Battarrea phalloides

Template:Taxobox
***Battarrea phalloides*** is an inedible species of mushroom in the family Agaricaceae, and the type species of the genus *Battarrea*. Known in the vernacular as the **scaley-stalked puffball**, **sandy stiltball**, or **desert stalked puffball**, it has a woody, slender, and shaggy or scaly stem that is typically up to Template:Convert in length. Although its general appearance resembles an agaric with stem and gills, atop the stem is a spore sac, consisting of an peridium and a powdery internal gleba. In maturity, the spore sac ruptures to release the spores. *Battarrea phalloides* is found in dry, sandy locations throughout the world, and has been collected from Africa, Asia, Australia, Europe, North America (primarily in western regions), and South America. There is currently some disagreement in the literature as to whether the European *B. stevensii* is the same species as *B. phalloides*.

## Taxonomy

The species was first mentioned in the scientific literature in 1784, when Thomas Jenkinson Woodward described a new species found by Mr. Humphreys of Norwich. It was named by James Dickson in 1785 as *Lycoperdon phalloides*, and the type locality was Suffolk, England. Christian Hendrik Persoon sanctioned the name when he transferred to *Battarrea* in his 1801 *Synopsis Methodica Fungorum*, a newly circumscribed genus named after the Italian mycologist Giovanni Antonio Battarra. Synonyms include *Phallus campanulatus*, published by Miles Berkeley in 1842 based on collections made by Charles Darwin in Maldonado (Uruguay) in 1833, and *Ithyphallus campanulatus*, a new combination based on Berkeley's name, published by Diederich Franz Leonhard von Schlechtendal in 1933.

The specific epithet *phalloides* means phallus-like, and refers to the similarity of the volva to the genus *Phallus*. *Battarrea phalloides* is commonly known as the "scaly-stalked puffball" and the "sandy stiltball"; a common name given to the species *B. stevenii* is the "desert drumstick". In Cyprus, it is known locally as "Donkey fungus", a name that may be derived from the fact that the spores were once used as a natural antiseptic and antipruritic agent applied to the backs of donkeys, or for their morphological resemblance to the animal's penis.

Historically, there has been uncertainty as to whether the European species known as *Battarrea stevenii* is a unique species or merely a polymorphic variant of *B. phalloides*. The taxon has been described as both a variety (as *B. phalloides* var. *stevenii* by Cleland and Cheel in 1916) and as a form (as *B. phalloides* f. *stevenii* by Calonge in 2004). In 1995, mycologist Roy Watling opined that *B. stevenii* differs in having spores that are colored more orange-tawny, slightly larger (5–6.5 by 5.75–7 µm, as opposed to 4.5–5.25 by 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. The presence or absence of mucilage has been traditionally considered the significant characteristic separating the species. In 1904, Lászlo Hollós proposed the idea of a single polymorphic species; 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. 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. The conspecificity of the species was supported in a later study that used traditional and molecular methods to compare field-collected English specimens and an extensive range of herbarium material collected from around the world. They found that material labeled *B. stevenii* generally had a greater range of cap and stem sizes, while that labeled *B. phalloides* was generally more consistent, and smaller. However, the *B. phalloides* material they studied was generally from similar habitats in the UK, while the *B. stevenii* material originated from a wide variety of locales and habitats, suggesting that environmental factors influence fruit body morphology. Evidence pointing to conspecificity included the continuum of spore sizes between the two, and the lack of any significant differences between their DNA. In 2006, however, based on a study of specimens collected in China, other scientists considered them to be independent species.

## Description

 File:Battarrea phalloides 40051.jpg
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 (exoperidium) that 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. The spores are sticky. 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.

The fruit body develops rapidly; when mature, it is rust-colored, with a hemispherical to somewhat conical "head" Template:Convert in diameter, and with a stalk up to Template:Convert long by Template:Convert thick.

One study reported a specimen found in Mexico with a length of Template:Convert. Typically, the spore case is Template:Convert broad by Template:Convert tall. The hollow stalk is pale brown to brown, woody, and has a fibrous, scaly, or even woolly surface. 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. The fragile sac-shaped volva is up to Template:Convert 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.

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. 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.

### Edibility

 File:Battarrea phalloides 40049.jpg
*Battarrea phalloides* (as well as *Battarrea stevenii*) is typically described as unknown edibility, or inedible. In Cyprus, the immature egg-form of the fruit body is eaten. Older specimens may smell unpleasant. The spores of *B. stevenii* are used as a cicatrizant—a product that promotes healing through the formation of scar tissue–by the Criollo herdsmen of Gran Chaco in northern Argentina.

### Similar species

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. *Battarrea stevenii* can grow taller, up to Template:Convert. *Podaxis pistillaris*, commonly known as the "desert shaggy mane", occurs in dry locales similar to *B. phalloides*, but can be distinguished by its shaggy, elongated cap.

## Habitat and distribution

*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. In Mexico, where it is only known from the north and central part of the country, it has been usually collected in arid and semiarid areas, on coastal dunes, found from sea level up to Template:Convert 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 *Opuntia* species, in coastal dunes. The largest fruit bodies were found on floodplains with halophilic (i.e., thriving in high-salt conditions) vegetation. In Belgium, specimens were found on sandy soil under dead elderflower bushes.

In North America, *Battarrea phalloides* has been collected from the Yukon Territory, western Canada; the USA, where it is confined to the west— Southern California, New Mexico, and Arizona— Mexico, and Hawaii. It has also been reported growing in South America (Brazil), Africa (Morocco), Europe (Belgium), China, and Australia. Due to a decline in sightings, *B. phalloides* was granted legal protection in Hungary in 2005, making it a finable offense to pick them. It received similar protected status in the United Kingdom in 1998.

The habitat and range of *Battarrea stevenii* include arid regions of the western and southwestern United States, Australia, South Africa, and several European countries, including Russia.

## Notes and references

Notes
1. ↑ Orthographic variations include *Batarrea*, *Battarea*, *Battarraea*, and *Battaraea*.

References

1. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Coetzee 1992`
2. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Kirk 2008`
3. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Woodward 1784`
4. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Dickson 1785`
5. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Persoon 1801`
6. ↑ 6.0 6.1 6.2 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Watling 1995`
7. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Berkeley 1842`
8. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlFungorum synonymy: Battarrea phalloides`
9. ↑ 9.0 9.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Jeffries 2004`
10. ↑ 10.0 10.1 10.2 10.3 10.4 10.5 10.6 10.7 10.8 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Arora 1986`
11. ↑ Authorities & Vocabularies (Library of Congress): Sandy stiltball. Library of Congress. Retrieved on July 20, 2009.
12. ↑ 12.0 12.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Schalkwijk-Barendsen 1991`
13. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Kyriakou 2009`
14. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlMycoBank: Battarrea phalloides var. stevenii`
15. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlMycoBank: Battarrea phalloides f. stevenii`
16. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Cunningham 1932`
17. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Hollós 1904`
18. ↑ 18.0 18.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Rea 1942`
19. ↑ 19.0 19.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Martin 2000`
20. ↑ 20.0 20.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Liu 2006`
21. ↑ 21.0 21.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Gaumann 1928`
22. ↑ 22.0 22.1 22.2 22.3 22.4 22.5 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Esqueda 2002`
23. ↑ 23.0 23.1 23.2 23.3 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Orr 1979`
24. ↑ 24.0 24.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Ellis 1990`
25. ↑ 25.0 25.1 25.2 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Miller 2006`
26. ↑ 26.0 26.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Roberts 2011`
27. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Scarpa 2004`
28. ↑ 28.0 28.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `Walleyn 2000`
29. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Dittmer 1978`
30. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `urlAMP - Checklists of Arizona Fungi`
31. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Cortez 2009`
32. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Hemmes 2002`
33. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Sobestiansky 2005`
34. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Outcoumit 2007`
35. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Orchard 1996`
36. ↑ 36.0 36.1 Cite error: Invalid `<ref>` tag; no text was provided for refs named `May2003.2B`
37. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Siller 2006`
38. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Evans 2006`
39. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `UKWildlife`
40. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Levin 1985`
41. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `Lloyd 1903`

## External links

* Template:IndexFungorum

Template:Good article
Template:Taxonbar