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Phallaceae is a family of fungi, commonly known as stinkhorns, within the order Phallales. Stinkhorns have a worldwide distribution, but are especially prevalent in tropical regions. They are known for their foul-smelling, sticky
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
masses, or gleba, borne on the end of a stalk called the receptaculum. The characteristic fruiting-body structure, a single, unbranched receptaculum with an externally attached gleba on the upper part, distinguishes the Phallaceae from other families in the Phallales. The spore mass typically smells of
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
or
dung Dung most often refers to animal feces. Dung may also refer to: Science and technology * Dry animal dung fuel * Manure * Cow dung * Coprolite, fossilized feces * Dung beetle Art * Mundungus Fletcher or "Dung", a character in the Harry Potter n ...
, and attracts flies,
beetles Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
and other insects to help disperse the spores. Although there is great diversity in body structure shape among the various
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, all species in the Phallaceae begin their development as oval or round structures known as "eggs". According to a 2008 estimate, the family contains 21 genera and 77 species.


Description

Species of stinkhorns have gasteroid, or internally produced spores. Fruit bodies originate as a gelatinous, spherical, or egg-shaped structure that may be completely or partially buried underground. The peridium, the outer layer of the egg, is white, or purple/red, with two or three layers. The outer layer is thin, membranous, and elastic, while the inner layer is thicker, gelatinous, and continuous. At maturity, the peridium opens up and remains as a volva at the base of the receptaculum. The fertile portion of the fruiting body is often borne on the end of a wide, fleshy or spongy stalk (as in the Phallales), which may be cylindrical, star-shaped, or reticulate (forming a network). They may be brightly colored, sometimes with a lattice- or veil-like membrane enclosing and protecting the spores. The spore-containing substance, the gleba, is typically
gelatin Gelatin or gelatine (from la, gelatus meaning "stiff" or "frozen") is a translucent, colorless, flavorless food ingredient, commonly derived from collagen taken from animal body parts. It is brittle when dry and rubbery when moist. It may also ...
ous, often foul-smelling, and deliquescent (becoming liquid from the absorption of water). The gleba is formed on the exterior face of the cap or the upper part of the receptacle. The basidia are small and narrowly club-shaped or
fusiform Fusiform means having a spindle-like shape that is wide in the middle and tapers at both ends. It is similar to the lemon-shape, but often implies a focal broadening of a structure that continues from one or both ends, such as an aneurysm on a b ...
, short-lived (evanescent), with four to eight sterigmata. The spores are usually ellipsoid or cylindrical in shape,
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
or pale brown, smooth, more or less smooth-walled, and truncated at the base. The gelatinous layer of the stinkhorn '' (Clathrus ruber)'' contains high concentrations of potassium, calcium, manganese, and iron. A moderate amount of calcium is found in the
receptacle Receptacle may refer to: Biology * Receptacle (botany), a plant anatomical part * Seminal receptacle, a sperm storage site in some insects Electrical engineering * Automobile auxiliary power outlet, formerly known as ''cigarette lighter recep ...
and gleba as well as a high amount of manganese is also found in the gleba. Retaining a high potassium concentration is important for the growth of the stem of the fruiting body called the carpophore, where potassium plays a key component in regulating osmotic pressure. This osmotic pressure helps maintain the shape of the carpophore. The high concentration of potassium in the plant cell and the low permeability of the cell membrane also drives water into the cell through
flux Flux describes any effect that appears to pass or travel (whether it actually moves or not) through a surface or substance. Flux is a concept in applied mathematics and vector calculus which has many applications to physics. For transport ph ...
. The calcium found in the gelatinous layer contributes to the slimy characteristics of the gel and protects the carpophore during growth. The strong adhesive gelatinous layer consists of
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with wa ...
s that are formed by the sequestration of calcium ions. The high manganese and iron concentrations in the gelatinous layer and gleba are theorized to produce enzymes that produce sugars and odorous compounds that play a role in attracting insects.


Genera

; ''
Anthurus ''Anthurus'' is a genus of fungi in the family Phallaceae. Species , Species Fungorum ''Index Fungorum'' is an international project to index all formal names (scientific names) in the fungus kingdom. the project is based at the Royal Bo ...
'' Kalchbr. & MacOwan (1880):Fruiting bodies have a short stalk from which arises a spore-bearing structure (the receptaculum) of 5–8 arched arms. These arms, initially joined at the top, disconnect and curve irregularly to expose the inner surface of each arm, which is covered with green spore-containing gleba. Spores are 3–4 × 1–1.5 μm.Ellis and Ellis, p. 219. ; '' Aporophallus'' Möller (1895) ; '' Aseroë'' Labill. (1800):Mature fruiting bodies contain a roughly cylindrical white or pinkish stalk approximately 6 × 2 cm, with a volva at the base. At the top is a bright red disc with a variable number of arms, typically 3–7 cm long. The gleba found on the disc and inner side of the arms is slimy, foetid, and green colored. Spores are
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
, with dimensions of 4–6 1.5–2 μm. ''
Aseroë rubra ''Aseroe rubra'', commonly known as the anemone stinkhorn, sea anemone fungus and starfish fungus, is a common and widespread basidiomycete fungus recognizable for its foul odour of carrion and its sea anemone shape when mature. Found in garden ...
'', an Australian and Pacific species which has spread to Europe and North America. ;''
Blumenavia ''Blumenavia'' is a genus of fungi in the family Phallaceae. The genus contains three species found in South America and Africa. Species References External links * Phallales Agaricomycetes genera {{Phallales-stub ...
'' Möller (1895) ; '' Clathrus'' P. Micheli ex L. (1753):Fruiting bodies are latticed (clathrate), and made of hollow tubular arms that originate from the basal tissue within the volva. Spores are elliptical, smooth, hyaline, with dimensions of 4–6 ×1.5–2.5 μm.Miller and Miller, p. 80. Examples include '' Clathrus ruber'', the lattice stinkhorn, ''
Clathrus archeri ''Clathrus archeri'' (synonyms ''Lysurus archeri'', ''Anthurus archeri'', ''Pseudocolus archeri''), commonly known as octopus stinkhorn or devil's fingers, is a fungus which has a global distribution. Using rDNA, Geastrales, Gauteriales and Phal ...
'', the octopus stinkhorn, and ''
Clathrus columnatus ''Clathrus columnatus'', commonly known as the column stinkhorn, is a saprobic species of basidiomycete fungus in the family Phallaceae. It has a widespread distribution, and has been found in Africa, Australasia, and the Americas. It may have ...
'', the columned stinkhorn. ;''
Colus ''Colus'' (lat. ''colus'', distaff) is a genus of sea snails, marine gastropod mollusks in the family Colidae, the true whelks and the like. Evolution ''Colus'' is related to the New Zealand genus ''Austrofusus''. Taxonomy The name ''Colus'' ...
'' Cavalier & Séchier (1835) : A genus of four species with fruit bodies that are a short stalk from which six columns arise, joining at the top to form a red, arching, clathrate receptaculum. The gleba is smeared on the inside surface of the receptaculum.Miller and Miller, p. 91. ; '' Echinophalus'' Henn. (1898) ; ''
Endophallus A drone is a male honey bee. Unlike the female worker bee, drones do not have stingers. They gather neither nectar nor pollen and are unable to feed without assistance from worker bees. A drone's only role is to mate with a maiden queen in n ...
'' Zang & Petersen (1989): Contains a single species, ''E. yunnanensis'' found in China, that resembles ''Phallus'' except for a peridium that is separated from the base of the stem and which does not persist as a volva. ; ''
Ileodictyon ''Ileodictyon'' is a genus of fungi in the family Phallaceae. Basidiocarps (fruit bodies) are clathroid (resembling those in the genus ''Clathrus''), emerging from egg-like peridia and forming spongy "arms" which connect to create a cage-like la ...
''
Tul. Louis René Étienne Tulasne, a.k.a. Edmond Tulasne (12 September 1815 – 22 December 1885) was a French botanist and mycologist born in Azay-le-Rideau. He originally studied law at Poitiers, but his interest later turned to botany. As a yo ...
ex M. Raoul (1844)
:Fruiting bodies are latticed (clathrate), and have gelatinous arms that lie sessile within the volva. Spores are elliptical, and have dimensions of 4–6 1.5–2.5 μm. The New Zealand native, ''
Ileodictyon cibarium ''Ileodictyon cibarium'' is a saprotrophic species of fungus in the family Phallaceae. It is native to Australia and New Zealand, where it is commonly known as the basket fungus or the white basket fungus, alluding to its fruit bodies, shaped li ...
'', known as the basket fungus, has a fruiting body shaped somewhat like a round or oval ball with interlaced or latticed branches. ; '' Itajahya'' Möller (1895):Characters in this genus include a white calyptra (tissue which covers the top of the fruiting body to which the gleba is attached), lamellate plates covered with gleba. The gleba has a white mottled surface, and the pileus appears wig-like when removed of the gleba. The thick, stout stalk has many chambered walls. ; ''
Kobayasia ''Kobayasia'' is a genus of fungi in the family Phallaceae. It contains the species ''Kobayasia nipponica'' and ''Kobayasia kunmingica'' . It is known as シラタマタケ ''shiratamatake''; "white ball mushroom" in the Japanese language. Th ...
'' (Kobayasi) S. Imai & A. Kawam. (1958): This genus circumscribes the single species ''Kobayasia nipponica'', found in Japan in 1958. ; '' Laternea'' Turpin (1822): ; '' Ligiella'' J.A. Sáenz (1980):Fruiting bodies are 4.5–6.5 cm long, 2.2–5 cm in diameter, and have four or five thick white chambered arms that are joined at the top, but free at the base. This monotypic genus, containing the single species ''L. rodrigueziana'', is known only from
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
. ; ''
Lysurus ''Arremon'' is a genus of neotropical birds in the family Passerellidae. With the exception of the green-striped brushfinch which is endemic to Mexico, all species are found in South America, with a few reaching Central America. These spa ...
'' Fr. (1823):Fruiting bodies consist of a long stalk with 3 or 4 short, thick arms. These arms, initially joined together, but usually separate at maturity, are covered with the brownish-olivaceous gleba. Spores are ellipsoidal, with dimensions of 4–5 × 1.5–2 μm. ; ''
Mutinus ''Mutinus'' is a genus of fungi in the family Phallaceae. The genus was first described by Elias Magnus Fries in 1849. According to the ''Dictionary of the Fungi'' (10th edition, 2008), the widespread genus contains 12 species. Species Etymo ...
'' (Huds.) Fr. (1849):The mature fruiting body has a spongy, cylindrical hollow stalk which ends in a slender, tapered, sometimes curved head covered with the dark olivaceous, slimy gleba. In older specimens, the gleba may be washed or worn off to show the orange or red color of the head itself. Notable species include the dog stinkhorn ''
Mutinus caninus ''Mutinus caninus'', commonly known as the dog stinkhorn, is a small thin, phallus-shaped woodland fungus, with a dark tip. It is often found growing in small groups on wood debris, or in leaf litter, during summer and autumn in Europe, Asia, ...
''. ; ''
Neolysurus ''Neolysurus'' is a genus of fungi in the family Phallaceae. A monotypic genus, it contains the single species ''Neolysurus arcipulvinus''. Species References External links Index Fungorum
Phallales Monotypic Basidiomycota genera ...
'' O.K. Mill., Ovrebo & Burk (1991): This genus contains the single species ''Neolysurus arcipulvinus'', described from
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
. The stipe and arms have a tubular construction, similar to species from ''Lysurus''. ''Neolysurus'' is unique in having a long stipe, ending in arms or columns that branch and interconnect to support a cushion-shaped, olive green gleba. The glebal cushion is divided into polygonal compartments by a fine pinkish white, solid mesh. The
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some ...
is continuous between the mesh. ; ''
Phallus A phallus is a penis (especially when erect), an object that resembles a penis, or a mimetic image of an erect penis. In art history a figure with an erect penis is described as ithyphallic. Any object that symbolically—or, more precisel ...
'' Junius ex L. (1753):In species of ''Phallus'', the receptaculum is a tall unbranched stalk that ends in a cap-like structure that bears the gleba. Notable species include '' Phallus impudicus'', the common stinkhorn, ''
Phallus hadriani ''Phallus hadriani'', commonly known as the dune stinkhorn or the sand stinkhorn, is a species of fungus in the Phallaceae (stinkhorn) family. It is a widely distributed species, and is native to Asia, Europe, and North America. In Australia, ...
'', ''
Phallus ravenelii ''Phallus ravenelii'', commonly known as Ravenel's stinkhorn, is a fungus in the Phallaceae (stinkhorn) family. It is found in eastern North America. Its mushrooms commonly grow in large clusters and are noted for their foul odor and phallic ...
'', and '' Phallus indusiatus'' (syn. ''Dictyophora indusiata''), the Chinese "bamboo fungus", eaten as a food in southwestern China after the foul smelling cap is removed. ; ''
Protubera ''Protubera'' is a genus of fungi in the family Phallogastraceae. The genus has a widespread distribution in tropical and subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification ...
'' Möller (1895):Fruiting bodies are egg-shaped, 2.2–3.5 cm in diameter by 3–4 cm high, with a dull white, soft exoperidium (roughly 1 mm thick) and a grayish exoperidium (3–4 mm thick). The gleba is contained within internal chambers that are separated by whitish, gelatinous tissue that originates from a columella-like, gelatinous central core. Spores are elliptical, smooth, nearly hyaline, and 3.5–4.5 1.5–2 μm.Miller and Miller, p. 83. ; ''
Protuberella ''Protuberella'' is a genus of fungi in the family Phallaceae. A monotypic genus, it contains the single species ''Protuberella borealis''. Species References External links Index Fungorum
Phallales Monotypic Basidiomycota genera ...
'' (S. Imai) S. Imai & Kawam. (1958):This genus resembles the ''Protubera'', except that the tissue is "gelatinous fleshy in the peridium and not so distinctly gelatinous as in ''Protubera maracuja'', they are less gelatinous in the nature of the sterile strands than those of ''P. maracuja''". Spores are clylindric, thin-walled, and 3.7–5 2–2.5 μm. This monotypic genus contains the single species ''Protuberella borealis'', known only from Asia. ;'' Pseudoclathrus'' B. Liu & Y.S. Bau (1980) ; ''
Pseudocolus ''Pseudocolus'' is a genus of fungi in the stinkhorn family. The sporocarp (fungi), fruit bodies have three or four simple arms that are initially joined at the tip, but often break apart. The tips of the arms are covered with a slimy, foul-smel ...
'' Lloyd (1907) ; '' Staheliomyces'' E. Fisch. (1921)


Volatile compounds

The number and type of volatile compounds produced by stinkhorn mushrooms changes throughout its life course. When the stinkhorn is still in its egg stage, the total amount of volatile substances is relatively low. The sulfuric odorous compounds (that attract insects) is absent. During this stage the mushroom’s veil covers up the slime covered cap that contains the spores. When the stinkhorn grows into a more mature-fruit body, there are over twenty-two volatile compounds found in the mature-fruit bodies including dimethyl oligosulfides, aroma-active
phenylalanine Phenylalanine (symbol Phe or F) is an essential α-amino acid with the formula . It can be viewed as a benzyl group substituted for the methyl group of alanine, or a phenyl group in place of a terminal hydrogen of alanine. This essential amino a ...
s and various terpenoids that act as insect attractants. In an over-ripe stinkhorn, forty-one volatile components are present, many of which also act as insect attractants. Dimethyl oligosulfides are not as commonly found in over-ripe fruit as they were in the mature fruit bodies since many of the insects already carried the smelly slime away. The stinkhorn contains compounds of
carrion Carrion () is the decaying flesh of dead animals, including human flesh. Overview Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters (or scavengers) include crows, vultures, c ...
, such as oligosulfides, and of feces, such as phenol, indole and p-cresol. This provides evidence for mimicry of carrion and feces and convergence in the putrid scents of fungus. The stinkhorn’s scent is similar to that of rotten meat and dog feces. This scent profile supports the assumption that the stinkhorn mimics fly brood sites and food sources. The flies are being lured in by the exploitation of their innate attraction to smell of the decaying animal matter that the stinkhorn gives off.


References


Books cited

* *


External links

* *
Index Fungorum


* ttp://www.mushroomexpert.com/phallaceae.html Stinkhorns: The Phallaceae and Clathraceae {{Authority control Mushroom types Phallales Basidiomycota families Taxa named by August Carl Joseph Corda Taxa described in 1842