Battarrea phalloides
Encyclopedia
Battarrea phalloides is an inedible species of mushroom
Mushroom
A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or on its food source. The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus; hence the word "mushroom" is most often applied to those fungi that...

 in the family Agaricaceae
Agaricaceae
The Agaricaceae are a family of basidiomycete fungi and includes the genus Agaricus, as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. The genus contains 85 genera and 1340 species.-Genera:...

, and the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 of the genus Battarrea
Battarrea
Battarrea is a genus of mushrooms in the Tulostomataceae family.-Species list:*Battarrea arenicola Copel. *Battarrea franciscana Copel. *Battarrea gaudichaudii Mont. *Battarrea guachiparum Speg....

. Known in the vernacular
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 as the scaley-stalked puffball or the sandy stiltball, it has a woody, slender, and scaly stem that can typically reach 40 centimetres (15.7 in) in length. Although its general appearance resembles a more typical agaric
Agaric
An agaric is a type of fungal fruiting body characterized by the presence of a pileus that is clearly differentiated from the stipe , with lamellae on the underside of the pileus. "Agaric" can also refer to a basidiomycete species characterized by an agaric-type fruiting body...

 with stem and gills, atop the stem is a spore
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...

 sac, consisting of an endoperidium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...

 and a gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

. 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.

Taxonomy

This species was first named by James Dickson
James Dickson (biologist)
James Dickson was a Scottish nurseryman, plant collector, botanist and mycologist. Between 1785 and 1801 he published his Fasciculus plantarum cryptogamicarum Britanniae, a four-volume work in which he published over 400 species of algae and fungi that occur in the British Isles...

 in 1785 as Lycoperdon phalloides, based on specimens collected in England. In 1801, Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 transferred it into the newly described genus Battarrea
Battarrea
Battarrea is a genus of mushrooms in the Tulostomataceae family.-Species list:*Battarrea arenicola Copel. *Battarrea franciscana Copel. *Battarrea gaudichaudii Mont. *Battarrea guachiparum Speg....

, named after the Italian mycologist Giovanni Antonio Battarra
Giovanni Antonio Battarra
Giovanni Antonio Battarra was an Italian mycologist. In 1755, he published Fungorum Agri Ariminensis Historia.- External links :* Online edition of Fungorum Agri Ariminensis Historia...

. Battarrea phalloides is also known
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 as the "scaly-stalked puffball" and the "sandy stiltball"; a common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 given to the species B. stevenii is the "desert drumstick".

Historically, there has been uncertainty as to whether the European species known as Battarrea stevenii was a unique species or merely a polymorphic
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...

 variant of B. phalloides. In 1995, mycologist Roy Watling opined that B. stevenii differs in having spores that are colored more orange-tawny
Tawny (color)
Tawny is a yellowish brown color. The word means "tan-colored," from Anglo-Norman tauné "associated with the brownish-yellow of tanned leather," from Old French tané "to tan hides," from Medieval Latin tannare, from tannum "crushed oak bark," used in tanning leather, probably from a Celtic source...

, slightly larger (5–6.5 x 5.75–7 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is 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
Mucilage
Mucilage is a thick, gluey substance produced by most plants and some microorganisms. It is a polar glycoprotein and an exopolysaccharide.It occurs in various parts of nearly all classes of plant, usually in relatively small percentages, and is frequently associated with other substances, such as...

 in the volva
Volva
The word volva can refer to:* Völva, a pagan Norse shaman* Volva * Volva , a genus of sea snails....

 and the innermost parts of the stem
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...

. 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
Internal transcribed spacer
ITS refers to a piece of non-functional RNA situated between structural ribosomal RNAs on a common precursor transcript. Read from 5' to 3', this polycistronic rRNA precursor transcript contains the 5' external transcribed sequence , 18S rRNA, ITS1, 5.8S rRNA, ITS2, 28S rRNA and finally the 3'ETS...

 regions of the 5.8S rDNA
Ribosomal DNA
Ribosomal DNA codes for ribosomal RNA. The ribosome is an intracellular macromolecule that produces proteins or polypeptide chains. The ribosome itself consists of a composite of proteins and RNA. As shown in the figure, rDNA consists of a tandem repeat of a unit segment, an operon, composed of...

—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. Several years later in 2006, based on a study of specimens collected in China, other scientists considered them to be independent species.

Description

Mature specimens of Battarrea phalloides roughly resemble the typical agaric
Agaric
An agaric is a type of fungal fruiting body characterized by the presence of a pileus that is clearly differentiated from the stipe , with lamellae on the underside of the pileus. "Agaric" can also refer to a basidiomycete species characterized by an agaric-type fruiting body...

 mushroom stature of stem and cap
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...

. 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
Volva (mycology)
The volva is a mycological term to describe a cup-like structure at the base of a mushroom that is a remnant of the universal veil. This macrofeature is important in wild mushroom identification due to it being an easily observed, taxonomically significant feature which frequently signifies a...

 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
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...

. 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
Cone (geometry)
A cone is an n-dimensional geometric shape that tapers smoothly from a base to a point called the apex or vertex. Formally, it is the solid figure formed by the locus of all straight line segments that join the apex to the base...

 "head" 1 to 3 cm (0.393700787401575 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 (0.15748031496063 to 0.590551181102362 in) thick.

One study reported a specimen found in Mexico with a length of 65 cm (25.6 in). Typically, the spore case is 3 to 5 cm (1.2 to 2 in) broad by 1 to 3.5 cm (0.393700787401575 to 1.4 ) tall. The hollow stalk
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...

 is pale brown to brown, woody, and has a fibrous, scaly, or even woolly surface. The mature gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

, 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
Volva (mycology)
The volva is a mycological term to describe a cup-like structure at the base of a mushroom that is a remnant of the universal veil. This macrofeature is important in wild mushroom identification due to it being an easily observed, taxonomically significant feature which frequently signifies a...

 is up to 15 by broad, unattached to the stalk, and formed by two distinct, separated tissue layers. The inner layer resembles the scales of the stem, consisting of hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

e that are 3–18 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...

 in diameter, closely arranged (nearly parallel), septate, sparsely branched, yellowish ochre
Ochre
Ochre is the term for both a golden-yellow or light yellow brown color and for a form of earth pigment which produces the color. The pigment can also be used to create a reddish tint known as "red ochre". The more rarely used terms "purple ochre" and "brown ochre" also exist for variant hues...

, with clamps
Clamp connection
A clamp connection is a structure formed by growing hyphal cells of certain fungi. It is created to ensure each septum, or segment of hypha separated by crossed walls, receives a set of differing nuclei, which are obtained through mating of hyphae of differing sexual types...

 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.

Microscopic characteristics

The thick-walled spores
Basidiospore
A basidiospore is a reproductive spore produced by Basidiomycete fungi. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are produced by specialized fungal cells called basidia. In grills under a cap of one common species in the phylum of...

 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 hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

e 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
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

 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
Hyaline
The term hyaline denotes a substance with a glass-like appearance.-Histopathology:In histopathological medical usage, a hyaline substance appears glassy and pink after being stained with haematoxylin and eosin — usually it is an acellular, proteinaceous material...

 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.

Similar species

The closely related species Battarrea diguettii is known in the United States from the Mojave desert
Mojave Desert
The Mojave Desert occupies a significant portion of southeastern California and smaller parts of central California, southern Nevada, southwestern Utah and northwestern Arizona, in the United States...

, 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 70 centimetres (27.6 in).

Edibility

Battarrea phalloides (as well as Battarrea stevenii) is typically described as unknown edibility, or inedible. Older specimens may smell unpleasant.

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 hedge
Hedge
Hedge may refer to:* Hedge or hedgerow, line of closely spaced shrubs planted to act as a barrier* Hedge , investment made to limit loss* Hedge , intentionally non-committal or ambiguous sentence fragments-See also:...

row), sometimes growing amongst elm suckers
Basal shoot
A basal shoot, root sprout, adventitious shoot, water sprout or sucker is a shoot or cane which grows from a bud at the base of a tree or shrub or from its roots. This shoot then becomes, or takes the form of, a singular plant. A plant that produces suckers is referred to as surculose...

. 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
Arid
A region is said to be arid when it is characterized by a severe lack of available water, to the extent of hindering or even preventing the growth and development of plant and animal life...

 and semiarid areas, on coastal dunes, found from sea level up to 2550 m (8,366.1 ft) high. The mushroom has been associated with the quick-growing evergreen tree Schinus molle
Schinus molle
Peruvian Pepper is an evergreen tree that grows to 15 meters . It is native to the Peruvian Andes...

, as well as Lycium brevipes
Lycium brevipes
Lycium brevipes is a species of flowering plant in the nightshade family known by the common name Baja desert-thorn. It is native to northwestern Mexico and it occurs in California as far as the Sonoran Desert as well as some of the Channel Islands. It grows in the scrub of desert and coastline. It...

, Solanum hindsianum, Salicornia subterminalis, Atriplex linearis, Quercus agrifolia and Optunia species, in coastal dunes. The largest fruit bodies were found on floodplain
Floodplain
A floodplain, or flood plain, is a flat or nearly flat land adjacent a stream or river that stretches from the banks of its channel to the base of the enclosing valley walls and experiences flooding during periods of high discharge...

s with halophilic
Halophile
Halophiles are extremophile organisms that thrive in environments with very high concentrations of salt. The name comes from the Greek for "salt-loving". While the term is perhaps most often applied to some halophiles classified into the Archaea domain, there are also bacterial halophiles and some...

 (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
Western Canada
Western Canada, also referred to as the Western provinces and commonly as the West, is a region of Canada that includes the four provinces west of the province of Ontario.- Provinces :...

; the USA, where it is confined to the west— Southern California, New Mexico
New Mexico
New Mexico is a state located in the southwest and western regions of the United States. New Mexico is also usually considered one of the Mountain States. With a population density of 16 per square mile, New Mexico is the sixth-most sparsely inhabited U.S...

, and Arizona— and Mexico. It has also been reported growing in South America (Brazil), Africa (Morocco
Morocco
Morocco , officially the Kingdom of Morocco , is a country located in North Africa. It has a population of more than 32 million and an area of 710,850 km², and also primarily administers the disputed region of the Western Sahara...

), Europe (Belgium), China, and Australia. Due to a decline in sightings, B. phalloides was granted legal protection in Hungary
Hungary
Hungary , officially the Republic of Hungary , is a landlocked country in Central Europe. It is situated in the Carpathian Basin and is bordered by Slovakia to the north, Ukraine and Romania to the east, Serbia and Croatia to the south, Slovenia to the southwest and Austria to the west. The...

in 2005.

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.

External links

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