Clathrus columnatus
Encyclopedia
Clathrus columnatus, commonly known as the column stinkhorn, is a saprobic species of basidiomycete fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 in the family Phallaceae. It has a widespread distribution, and has been found in Africa, Australasia, and the Americas. It may have been introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 to North America with exotic plants. Similar to other stinkhorn fungi, the fruiting body, known as the receptaculum, starts out as a subterranean "egg" form. As the fungus develops, the receptaculum expands and erupts out of the protective 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...

, ultimately developing into mature structures characterized by two to five long vertical orange or red spongy columns, joined together at the apex. The fully grown receptaculum reaches heights of 8 cm (3.1 in) tall. The inside surfaces of the columns are covered with a fetid olive-brown spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

-containing slime, which attracts flies and other insects that help disseminate the spores. Although once considered undesirable, the fungus is listed as edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

. It is found commonly in mulch
Mulch
In agriculture and gardening, is a protective cover placed over the soil to retain moisture, reduce erosion, provide nutrients, and suppress weed growth and seed germination. Mulching in gardens and landscaping mimics the leaf cover that is found on forest floors....

.

Taxonomy and naming

The species was first named by the French botanist Louis Augustin Guillaume Bosc
Louis Augustin Guillaume Bosc
Louis Augustin Guillaume Bosc was a French botanist, invertebrate zoologist, and entomologist.-Biography:...

 in 1811. Christian Gottfried Daniel Nees von Esenbeck
Christian Gottfried Daniel Nees von Esenbeck
Christian Gottfried Daniel Nees von Esenbeck was a prolific German botanist, physician, zoologist, and natural philosopher. He was a contemporary of Goethe and was born within the lifetime of Linnaeus. He described approximately 7,000 plant species...

 transferred it into Laternea
Laternea
Laternea is a genus of fungi in the Phallaceae family. The genus was first described by French botanist Pierre Jean François Turpin in 1822...

in 1858, a genus intended to accommodate those Clathrus-like species with arms arranged in columns rather than a network; in its current meaning, Laternea includes species that have gleba suspended below the arch of the receptaculum by trabeculae (columns that extend from the peridium
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:...

 to the central core of the receptaculum). Other genera to which the species has been transferred include Linderia by Gordon Herriot Cunningham in 1932, Colonnaria by Eduard Fischer
Eduard Fischer
Major GeneralDr. h. c. Eduard von Fischer, 1862 - 1935,Knight of the Maria Theresa Order, was a Jewish colonel commanding the Austrian gendarmerie in Bukovina....

 in 1933, and Linderiella by Cunningham in 1942. Colonnaria, Linderia and Linderiella are now considered obsolete genera, as they have been subsumed into Clathrus
Clathrus
Clathrus is a genus of basidiomycete fungi of the family Phallaceae. As with other stinkhorn fungi, mature fruit bodies are covered with olive-brown slimy gleba, containing spores, which attracts flies...

.

The specific epithet columnatus is Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

, meaning "supported by pillars". The mushroom is commonly
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...

 known as the "column stinkhorn". Curtis Gates Lloyd
Curtis Gates Lloyd
Curtis Gates Lloyd was an American mycologist known for both his research on the Gasteromycetes, as well as his controversial views on naming conventions in taxonomy. He had a herbarium with over 59,000 fungal specimens, and published over a thousand new species of fungi...

 wrote in 1906 "in Florida, it is known to the natives as "Dead Men's Fingers."

Description

The fruit body
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

, or receptaculum, of Clathrus columnatus consists of two to five (usually four) spongy vertical columns, which are separate where they arise from the 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...

, but joined together at the top in an arch. The columns are joined together in pairs; the opposite pairs are joined by a short and broad arch similar in structure to the columns. The columns, which are narrower at the base than above, are reddish-orange above and yellowish-pink below. Young specimens have the fruit body compressed into the small interior space of an "egg", which consists of a peridium
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:...

 that is surrounded by a gelatinous layer that encloses the compressed fruiting body. The egg, usually gray or grayish-brown, typically reaches diameters of 3 to 5 cm (1.2 to 2 in) before the columns grow. The 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...

 remains at the base of the fruit body as a thick, loose, whitish sack. The mycelial cords found at the base of the volva are made of two types of tissues: a central bundle of fine 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 extend in a longitudinal direction, and an outer cortical
Cortex (botany)
In botany, the cortex is the outer layer of the stem or root of a plant, bounded on the outside by the epidermis and on the inside by the endodermis. It is composed mostly of undifferentiated cells, usually large thin-walled parenchyma cells of the ground tissue system. The outer cortical cells...

 layer of coarser hyphae that form a loose but highly interwoven structure. Full-grown columns can extend to 5 to 8 cm (2 to 3.1 in) above the ground, a location that optimizes spore dispersal. The fetid-smelling 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...

, the spore-bearing mass, is smeared on the upper inner surface of the columns.

The spores are elliptical, smooth, and have dimensions of 3.5–5 by 2–2.5 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...

. They have thin walls, and are covered by a transparent envelope.

Edibility

The words of William Gilson Farlow
William Gilson Farlow
William Gilson Farlow was an American botanist, born in Boston, Massachusetts, and educated at Harvard , where, after several years of European study, he became adjunct professor of botany in 1874 and professor of cryptogamic botany in 1879.In 1899 he was president of the American Society of...

, published in 1890, serve as a warning to those who might be inclined to consume Clathrus columnatus: "The odor of fully grown specimens of the order Phalloidea is so repulsive that the question as to their poisonous character when eaten by men has not often been the subject of experiment." Farlow described two cases of poisoning, one involving a young girl "who ate a small piece of the fungus, and was seized with violent convulsions followed by loss of speech and a deep sleep lasting 52 hours"; the other case involved hog
Domestic pig
The domestic pig is a domesticated animal that traces its ancestry to the wild boar, and is considered a subspecies of the wild boar or a distinct species in its own right. It is likely the wild boar was domesticated as early as 13,000 BC in the Tigris River basin...

s that ate the fungus found in patches in oak woods, and died 12–15 hours later. Despite this early report of poisoning, Orson K. Miller, Jr.
Orson K. Miller, Jr.
Orson Knapp Miller, Jr., born December 19, 1930, died June 9, 2006, is an American mycologist. He has published numerous papers in mycology and is responsible for the naming of many taxa, as well as being one of the authors erecting the genus Chroogomphus:...

 notes that the taste of the egg is mild, and lists the species as edible
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

.

Similar species

Pseudocolus fusiformis
Pseudocolus fusiformis
Pseudocolus fusiformis is a stinkhorn mushroom in the Phallaceae family, a family well-known for a remarkable range of fruit body types. It is the most widely distributed member of the genus Pseudocolus and has been found in the United States, Australia, Japan, Java, and the Philippines...

, Clathrus bicolumnatus
Clathrus bicolumnatus
Clathrus bicolumnatus is a species of fungus in the stinkhorn family. It is known only from Japan....

, and C. ruber
Clathrus ruber
Clathrus ruber is a species of fungus in the stinkhorn family, and the type species of the genus Clathrus. It is commonly known as the latticed stinkhorn, the basket stinkhorn, or the red cage, alluding to the striking fruit bodies that are shaped somewhat like a round or oval hollow sphere with...

have similarities to Clathrus columnatus. The lattice stinkhorn, C. ruber, has a larger, more globular, lattice-like receptacle. The "stinky squid", P. fusiformis, has arms that are attached at the bases, and free at the top. It grows on rotting logs and chip-mulched soil, in contrast to C. columnatus, which grows on sandy soil. C. bicolumnatus has a smaller stature (up to 9 cm tall), and only has two columns.

Development

The American botanist Edward Angus Burt published a detailed description of the development of C. columnatus in 1896. He found that the egg consists of cortical and medullary systems continued upward from the mycelial strand in the earliest stage. The cortical layer gives rise to the outer layer of the volva, the cortical plates and the pseudoparenchyma (thin-walled, usually angular, randomly-arranged cells that are tightly packed) of the receptaculum. The medullary portion gives rise to the gelatinous masses of the gelatinous layer of the volva, to the gleba, and to the gelatinous tissue of the chambers of the receptaculum. The elongation of the receptacle begins at the base and after its elongation, the gleba hangs suspended from the arch of the receptaculum by medullary tissue constituting the chamber masses of the receptacle.

Ecology, habitat and distribution

Like all Phallaceae species, C. columnatus is saprobic, and uses extracellular digestion to acquire nutrients from dead and decaying organic matter
Organic matter
Organic matter is matter that has come from a once-living organism; is capable of decay, or the product of decay; or is composed of organic compounds...

, like wood. As a consequence of its predilection for dead wood, the fungus is often associated with disturbed habitats. It can often be found growing in and around gardens and residences where areas of cultivation or landscaping have resulted in accumulations of mulch
Mulch
In agriculture and gardening, is a protective cover placed over the soil to retain moisture, reduce erosion, provide nutrients, and suppress weed growth and seed germination. Mulching in gardens and landscaping mimics the leaf cover that is found on forest floors....

, wood chips or other cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

-rich materials. The mycelial cords can be traced to buried roots, stumps, and other woody material. The species grows in sandy soil, near woody debris, in lawns, gardens, and cultivated soil. Fruit bodies appear singly, or scattered, and can arise in the summer, autumn, and early winter, especially after wet weather.

Like other member of the Phallaceae family, the mature fungus attracts insects with its smell to help disperse its spores. Psilopyga fasciata, a stinkhorn beetle of the sap beetle
Sap beetle
The sap beetles are a family of beetles.They are small ovoid, usually dull-coloured beetles, with knobbed antennae. Some have red or yellow spots or bands. They feed mainly on decaying vegetable matter, over-ripe fruit, and sap...

 family, has been recorded feeding on the gleba of Mexican specimens.

In 1980 Donald Malcolm Dring summarized the known geographical distribution of C. columnatus; the fungus has been collected in Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

, Oceania
Oceania
Oceania is a region centered on the islands of the tropical Pacific Ocean. Conceptions of what constitutes Oceania range from the coral atolls and volcanic islands of the South Pacific to the entire insular region between Asia and the Americas, including Australasia and the Malay Archipelago...

, New Guinea
New Guinea
New Guinea is the world's second largest island, after Greenland, covering a land area of 786,000 km2. Located in the southwest Pacific Ocean, it lies geographically to the east of the Malay Archipelago, with which it is sometimes included as part of a greater Indo-Australian Archipelago...

, Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

, and North
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

 and South America
South America
South America is a continent situated in the Western Hemisphere, mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere. The continent is also considered a subcontinent of the Americas. It is bordered on the west by the Pacific Ocean and on the north and east...

; According to Australian mycologist Tom May
Tom May (mycologist)
Thomas William May is a mycologist and the curator of the fungal collection at the National Herbarium of Victoria where he specialises in the taxonomy and ecology of Australian macrofungi. He is most notable for the comprehensive bibliographical lists of all Australian fungi published thus far;...

, the Australian distribution is "presumably erroneus", as it is based on only a single collection in 1948. In Central America, it has been collected from Costa Rica. It is also found in China, in Jiangsu
Jiangsu
' is a province of the People's Republic of China, located along the east coast of the country. The name comes from jiang, short for the city of Jiangning , and su, for the city of Suzhou. The abbreviation for this province is "苏" , the second character of its name...

, Fujian
Fujian
' , formerly romanised as Fukien or Huguing or Foukien, is a province on the southeast coast of mainland China. Fujian is bordered by Zhejiang to the north, Jiangxi to the west, and Guangdong to the south. Taiwan lies to the east, across the Taiwan Strait...

, and Guangdong
Guangdong
Guangdong is a province on the South China Sea coast of the People's Republic of China. The province was previously often written with the alternative English name Kwangtung Province...

. The North American distribution extends south to Mexico, and north to New York
New York
New York is a state in the Northeastern region of the United States. It is the nation's third most populous state. New York is bordered by New Jersey and Pennsylvania to the south, and by Connecticut, Massachusetts and Vermont to the east...

; it is also in Hawaii
Hawaii
Hawaii is the newest of the 50 U.S. states , and is the only U.S. state made up entirely of islands. It is the northernmost island group in Polynesia, occupying most of an archipelago in the central Pacific Ocean, southwest of the continental United States, southeast of Japan, and northeast of...

. The fungus is less common in the southeastern and southern United States. It is thought to have been introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

to North America, as it typically appear in landscaped areas or other locations where exotic plants have been established.
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