| Battarrea phalloides | |
|---|---|
| Scientific classification | |
| Kingdom: | Fungi |
| Division: | Basidiomycota |
| Class: | Agaricomycetes |
| Order: | Agaricales |
| Family: | Agaricaceae |
| Genus: | Battarrea |
| Species: | B. phalloides |
| Binomial name | |
| Battarrea phalloides (Dicks.) Pers. |
|
| Synonyms | |
|
Lycoperdon phalloides Dicks. |
|
Battarrea phalloides[1] is an inedible species of mushroom in the family Agaricaceae,[2] and the type species of the genus Battarrea. Known in the vernacular as the scaley-stalked puffball or the sandy stiltball, it has a woody, slender, and scaly stem that can typically reach 40 centimeters (15.7 in) in length. Although its general appearance resembles a more typical agaric with stem and gills, atop the stem is a spore sac, consisting of an endoperidium and a gleba. Battarrea phalloides is found in dry, sandy locations throughout North America, primarily in western regions; it has also been collected in South America, Africa, Australia, Europe, and China. There is currently some disagreement in the literature as to whether the European species B. stevensii is the same species as B. phalloides.
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This species was first named by James Dickson in 1785 as Lycoperdon phalloides, based on specimens collected in England. In 1801, Christian Hendrik Persoon transferred it into the newly described genus Battarrea,[3] named after the Italian mycologist Giovanni Antonio Battarra.[4] Battarrea phalloides is also known as the "scaly-stalked puffball"[5] and the "sandy stiltball";[6] a common name given to the species B. stevenii is the "desert drumstick".[7]
Historically, there has been uncertainty as to whether the European species known as Battarrea stevenii was a unique species or merely a polymorphic variant of B. phalloides. In 1995, mycologist Roy Watling opined that B. stevenii differs in having spores that are colored more orange-tawny, slightly larger (5–6.5 x 5.75–7 µm, as opposed to 4.5–5.25 x 4.5–5.75 µm), and less ornamented. Further, B. stevenii is thought to have a larger fruit body size, a more coarsely scaly stipe, and lack of mucilage in the volva and the innermost parts of the stem.[4] The presence or absence of mucilage has been traditionally considered the significant characteristic separating the species.[8] In 1904, Lászlo Hollós proposed the idea of a single polymorphic species;[9] in 1942, Paul Marshall Rea, after studying 25 specimens from southern California, concluded that B. stevenii was conspecific with B. phalloides and represented a single species.[10] An analysis of a number of European specimens—using both macroscopic and microscopic characteristics in addition to molecular analysis of ITS regions of the 5.8S rDNA—also suggested both to be conspecific. The authors of this study considered the differences in spore ornamentation and stipe hyphae to be insufficient to discriminate them as two species; they did, however, note that their inability to locate the type material for both species limited the conclusiveness of their suggestion.[11] Several years later in 2006, based on a study of specimens collected in China, other scientists considered them to be independent species.[12]
Mature specimens of Battarrea phalloides roughly resemble the typical agaric mushroom stature of stem and cap. However, rather than a cap with gills, this species has a spore sac atop the stem. When young, the fruit body is roughly spherical and completely encased in an outer wall which later splits in a circumscissile fashion (along a circular or equatorial line), the lower wall forming a volva and the upper part forming scales that cover the inner wall. The upper part rolls upward and backward and eventually falls away in one piece, exposing a spore sac lined with a narrow ring of capillitium and spores.[13] The spores are sticky.[5] As these are carried away by the wind, the drying action of the latter cause the edges of the peridium to shrivel and roll up more, exposing more spores. This is continued until the upper half of the peridium has shriveled and blown away and there remains only a few spores, which may be washed away by rain.[13]
The fruit body develops rapidly;[4] when mature, it is rust-colored, with a hemispherical to somewhat conical "head" 1 to 3 cm (0.4 to 1.2 in) in diameter, and with a stalk up to 40 cm (15.7 in) long by 0.4 to 1.5 cm (0.2 to 0.6 in) thick.[5]
One study reported a specimen found in Mexico with a length of 65 cm (25.6 in).[14] Typically, the spore case is 3 to 5 cm (1.2 to 2.0 in) broad by 1 to 3.5 cm (0.4 to 1.4 in) tall.[5] The hollow stalk is pale brown to brown, woody, and has a fibrous, scaly, or even woolly surface.[14] The mature gleba, which is eventually exposed when the peridial cap is shed, has a rust-brown color. The cap may persist after the spore mass is dispersed and form a disc-like unit that slides down the stalk like a ring.[5] The fragile sac-shaped volva is up to 15 by 13 cm (5.9 by 5.1 in) broad, unattached to the stalk, and formed by two distinct, separated tissue layers. The inner layer resembles the scales of the stem, consisting of hyphae that are 3–18 µm in diameter, closely arranged (nearly parallel), septate, sparsely branched, yellowish ochre, with clamps at some septa. The outer layer of fungal tissue is thicker, membranous, sometimes with a corky texture when dry, and dirty white. It consists of pale yellow intertwined hyphae that are difficult to distinguish individually, and without remains of a gelatinous matrix. Fruit bodies may persist for several months after they have dried.[15]
The thick-walled spores are roughly spherical, rusty-brown, finely and densely warted, and have diameters of 5–6.5 µm. Elaters are 50–80 by 4–6 µm, and have ring-like or spiral thickenings.[16] The endoperidium consists of densely interwoven hyphae that are 3–9 µm in diameter and walls less than 1 µm thick; they are septate, branched, pale yellow, with clamp connections.
The gleba is largely made up of two types of threads. The pseudocapillitium has hyphae up to 5 µm diameter, mostly thin walled, smooth, septate, sparsely branched, hyaline to pale yellow, with clamps. The elaters have diameters of 3.5–7 µm and are 32–70 µm long; they are pale yellow, smooth-walled, tapered and cylindrical with spiral thickenings. Glebal elaters are aseptate and not branched.[14]
The closely related species Battarrea diguettii is known in the United States from the Mojave desert, and differs from B. phalloides in that the spore sac emerges by ripping through the top of the exoperidium, rather than by circumscissile rupture. The endoperidium of B. diguettii is also smaller, and the spores emerge through a number of pores on the upper surface of the spore sac.[15] Battarrea stevenii can grow taller, up to 70 centimeters (27.6 in).[17]
Battarrea phalloides (as well as Battarrea stevenii) is typically described as unknown edibility,[15][17] or inedible.[5] Older specimens may smell unpleasant.[5]
Battarrea phalloides may be found growing solitary to scattered on dry, sandy hedgebanks (raised or mounded boundary feature, often topped by a hedgerow), sometimes growing amongst elm suckers. It is a relatively rare species, but may be locally abundant in some locations.[5][16] In Mexico, where it is only known from the north and central part of the country,[14] it has been usually collected in arid and semiarid areas, on coastal dunes, found from sea level up to 2,550 m (8,370 ft) high. The mushroom has been associated with the quick-growing evergreen tree Schinus molle, as well as Lycium brevipes, Solanum hindsianum, Salicornia subterminalis, Atriplex linearis, Quercus agrifolia and Optunia species, in coastal dunes.[14] The largest fruit bodies were found on floodplains with halophilic (i.e., thriving in high-salt conditions) vegetation.[14] In Belgium, specimens were found on sandy soil under dead elderflower bushes.[18]
In North America, Battarrea phalloides has been collected from the Yukon Territory,[15] western Canada;[7] the USA, where it is confined to the west—[5] Southern California,[10] New Mexico,[19] and Arizona—[20] and Mexico.[21] It has also been reported growing in South America (Brazil),[22] Africa (Morocco),[23] Europe (Belgium),[18] China,[12] and Australia.[24][25] Due to a decline in sightings, B. phalloides was granted legal protection in Hungary in 2005.[26]
The habitat and range of Battarrea stevenii include arid regions of the western and southwestern United States,[17] Australia,[25] South Africa,[27] and several European countries,[11] including Russia.[28]