 June 2008, S.S. Jarvis (SSJ 209); Yuba Pass, Chapman Creek, 2 June 2008, S Jarvis (SSJ 210); Yuba Pass, Highway 49, 2 June 2008, S.S. Jarvis (SSJ 211); Yuba Pass, Highway 49, terrestrial, 2 June 2008, S.S. Jarvis (SSJ 213); Wild Plum Road, Girl Scout Camp, 3 June 2008, S.S. Jarvis (SSJ 216); Yuba Pass, north side of the highway, 3 June 2008, S.S. Jarvis (SSJ 219); Yuba Pass Summit, terrestrial in the campground, 3 June 2008, S.S. Jarvis (SSJ 221); Green Acres, Highway 48, terrestrial, 3 June 2008, S.S. Jarvis (SSJ 222); Sierra Nevada Field Campus, terrestrial near the back kitchen door, 3 June 2008, S.S. Jarvis (SSJ 224). Siskiyou Co.: Caribou Basin, 1,981 meters (6,500 feet) elev., terrestrial, 27 July 1937, coll. unknown (UCB585263)(UC); Mt. Shasta, along the ski trail below horse camp, terrestrial, 3 July 1946, W.M. Cooke (UCB1138972)(UC); Mt. Shasta, jeep trail to Clear Creek, 3 August 1955, W.M. Cooke (UCB1139042)(UC); Between Etna and Sawyers Pass, solitary in soil under conifers, June 1980, H.D. Thiers (HDT 41211); Mt Shasta, gregarious in humus in conifer woods, July 1980, H.D. Thiers (HDT 41090); Mt Shasta, solitary in soil under conifers, July 1982, H.D. Thiers (HDT 44585); Mt. Shasta, Sand Flats, on the ground, 24 June 1989, W.M. Cooke (UCB1575837)(UC). Tehama Co.: Gurnsey Creek Campground, Highway 89, solitary in soil under fir trees, 10 October 1976, H.D. Thiers (HDT 36618); Highway 89, 20 miles south of Lassen National Park, gregarious in soil under conifers, July 1982, H.D. Thiers (HDT 44612); Gurnsey Creek campgrounds, Highway 36, June 1988, H.D. Thiers (HDT 51693); Mineral, Highway 36, Lassen Peak junction, 31 May 2008, S.S. Jarvis (SSJ 201); Mineral, Highway 36, terrestrial under pine, 31 May 2008, S.S. Jarvis (SSJ 202). Tulare Co.: Sequoia National Park, 2 June 1947, M.T. Cooke (UCB759900)(UC). Tuolumne Co.: Pinecrest campground, scattered in humus under conifers, June 1965, H.D. Thiers (HDT 12638); Yosemite National Park, Tuolumne Meadow campgrounds, gregarious in humus under Pinus contorta (lodgepole pines), June 1966, coll. unknown (DEM); Pinecrest, 1,676 meters (5,500 feet) elev., gregarious in soil under conifers, June 1980, H.D. Thiers (HDT 47658).

            Comments: In 1947 S.M. Zeller described Calvatia subcretacea as a new species based on Calvatia-type eucapillitium, lack of subgleba, and other Calvatia-like characteristics. Ponce de Leon (1976) transferred Calvatia subcretaceae to Gastropila subcretaceae following the publication of Gastropila as a new genus by Homrich & Wright (1973). Gastropila was circumscribed by having smooth spores, a three-layered peridium, and smooth, sparsely branched, not easily broken, much entangled eucapillitium (Homrich and Wright 1973) . The use of SEM imaging on California material debunks the transfer of this species to Gastropila, showing echinate spores and Calvatia-type eucapillitium. Kreisel (1989) erected the genus Handkea based on the slit-like nature of eucapillitium pits, and transferred Calvatia subcretacea into Handkea. Calvatia subcretacea has abundant slit-like pits on the capillitium threads when seen under light microscopy. With the use of SEM imaging, these slit-like pits clearly resemble cracks and breaks in the fragile capillitium walls. They do not seem to be genetically derived morphological characters. Additionally, phylogenetic results show that Handkea is a polyphyletic assemblage and that it’s members fall at different locations within the Lycoperdon clade (Bates 2004, Gube 2009, Larsson and Jeppson 2008). Jeppson and Larsson (2010) recorded Lycoperdon subcretaceum in Norway for the first time, with results showing that two different sequences of Lycoperdon subcretaceum and Handkea subcretacea from different regions of the world were identical to each other. They cluster together with 99% bootstrap support within the Lycoperdon clade, and quite distant from the Calvatia clade. Jeppson and Larsson (2010) reports fragile Lycoperdon-type eucapillitium in the material collected from Norway, and finely warted spores; whereas the material from California has Calvatia-type eucapillitium, and the exoperidium ornamentation seems much more exaggerated than that of the European collections. ITS sequence data comparing a collection from California and a sequence from Jeppson and Larsson (JN572908) study reiterates these findings. The molecular data shows Lycoperdon subcretaceum basal to the Lycoperdon clade, and shows these two sequences to be the same species with 100% bootstrap and 100% PP support, Because molecular data place this species in the Lycoperdon clade, it will be treated here as Lycoperdon subcretaceum.

 

Lycoperdon umbrinum            Pers., Syn. meth. fung.(Göttingen) 1: 147. 1801.                                                                                                                                                                        (fig. 34, 74)

Reported synonyms            

            = Lycoperdon umbrinum ß hirtum Pers..1801.

            = Lycoperdon umbrinum var. hirsutum Alb. & Schwein., Consp. fung. (Leipzig): 80. 1805.

            = Lycoperdon hirtum (Pers.) Mart., Fl. crypt. erlang. (Nürnberg): 386. 1817.

            = Lycoperdon umbrinum var. curtisiiforme Hollós,. 1904.

            = Lycoperdon hirtum var. curtisiiforme (Hollós) Sacc. & Traverso, Syll. fung. (Abellini)

19: 1152. 1910.

            = Lycoperdon hirtum var. hirtellum (Peck) Sacc. & Traverso, Syll. fung. (Abellini) 19: 1152. 1910.

            = Lycoperdon umbrinum f. fissispinum Kreisel, Feddes Repert. 64: 141. 1962.

Type: The type material is located at the Herbier de l’Universite Montpellier II (MPU), MPUB03504, the type locality is Collsacreu, Spain.

            Gasterocarp 30-52 mm tall x 11-40 mm broad; ovoid to globose when young, expanding into obpyriform with maturation and elongation of the subgleba; rhizomorph composed of white thick entangled hyphae creating a mat-like structure, clinging to vegetal debris, up to 11 mm long x 4mm broad; ostiole slow to develop, cracking and splitting to open up at the apex, developing as a circular to ovular opening, top becoming semi-flat to plane with age, ostiole with fibers radiating up and outward, sometime recurved outward. Exoperidium white cream when young (4A3), becoming yellow brown to dark yellowish to dark brown (5F5-8, 6E4, 6F7, 7F3) very early in maturation, lighter shades of pink buff to brownish orange towards the base (5A3, 5C4); entire fruitbody covered in a finely warted to minute furfuraceous-spinose and granular coating which takes up the darkest pigments, spines of various shapes, ornamentation more glabrous at the base, spines falling away around the apex, leaving a vey minute semi circular to oval or rectangular-shaped scar, spines more concentrated in some spots and larger in size where concentrated in folds and at the apex, some spines merging at the tips, spines wearing away completely with maturation, scars disappearing with age and weather, easily peeling away from the endoperidium when moist. Endoperidium white when young, becoming orange gray (5B2-4), to brownish orange (5C4), then grayish brown (5D2, 6D2) in old age, sometimes with a golden green (4D8) shiny metallic appearance; smooth, persistent, and parchment-like. Gleba white and firm when young, dark olive green brown to olive brown (4F4-6) at the outer edges as the spores mature, becoming yellow brown to brown (5E4-5, 6E6) when mature; cottony, maturing from the top down. Subgleba white when young, becoming olive brown (4E6, 4F6) from the base up, with a metallic shine; chambered, 5-9 mm broad at the base of the gleba, tapered to a pinched base at the point of rhizomorph attachment, comprising the lower bottom half of the fruitbody. Diaphragm absent.

Basidiospores globose to subglobose;  (3.2) 4-4.8 (5.6) X (3.2) 4.0-4.8 (5.6) µm [xmr = 4.1-4.5 X 4.2-4.6 µm, xmm = 4.3 ± 0.3 X 4.4 ± 0.3 µm, Q = 0.8-1.2, Qmr = 0.9-1.0, Qmm = 0.9 ± 0.1, n = 25, s = 2]; golden brown in wet mounts; verrucose spores seen at 100x under oil emersion, prominent truncate verrucose spores seen with SEM; oil drop absent; spores thick-walled; pedicel rudimentary, up to 0.8 μm long; free-floating sterigmata absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Lycoperdon-type; thick up to 4-6 μm broad, walls up to 1 μm thick; golden brown in wet mounts, thicker walled threads with more pigmentation, thinner walls hyaline; elastic; incrusted with cellular debris, frequent irregular dichotomous branching, sinuous to straight, or undulate with sinuous edges along thin hyphal walls, knob-like projections present; tips attenuate to a round terminus. Pores abundant, round, and large, up to 1-2 μm broad, well-defined pores obvious and with a circular lip wrapping around the opening. Septa rare or absent. Paracapillitium present, rare to scarce in mid-aged fruitbodies. Exoperidium a combination of textura globulosa and texture intricata; composed of inflated thin-walled globose sphaerocysts, undifferentiated cellular tissue present, globose to hexagonal-shaped and brown pigmented; intertwined with hyphal elements, Endoperidium a combination of textura globulosa and textura intricata; composed of globose cells and entangled with loosely woven, yellow to golden pigmented dichotomously branching hyphae.

            Habitat: A common puffball in dense moist conifer forests, fruiting with winter rains. Growing as a solitary fruitbody to gregariously on soil under conifer duff, or among damp moss and other soil humus. Among the Geographic Subdivisions of California, this species is found in the Northwestern California, in the Cascade Range, in the Sierra Nevada, in Central Western California, and in Southwestern California.

Distribution: Known from many parts of the United States, and previously reported from Alaska, Arizona, Arkansas, California, Colorado, Connecticut, Delaware, Florida, Georgia, Hawaii, Idaho, Illinois, Indiana, Iowa, Louisiana, Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Montana, Nebraska, New Hampshire, New Mexico, New Jersey, New York, North Carolina, Ohio, Oregon, Pennsylvania, Tennessee, Texas, Utah, Vermont, Virginia, Washington, Washington DC, West Virginia, Wisconsin, and Wyoming. Also known from Austria, Belgium, Brazil, Canada, China, Costa Rica, Czechoslovakia, Denmark, England, France, Germany, Greece, Guatemala, Hungary, India, Italy, Japan, Luxembourg, Portugal, South Africa, Spain, Sweden, Mexico, Russia, Uruguay, and the USSR.

            Material Examined: CALIFORNIA, Calaveras Co.: Camp Connell, Highway 4, 1 mile east, 1,524 meters (5,000 feet) elev., gregarious in litter under conifers, 23 October 1983, H.D. Thiers (HDT 46653). Del Norte Co.: Highway 199 near the Oregon boarder, solitary in soil under conifers, 30 October 1971, H.D. Thiers (HDT 28516). El Dorado Co.: Crystal Basin Recreation Area, solitary in soil near rotten log, 16 October 1976, H.D. Thiers (HDT 36748). Humboldt Co.: Patrick’s Point State Park, gregarious in soil under conifers, 16 October 1971, H.D. Thiers (HDT 28363); Hoopa Indian Reservation, SE of Hoopa, along hospital road, scattered beneath Pseudotsuga menziesii (Douglas fir), 3 December 1972, D. Largent (DL 5563-2128)(HSU); Samra Forest, terrestrial under spruce and pine, 31 March 1973, D. Largent (DLL 5756-2126)(HSU); Manilla pine forest, terrestrial, 31 March 1973, D. Largent (Baroni1 204-5370); Samra Forest, solitary on mossy humus, 7 April 1973, D. Largent (DLL 5783-2127)(HSU); no notes, 4 December 1975, coll. unknown (380H00388EWT1)(HSU); Patrick’s Point State Park, area b, trail 8, terrestrial, 20 October 1986, D. Largent (DD 207-10277)(HSU); Humboldt Coast, terrestrial in damp thick humus under conifers, 2 November 2009, S.S. Jarvis (SSJ 380); Gray’s Falls Campground, Highway 299, growing on damp moss and humus, 15 December 2009, S.S. Jarvis (SSJ 368). Marin Co.: Audubon Canyon Ranch, Bolinas Ridge Road, solitary in duff under Pseudotsuga menziesii (Douglas fir), 27 December 1979, C. Calhoun (Calhoun 79-1296); Audubon Canyon Ranch, Bolinas Ridge Road, solitary in humus under mixed evergreen forest, 28 February 1981, C. Calhoun (Calhoun 81-2010); Audubon Canyon Ranch, Bolinas Ridge Road, scattered in soil in mixed evergreen forest, 28 March 1981, C. Calhoun (Calhoun 81-2313). Mendocino Co.: Jackson State Demonstration Forest, terrestrial on decomposed wood, 17 November 2009, D.E. Desjardin (SSJ 471). Monterey Co.: Aquahito Road, solitary in humus in pine woods, 19 January 1975, H.D. Thiers (HDT 33349). Napa Co.: Mt. Saint Helena, Lake County Highway, solitary under conifers in grass, 5 December 2007, S.S. Jarvis (SSJ 240); Angwin, Las Posadas Park, among damp moss and hard packed soil, 16 February 2010, S.S. Jarvis (SSJ 370); Angwin, Las Posadas Park, terrestrial along foot path in damp conifer woods, 28 December 2012, S.S. Jarvis (SSJ 481). Nevada Co.: Nevada City, Tahoe National Forest, Skillman Flat Camp, solitary in humus under conifer, 14 May 1960, H.D. Thiers (HDT 7612); Tahoe National Forest, East of Nevada City, Highway 20, solitary in humus in mixed woods, 11 November 1972, H.D. Thiers (HDT 30603). San Bernardino Co.: San Bernardino National Forest, subhypogeous protected under a bush, 7 October 2008, S.S. Jarvis (SSJ 254). San Mateo Co.: San Francisco Watershed, terrestrial, December 2009, MSSF collection (SSJ 353); San Francisco Watershed, terrestrial, December 2009, MSSF collection (SSJ 480). Sierra Co.: Yuba Pass, Highway 49, gregarious in soil under conifer, 15 September 1983, H.D. Thiers (HDT 46321); Bullards Bar Reservoir, Yuba Pass, growing on old mulch, 7 December 2008, F. Stevens (SSJ 277). Sonoma Co.: Santa Rosa, Calistoga Road, on hard packed old dirt road among grass, 5 December 2008, S.S. Jarvis (SSJ 273); Salt Point State Park, terrestrial, 7 March 2009, D. Viess (SSJ 445); Occidental, CYO Camp along the Bohemian Highway, terrestrial under conifers, 15 November 2009, D. Deshazer (SSJ 457); Occidental, CYO Camp along the Bohemian Highway, under conifers in damp forest, 17 January 2010, S.S. Jarvis (SSJ 376). Tehama Co.: Gurnsey Creek Campground, Highway 89, gregarious in soil under conifers, 11 October 1976, H.D. Thiers (HDT 36623). Tulare Co.: Sequoia National Park, Halstead Creek Trail, on dry duff under pine, 29 July 1945, L. Bonar (UCB695791)(UC). Yuba Co.: Bullards Bar Recreation Area, Schoolhouse Campground, solitary in soil in dense mixed conifer forest, H.D. Thiers (HDT 53986).

            Comments: There are four puffballs in California with similar features, requiring a light microscope to compare spore ornamentation and careful observation of the peridium characters for proper identification: Lycoperdon molle, Lycoperdon nigrescens, Lycoperdon perlatum, and Lycoperdon umbrinum. Lycoperdon molle can be distinguished by the granular and spinose exoperidium that does not leave scars on the endoperidium, and verrucose spores. Lycoperdon nigrescens is a puffball with a large sterile base, becomes dark brown to almost black, has prominent scars on the endoperidium where the spinose exoperidium has sloughed off, and verruculose spores. Lycoperdon perlatum is set apart from the others due to the exaggerated pseudostipe, spinose exoperidium that leaves prominent scars on the endoperidium, white to cream colors of the peridium layers throughout the life of the fruitbody, and echinulate spores. Lycoperdon umbrinum has a prominent pseudostipe, and a dark pigmented exoperidium that may seem glabrous to furfuraceous depending on the development of the granular and spinose ornamentation, in addition to verruculose spores. The very small spines of Lycoperdon umbrinum will fall off and leave minute scars on the endoperidium, which are much smaller than what is found on Lycoperdon nigrescens or Lycoperdon perlatum. Lycoperdon umbrinum has verrucose spores similar to Lycoperdon molle, however L. molle can grow much larger and has a much more prominent ornamentation on the exoperidium. The ITS sequence data from this study shows Lycoperdon umbrinum nested with Lycoperdon vernimontanum within the Lycoperdon clade with 100% bootstrap and 100% PP support. One of the sequences of Lycoperdon umbrinum more closely related to Lycoperdon vernimontanum than it does to another sequence of its same species. This suggests that additional genes should be sequenced to further clarify the taxonomic position of Lycoperdon umbrinum amongst the other taxa in the Lycoperdaceae in order to better understand the relationships of these fungi.

 

Lycoperdon utriforme Bull., 1790.

                                                                                                            (fig. 35, 42, 75)

Reported synonyms

= Lycoperdon bovista var. bovista L., Sp. pl. 2: 1183. 1753. 


= Lycoperdon echinatum Schaeff., Fung. bavar. palat. nasc. (Ratisbonae) 4: 128, tab. 186. 1774.

= Lycoperdon bovista var. echinatum (Schaeff.) Huds., Fl. Angl., Edn 2 2: 642. 1778.

= Lycoperdon caelatum Bull., Herb. Fr. 9: tab. 430. 1789.


= Lycoperdon bovista Pers., Observ. mycol. (Lipsiae) 1: 4. 1796.


= Bovista utriformis (Bull.) Fr., Syst. mycol. (Lundae) 3(1): 25. 1829.


= Utraria caelata (Bull.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 2 5: 369. 1873.


= Utraria utriformis (Bull.) Quél., Mém. Soc. Émul. Montbéliard, Sér. 2 5: 369. 1873.

= Lycoperdon sinclairii Berk., J. Roy. Microscop. Soc.: 716. 1887.


= Calvatia caelata (Bull.) Morgan, J. Cincinnati Soc. Nat. Hist. 12: 169. 1890.


= Calvatia hungarica Hollós, 19: 84. 1904.

= Calvatia utriformis (Bull.) Jaap, Verh. bot. Ver. Prov. Brandenb. 59: 37. 1918.


= Calvatia caelata var. hungarica (Hollós) F. Šmarda, Fl. ČSR, B-1, Gasteromycetes: 285. 1958.

= Handkea utriformis (Bull.) Kreisel, Nova Hedwigia 48(3-4): 288. 1989.


Type: According to Kreisel (1962), the type locality of this species is France. No holotype specimen was designated in the protologue. Plate 450 of Herbier de la France, Bulliard 1790, can serve as an iconotype (http://raf.dessins.free.fr/2bgal/img.php?id_img=11076).

Gasterocarp 75-100 mm tall x 60-100 mm broad; obpyriform, depressed at the apex, bulbous at the base; rhizomorph composed of a thick mat of mycelium incrusted with soil and vegetal debris, tapered and root-like; ostiole slow to develop through a lateral laceration at the apex, developing into a round hole. Exoperidium white to cream when young (4A2-3), becoming yellow to yellow gray (4A5, 4B5-6, 4C6), with patches of grayish brown (6E3-4) in old and where sun exposed; ornamentation of fine minute furfuraceous spines and granules covering on the entire exoperidium, quickly rubbing off when moist and when handled, larger spines and dense patches of granules covering the top 2/3 of the gasterocarp, smoother along the lower regions of the fruitbody, but with spines within the plicate folds and crevices, the apex becoming heavy and collapsing upon its own weight with moisture. Endoperidium cream white at first (4A3), becoming brownish orange, then yellow brown to dark yellow brown (5C4-5, 5E4-5, 5F5-6); glabrous, waxy when moist, parchment-like when dry, persistent, staining yellow when moist and cut (4A5). Gleba white and firm at first, turning grayish yellow (4B4-5), then grayish yellow to shades of brownish green (4C6-7, 4E5, 4F8), then becoming dark brown (4F4, 5F4) when spores are mature; chambered, becoming chunky as the gleba matures, to completely pulverulent with age, maturing from the center outward. Subgleba cream color when young (4A3), turning yellow to olive green in the center (4A6, 4D5, 4E6), brown at the edges (4F8), becoming metallic and shinny with age; robust, composing up to 2/3 of the entire gasterocarp, chambered with large locules, larger chambers along the diaphragm and more compact chambers at the base. Diaphragm well-defined, membranous, up to 2mm thick when young and moist, thinning with age and desiccation.

            Basidiospores globose; (3.5) 4-5.5 X 4.0-5.5 (6.0) µm [xmr = 4.6 X 5.2 µm, xmm = 4.6 ± 0.0 X 5.2 ± 0.0 µm, Q = 0.8, n = 35, s = 1]; dark amber in wet mounts; minutely verruculose under light microscopy, under SEM densely packed knob-like bumps with a flattened apex; large prominent oil drop; spores thick-walled; pedicel short with a torn end up to ± 0.8 µm long; free-floating sterigmata not present in wet mounts; spores of equal size in wet mounts. Eucapillitium Calvatia-type; 1-6 µm broad, sometimes up to 7 µm broad at dichotomous branch junctures, with walls up to 0.8 µm thick; eucapillitium dark amber in wet mounts; fragile, fragmenting easily; eucapillitium threads heavily incrusted with cellular debris, dichotomous branches abundant, hyphal strands straight, sinuous along thin hyphal walls, tips taper to a small thin rounded terminus. Pores and pore-like slits present in wet mounts; pores round and small to medium-sized, abundant; slit-like cracks abundant, slits stellate-shaped to sinuous or straight, in various directions, under SEM these cracks appear to be the results of fragile hyphae cracking and not morphologically derived, with these eucapillitium often disarticulating at the cracks. Septa present, not abundant. Paracapillitium present to scarce or absent in the center of the gleba, abundant at the inner endoperidium wall. Exoperidium a combination of textura intricata and textura globulosa; composed of thin-walled, hyaline, short, septate, linking hyphal elements, and intermixed with thin-walled globose to subglobose inflated sphaerocyst cells. Endoperidium a combination of textura intricata and textura globulosa; composed of long hyphal elements in a parallel alignment, 6-8 µm broad with walls up to 0.8 µm thick, intertwined with large thin-walled inflated globose cells. Subgleba composed of long interwoven hyphal elements; hyaline, cream to green colored in KOH; un-branched, threads up to 4 µm broad with walls up to 2 µm thick, hyphae tapering to a fine thin blunt terminus, non-septate, elastic, some threads making a u-shape without breaking, developing into the chambers of the subgleba. Diaphragm composed of fragile hyphal elements, golden brown in KOH mounts, septa scarce, knob-like projections present, slit-like pores present, and simple dichotomous branching.

Habitat: A rare species that does not fruit every year. Found growing on soil or in dry duff, solitarily or in small gregarious groups. Mostly fruiting in the spring, with spring rain, among oak woodland. Among the Geographic Subdivisions of California, this species can be found in Northwestern California, and in Central Western California.

Distribution: Known from many parts of the United States, and previously reported from California, Colorado, Idaho, Kansas, Pennsylvania, Michigan, Montana, New York, Oregon, South Dakota, Washington, Wisconsin, Wyoming, Virginia. Also known from Austria, Canada, Germany, Iceland, India, Sweden, Switzerland, and Uruguay.

Material Examined: CALIFORNIA, Alameda Co.: Oakland, Piedmont Cemetery, on soil of a steep hill above the creek, in shade under oaks and bay laurel, 28 January 2009, S.S. Jarvis (SSJ 280). Marin Co.: Pt. Reyes National Seashore, in grass under coast live oak, January 2014, S.S. Jarvis (SSJ 486). Mendocino Co.: Masonite Redwood Grave, Highway 128, gregarious and terrestrial under redwood, 7 December 1969, A. Wenneken (Wenneken 211). San Mateo Co.: San Francisco Watershed, solitary in soil under Cupressus macrocarpa (Monterey cypress), 10 February 1970, H.D. Thiers (HDT 21303); San Francisco Watershed, solitary in soil under hardwood, 19 March 1970. Sonoma Co.: Occidental, Camp Meeker, solitary in humus in mixed woods, 13 April 1963, D. Largent (Largent 10234).

            Comments: Lycoperdon utriforme has many features in common with Lycoperdon lloydianum and Lycoperdon pratense. All three of these species have minute furfuraceous and granulose ornamentation on the exoperidium and a prominent diaphragm. However, Lycoperdon utriforme will get about four times larger in overall size than the other two diaphragm-forming meadow puffballs. Lycoperdon utriforme has Calvatia-type eucapillitium, which is fragile and forms slit-like pits along the hyphal walls as the threads dry up and break apart with maturation. The nature of slits was also observed and discussed by Lang (1993), being a factor of fragile eucapillitium.

Kreisel (1989) transferred Lycoperdon utriforme into Handkea, and attempted to split it into three separate varieties based on smooth spores and exoperidium characters; Handkea utriformis var. hungarica (Hollós) Kreisel, Handkea utriformis var. utriformis (Bull.) Kreisel, and Handkea utriformis var. gruberi (A.H. Smith) Kreisel. However, as with most work done using only light microscopy, the reliability of this work is questionable based on the maturity of the fruitbodies that were examined to make these determinations. Under scanning electron microscopy, it is clear that most puffball spores begin relatively smooth, and as they swell and expand their ornamentation becomes apparent. Such is the case with Lycoperdon utriforme; most fruitbodies will exhibit relatively smooth spores until they are mature. Martin (1997) concludes that there is not sufficient evidence to have more than one variety of L. utriforme.

Historically, due to the size and shape of this puffball, it has been recognized in the genera Calvatia, Bovista, and Utraria. Recent molecular studies (Bates 2004, Larsson & Jeppson 2008) show that Handkea utriformis is placed within the Lycoperdon clade. Bates (2004) showed that H. utriformis was basal to the Lycoperdon clade in a strict consensus maximum-parsimony analysis. This evidence was supported when Larsson and Jeppson (2008) showed H. utriformis being basal to all Lycoperdon in a strict consensus tree, and L. utriforme being basal to the Utraria clade in a 50% majority rule consensus tree. Because of these results, this puffball was accepted in Lycoperdon as Lycoperdon utriforme (Larsson and Jeppson 2008, 2009). In the analysis of puffballs from California, supports most of these claims. The ML tree here shows Lycoperdon utriforme as a group within the Lycoperdon clade with 82% bootstrap and less than 70% PP, but not basal to it. This ML tree also shows Lycoperdon utriforme sister to Calbovista subsculpta, forming a small group with 93% bootstrap and 100% PP support. This suggests that multi loci gene analysis will be necessary to further clarify the taxonomic position of Lycoperdon utriforme and Calbovista subsculpta within the Lycoperdaceae.

 

Lycoperdon vernimontanum S. Jarvis & F. Stevens, sp. nov.                                                                                                                                                                        (fig. 36, 40, 41, 42, 76)

Misapplied names:

            = Calvatia lycoperdoides A.H. Sm. 1964, nom. illegit. (non Calvatia lycoperdoides Kościelny & Wojt., 1935.)

             = Handkea lycoperdoides Kreisel, Nova Hedwigia 48(3-4): 287. 1989.

Type: A holotype collection is located in the San Francisco State University Herbarium, California, Plumas County, Lake Davis, Jenkins point, 39’55’31.84” N, 120’32’12.57” W, elevation 6060, S. Jarvis (SSJ 450). A paratype for this species is located in the University of Michigan Herbarium (MICH 7736) from Karney Lakes, Boise Co., Idaho.

            Gasterocarp (11) 24-40 mm tall x (15) 22-45 mm broad; globose, depressed-globose, to obpyriforme, tapered and plicate at the base, wrinkled at the base; rhizomorph bulky and root-like, 15-26 mm long x 8-10 mm broad at the basal attachment, heavily incrusted with soil and vegetal debris, remaining attached to the soil; ostiole opening via peridium slowly breaking apart at the apex to form longitudinal to stellate cracks that expose the gleba, sometimes in more than one location. Exoperidium white to cream when young (4A3-4B5), grayish yellow (4B2), yellowish brown (5E5) to light yellowish brown (5D5-6), pale olive ochre (4D5), or tan-beige (5E5-6) to dark brown gray (6F3) with age, coloring in uneven patches; surface covered in hexagonal scales and small furfuraceous spines, tiny dense scurfy spines radiating out of the exoperidium and fusing at the tips to form larger spines in early development, becoming appressed with age, raised, flattened, hexagonal or irregular patches (< 1 mm broad) develop at the apex as the fruitbody expands, becoming stellate-cracked with desiccation and maturation, spinose-scurfy scales remain along the base and within the plicate folds in late maturity, overall exoperidium tissue persistent and remaining adherent to the endoperidium, eventually will crack and flake off in patches from the apex down to expose the endoperidium and gleba. Endoperidium yellow beige (4B5), becoming light brown (5D5), sometime with olive tints (4E7), unevenly colored and darkening with sun exposure; peridium layer up to 0.5mm thick, glabrous, firm, persistent and parchment-like when dry and mature, more or less remaining attached to the exoperidium, seen only in late development as the exoperidium partially sloughs off. Gleba yellow brown (5E4) to pale yellow brown (5E5) or dark yellowish brown (5F5), becoming dark olive brown in late maturity (4F5), turning bright olive green in KOH; somewhat cottony, becoming subpulverulent to powdery with age, fragile, crumbling apart in patches or clumps, maturing from the center outward. Subgleba bright yellow (4A5) when mature; rarely up to 2-3 mm thick, reduced and almost non-existent in most larger and older fruitbodies, compact, chambered with compressed locules, cellular type tissue.

            Basidiospores globose; (4-) 4.5-8 X (-4) 4.5-8 (-8.5) µm [xmr = 5.6-7.1 X 5.8-7.0 µm, xmm = 6.53 ± 0.6 X 6.56 ± 0.5 µm, Q = 0.9-1.1, Qmr = 0.96-1.02, Qmm = 0.99 ± 0.0, n = 25, s = 4]; spores golden yellow in KOH mounts; roughened to echinulate under light microscope, with SEM the ornamentation composed of loosely packed, truncate echinulate spines or round bumps connected at the top to form a tri-pod-like protuberance, some connected with ridges; with a central oil drop; spores thick-walled; pedicel rudimentary or ± 0.8-1.5 μm, long and broken at the end, under SEM the pedicel ends have a clean terminus; free-floating sterigmata fragments not present in wet mounts; spores of various size under light microscope. Eucapillitium Calvatia-type; 3.2-4.8 μm in diameter, walls ± 0.8 μm thick; golden brown in wet mounts; fragile and mostly broken up in the center of the gleba early in maturation; relatively narrow hyphal threads, glabrous, branching scarce to absent and dichotomous when occurring, straight to subundulate with knob-like projections; tips semi-attenuated with a rounded terminus. Pores small to medium-sized; of varying shapes, round, slit-like cracks, and sinuous cracks all present in a single fruitbody. Septa abundant, hyphae disarticulating at the septum wall. Paracapillitium present but not abundant, septate, incrusted with debris, staining light blue with a lactophenol cotton blue reaction. Exoperidium textura globulosa; composed of thin-walled globose sphaerocyst cells, 12-30 μm broad x 24-34 μm long, with walls up to 2 μm thick, orbicular to semi-round or club-shaped, septate linked cells, tightly compact. Endoperidium textura intricata; composed of septate, tightly interwoven hyphae, 2-6 μm broad, thin-walled.

            Habitat: Solitary to scattered, and terrestrial in leaf litter under conifers. Fruiting after the snow melts in spring through early summer, and with bouts of rain in higher elevations of the Sierra Nevada. Often found among the duff, soil, and leaf litter under the drip zones of Pinus jeffreyi (Jeffery pine), and Pinus contorta (lodgepole pine). Growing along the edges of dirt roads where the soil is compact and disturbed, suggesting a saprotrophic habit. Among the Geographic Subdivisions of California, this species is found in the Cascade Range, in the Sierra Nevada, and in the high elevations of the San Bernardino Mountains of South Western CA.

            Distribution: Known from parts of the United States, and previously reported from Alaska, California, Colorado, Idaho, Oregon, Utah, Mississippi, and Washington.

            Material Examined: CALIFORNIA, Plumas Co.: Grizzly Creek Campground, in Pinus jeffreyi (Jeffery pines), 10 June 2011, F. Stevens (SSJ 417); Davis Lake, Jenkins Point, in soil at the edge of the dirt road at 6060 elevation, 11 June 2011, F. Stevens (SSJ 450); Grizzly Creek Campground, site # 35, in Pinus jeffreyi (Jeffery pines), (23 June 2012, F. Stevens (SSJ 451; TYPE). San Bernardino Co.: Camp Osceola, between Ponderosa pine, Pinus ponderosa, and Quercus sp., 26 September 1976, Tom Tang (UCB1462464)(UC). Sierra Co.: Chapman Creek Campground, Yuba Pass, on soil with Abies concolor and Pinus lambertiana, 10 June 1961, V. Demoulin, (UCB1408395)(UC); Yuba Pass, Highway 49, open area, June 1986, coll. unknown (HS 3141, cf. vernimontanum); Wild Plum Campground, near Sierra City, gregarious on soil under mixed conifers, October 1989, H.D. Thiers (HDT 52785); Chapman Creek Campground, Highway 49, scattered in grassy area in Montane fir-pine forest, June 1996, D.E. Desjardin (DED 6476); Yuba Pass Campground, in soil and pine duff at 2,051 meters (6,730 feet) elevation, 5 June 2012, S.S. Jarvis (SSJ 463). Siskiyou Co.: Sisson southern trail, Mt. Shasta, on the ground, 7 April 1947, M. Cooke, (UCBM138940)(UC); Andesite Station, Mt. Shasta, on the ground, 9 April 1947, M. Cooke, (UCBM138941)(UC); Sisson southern trail, Mt. Shasta, on the ground, 17 June 1947, M. Cooke, (UCBM138944)(UC); N. Mt. Shasta, on sandy volcanic soil, 1 July 1968, V. Demoulin, (UCB1463208)(UC). Tulare Co.: Sequoia National Park, General Grant National Park, terrestrial, 1970, L. Bonar, (UCB506724)(UC). Tuolumne Co.: Pinecrest Campground, gregarious in humus under conifers, June 1965, H.D. Thiers (HDT 12629); Bumblebee Meadow, scattered under mixed conifers, 25 July 1965, W.J. Sundberg (HDT 282). IDAHO, Boise Co.: Karney Lakes, 1959, E. Trueblood 920, labeled “Calvatia lycoperdoides A.H. Smith” (MICH 7736). WASHINGTON, Glacier Snohomish Co.: on the ground, 8 July 1920, E. Morse (UCB977170)(UC).

            Comments:  In 1964, Smith described Calvatia lycoperdoides A.H. Sm. based on several specimens collected in: Butte Co., California; Boise Co., Idaho; Valley Co., Idaho; and Pierce Co., Washington. Smith’s name is an illegitimate homonym of Calvatia lycoperdoides Kościelny & Wojt. described from Europe in 1935. Kreisel (1989) transferred Smith's illegitimate name into Handkea where it was accepted as a new name Handkea lycoperdoides Kreisel. This transfer was based on the presence of slit-like pits reported by Smith (1964). Kreisel stated that he “did not examine material,” hence the species concept of Handkea lycoperdoides is questionable. The holotype specimen cited by Smith (1964) is missing. The paratype specimens reported in the prologue represent several different species, indicating that Calvatia lycoperdoides A.H. Sm. (= Handkea lycoperdoides Kreisel) represents a nomen dubium. One of the paratype specimens (MICH 7736) matches the description from California material, recognized here as the new species Lycoperdon vernimontanum.

Lycoperdon vernimontanum can be confused easily for a small Calvatia. The gasterocarp dehisces in a Calvatia-like manner, has Calvatia-type eucapillitium, and exoperidium ornamentation similar to another small sierra puffball, Calvatia lloydii. However, in contrast to C. lloydii, L. vernimontanum has a very compact, almost non-existant subgleba. Bates (2009) showed in his sequence analysis that Handkea lycoperdoides was sister to Lycoperdon umbrinum, and clearly part of the Lycoperdon clade. Supporting this claim, the ITS sequence data from this study on California taxa shows Lycoperdon umbrinum nested with Lycoperdon vernimontanum within the Lycoperdon clade with 100% bootstrap and 100% PP support. One of the sequences of Lycoperdon umbrinum is more closely related to Lycoperdon vernimontanum than it is to another sequence of its same species. This suggests that additional genes should be sequenced to further clarify the taxonomic position of Lycoperdon umbrinum amongst the other taxa in the Lycoperdaceae in order to better understand the relationships of these fungi. Furthermore, four sequences of Lycoperdon vernimontanum from various regions of the Sierra Nevada form their own group within the Lycoperdon clade with 96% bootstrap and 100% PP. Therefore supporting the claim that this puffball is a new taxon, Lycoperdon vernimontanum

 

MYCENASTRUM Desv.:

Mycenastrum corium (Guers.) Desv., Annls Sci. Nat., Bot., sér. 2 17: 147. 1842.                                                                                                                                                (fig. 37, 38, 39, 77)

Basionym º Lycoperdon corium Guers., in Lamarck & de Candolle, Fl. franç., Edn 3 (Paris) 2: 598. 1805.

Reported synonyms

= Scleroderma corium (Guers.) A.H. Graves, in Duby, Bot. Gall., Edn 2 (Paris) 2: 852. 1830.

= Mycenastrum corium (Guers.) Desv., Annls Sci. Nat., Bot., Sér. 2 17:147. 1842.


            = Mycenastrum chilense Mont., Ann. Sci. Nat. Bot., Sér. 2, 20: 375. 1843.

= Sterrebekia corium (Guers.) Fr., K. svenska Vetensk-Akad. Handl. 69: 150. 1849.

= Scleroderma chilense (Mont.) De Toni, inn Sacardo’s Syll. Fung. 7: 139. 1888.

Type: Kreisel (1962) and Calonge (1998) list the type locality of this species as Rouen-Sotteville, France. However, type material does not seem to exist in any of the herbaria collections in Europe. Bates (2004) lists the type material as “not extant.”

            Gasterocarp 10-125 mm tall x 25-140 mm broad; globose to subglobose, depressed globose to ovate at the apex, often tapered at the base to a point, can be plicate with large folds around the base; rhizomorph composed of a tuft of fine hyphal hairs that anchor the fruitbody; ostiole slow to develop, formed of horizontal or stellate crack along the apex and downward, breaking through the peridium layers, opening up wide with many cracks running vertically down each side, peridium curving outward, entire fruitbody forming into a stellate shape in late maturation, continuous cracking with peridium layers falling off until a stellate cup-like base remains. Exoperidium white to cream yellow (4A2-4), becoming to brownish orange to dark yellow brown (5C4, 5D4, 5F6-7, 5F4-8), becoming metallic grey brown with age (5F2-3), color sometimes uneven and mottled, darker where sun exposed; ornamentation composed of a thick matted-tomentose to flocculose or delicately cottony layer when young, up to 1 mm thick when young, thinning with age and drying, cracking and developing patch-like scales as the fruitbody swells and grows, patches at first becoming thin up-turned sheets with fruitbody desiccation, peeling off slowly and flaking off, smaller patches remaining adherent to the endoperidium, parchment-like and tough. Endoperidium white to cream when young (4A2-3), turning brownish orange (5A3, 5C4), becoming dark yellowish brown (5E5, 5F5), sometimes becoming bleached grey brown (5E3-4) at the apex with sun exposure, entirely or in large patches, bleached grey brown tones seen after sloughing of exoperidium and in very old fruitbodies; 1-3 mm thick, leather-like in texture when young and moist, tough and hardened with age, rigid when dry, large sections of the endoperidium falling away until only the rooted base of the fruitbody remains in the ground. Gleba firm and white when young (4A2), dark yellowish brown when mature (5F4-6); maturing from the upper middle of the gleba outward, becoming clumpy to powdery as spores mature, then completely pulverulent with full maturity; seldom long stalagmite-like formations present inside old specimens, these structures attached to the inner peridium wall at the base of the fruitbody and composed of hyphal elements radiating upwards, incrusted with spores, a feature unique to Mycenastrum corium. Subgleba absent. Paracapillitium absent.

Basidiospores globose; 8-12.6 X 8-12.6 µm [xmr = 10.8-11.2 X 11.2-11.5 µm, xmm = 11.2 ± 025 X 11.38 ± 0.21 µm, Q = 0.9-1.0, Qmr = 0.97, Qmm = 0.97 ± 0.0, n = 25, s = 2]; golden brown in wet mounts, turning green in KOH mounts; young spores with slight dextrinoid reaction; roughened to pitted ornamentation seen under light microscope with oil emersion, pitted ornamentation distinct with SEM; central oil drop absent; spores thick-walled; pedicel absent; free-floating sterigmata absent from wet mounts; spores of equal size in wet mounts. Eucapillitium Mycenastrum-type; thick-walled eucapillitium threads up to 8-12 µm broad, with walls 0.8-3 µm thick; golden brown in wet mounts; elastic; glabrous, with many ramified branches and abundant dichotomous branching to create an antler-like formation, hyphal segments straight, knob-like projections absent, separated individual short threads; tips attenuate to a round pointed terminus. Pores absent. Septa absent. Paracapillitium absent. Exoperidium textura intricata; composed of long hyphal strands with walls up to 2 µm thick. Endoperidium textura intricata; composed of tightly woven, long hyphae. Basidia developing in fascicles, not in a hymenium layer and collapsing early in development stages.

            Habitat: Mycenastrum corium is a species that seems to favor very fertile soils. It is often found growing on dung heaps, compost piles, in very tall grass, or in open fields that get fertilized on a regular basis. Often reoccurring in a perennial growing pattern, this puffball can be collected on disturbed ground where there is heavy foot traffic, and in sandy soil. Fruiting in early winter after light rain, or in spring after heavy rain, this is a common puffball in years when the conditions are correct. Occurring as caespitose clusters, in triplets or pairs, singularly, or in gregarious rings. Typically, up to one-third of the fruitbody will be rooted in the soil, but not completely subhypogeous. Sometimes breaking away from the point of attachment at maturity, the gasterocarp will roll along the ground with the wind and disperse spores, or remain attached to the soil. The original description describes its occurrence along sandy coastlines in France, so it may be possible to collect Mycenastrum corium along the coasts of California. Among these Geographic Subdivisions of California, this species is found in the Great Central Valley, rarely in the Sierra Nevada, in Central Western California, Southwestern California.

Distribution: Known only from the Western United States, and previously reported from Arizona, California, Colorado, Florida, Georgia, Illinois, Iowa, Kansas, Minnesota, Mississippi, Montana, New Mexico, North Dakota, Oregon, Texas, Wisconsin, and Wyoming. Also reported from Africa, Australia, Austria, British Colombia, Canada, Hungary, India, Mexico, New Zealand, Scandinavia, South Africa, and South America: Argentina, Chile, and Uruguay.

Material Examined: CALIFORNIA, Alameda Co.: Berkeley, December 1901, T.H. Gilbert (UCB506610)(UC); Berkeley, on a dung heap, December 1901, T.H. Gilbert (UCB506621)(UC); Knowland Park, above the Oakland Zoo, in an old compost pile, 30 November 2011, S.S. Jarvis (SSJ 418). Bute Co.: Chico, in an old corral, 29 December 1936, V. Mentzer (UCB564434)(UC). Fresno Co.: Coalinga, along road to Mendota, in shifting sand, 26 April 1949, H. Manson (UCB793320)(UC). Napa Co.: Napa, Conn Valley Road, 21 April 1942, E.E. Morse (UCB671505)(UC). Orange Co.: Pasadena, 27 August 1919, J. Pearson (UCB472080)(UC). San Diego Co.: El Centro, 1940, L. Bonar (UCB507211)(UC); San Diego, on lawn under cypress trees, October 1940, V. Miller (UCB660270)(UC). Stanislaus Co.: Oakdale, on bare elevated ground in a field, 2 January 1980, T. Tang (UCB1570965)(UC); Oakdale, on bare elevated ground in field, 4 January 1980, T. Tang (UCB1569809)(UC). Tulare Co.: Sequoia National Park, Tharp Long Meadow, terrestrial, 9 October 1931, E.E. Morse (UCB507208)(UC). Yolo Co.: Harris Farm, in pasture near Davis, 18 February 2002, D.E. Desjardin (DED 7353).

            Comments:  Mycenastrum corium once regarded as a monotypic genus (Homrich and Wright 1973), is now described as sixteen species on Index Fungorum. There is a wide range in gasterocarp shape and color of the gleba tissue, which is why there are many synonyms for Mycenastrum corium. Some have speculated that the red to orange color of the gleba in some specimens legitimized the recognition of a new subspecies, Mycenastrum corium ssp. ferrugineum (O.K. Miller 2005). The stellate splitting and hardening of the peridium layers is unique to this puffball, as well as the microscopic characters of the eucapillitium and spores. Calvatia pachyderma and Calvatia fragilis are both medium to large-sized puffballs, which might become misidentified as Mycenastrum corium when very old specimens are found. All three of these will grow in grasslands and have thick peridium layers that will slough off until only a cup-like base remains in the soil. Four sequences of Mycenastrum corium from both California and Europe form a monophyletic group outside of the Lycoperdaceae with 100% bootstrap and 100% PP support. The ITS sequence data suggests that this species is outside of the Lycoperdaceae clade, and closer related to Lepiota cristata. The next closest puffball in relation to Mycenastrum corium is Abstoma townei, (which also has pitted spores), but with quite a distance and no support. This suggests that additional genes should be sequenced to further clarify the taxonomic position of Mycenastrum corium amongst the Lycoperdaceae.

 

Additional Taxa

Bovista cacao P. Ponce de León, Fieldiana, Bot. 38(1): 1. 1975.

Reported synonyms:
           

            = Calvatia umbrina Lloyd, Mycol. Writ. 1: 2. 1902.

            Type: Erhorn s.n. Mountain View, California, Lloyd, catalog no. 20679 (BPI).

            This species was not noticed in any of the herbarium collections investigated for this study. We currently do not know what this is. However, it was described from California. This author has not collected any puffballs fitting the description for Bovista cacao, nor has material been examined at this time. According to the literature, this species was first described by Lloyd (1904) as Calvatia umbrina, and transferred to Bovista cacao by Ponce de Leon in 1975 based on it’s resemblance to Bovista pila, having Bovista-type eucapillitium, and smooth spores with a short pedicle. Lloyd compared this species to Calvatia hesperia in 1904, but Ponce de Leon suggests “there is no similarity between these two species.” There are not any reports of this fungi in the MyCoPortal website (http://mycoportal.org/portal/index.php), nor the BPI database (http://nt.ars-grin.gov/fungaldatabases/specimens/specimens.cfm)

 

Gastropila hesperia (J.B. Morgan) P. Ponce de León, Phytologia 33(7): 460. 1976.

Reported Synonyms:

                        = Calvatia hesperia Morgan, J. Cincinnati Soc. Nat. Hist. 18: 39. 1895.


            = Handkea hesperia (Morgan) Kreisel, Nova Hedwigia 48(3-4): 286. 1989.

            Type: McClatchie s.n., Los Angeles Co., Pasadena, CA, 1895 (NY)           

            The results from this study suggest that Calvatia hesperia may be synonymous with both Disciseda atra and Disciseda levispora, and that Calvatia hesperia might be the oldest name for the species (Morgan 1895). Calvatia hesperia was not noticed in any of the herbarium collections investigated for this study. This author has not collected any puffballs fitting the description for Calvatia hesperia, nor has material been examined at this time. According to the BPI database, it has been reported in San Francisco, Claremont, and California (Smithsonian Institute collection # 22578, 22579, 22580, respectively). The type collection is located at the New York Botanical Garden, with three collection numbers  (00291963, 00065667, 00068898), There are two collections determined as C. hesperia at UC Berkeley; from Yosemite, (UC1998626), and from Santa Barbara, Santa Cruz Island, Los Pinos del Sur ridge (UC1860653). There are other reports of this puffball from Fresno, California, and Mississippi. 

 

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