Solitary Dog Sculptor I

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 interlaced or latticed branches. The fungus is saprobic, feeding off decaying woody plant material, and is usually found alone or in groups in leaf litter on garden soil, grassy places, or on woodchip garden mulches. Although considered primarily a European species, C. ruber has been introduced to other areas, and now has a wide distribution that includes northern Africa, Asia, Australia, and North and South America. The species was illustrated in the scientific literature during the 1500s, but was not officially described until 1729.

The fruit body initially appears like a whitish "egg" attached to the ground at the base by cords called rhizomorphs. The egg has a delicate, leathery outer membrane enclosing the compressed lattice that surrounds a layer of olive-green spore-bearing slime called the gleba, which contains high levels of calcium that help protect the developing fruit body during development. As the egg ruptures and the fruit body expands, the gleba is carried upward on the inner surfaces of the spongy lattice, and the egg membrane remains as a volva around the base of the structure. The fruit body can reach heights of up to 20 cm (7.9 in). The color of the fruit body, which can range from pink to orange to red, results primarily from the carotenoid pigments lycopene and beta-carotene. The gleba has a fetid odor, somewhat like rotting meat, which attracts flies and other insects to help disperse its spores. Although the edibility of the fungus is not known with certainty, its odor would deter most from consuming it. C. ruber was not regarded highly in tales in southern European folklore, which suggested that those who handled the mushroom risked contracting various ailments.


Taxonomy, phylogeny, and naming

Clathrus ruber was illustrated as early as 1560 by the Swiss naturalist Conrad Gesner in his Nomenclator Aquatilium Animantium—Gesner mistook the mushroom for a marine organism.[4] It appeared in a woodcut in John Gerard's 1597 Great Herball,[5] shortly thereafter in Carolus Clusius '​ 1601 Fungorum in Pannoniis Observatorum Brevis Historia,[6] and was one of the species featured in Cassiano dal Pozzo's museo cartaceo ("paper museum") that consisted of thousands of illustrations of the natural world.[7]

The fungus was first described scientifically in 1729, by the Italian Pier Antonio Micheli in his Nova plantarum genera iuxta Tournefortii methodum disposita, who gave it its current scientific name.[8] The species was once referred to by American authors as Clathrus cancellatus L., as they used a system of nomenclature based on the former American Code of Botanical Nomenclature, in which the starting point for naming species was Linnaeus's 1753 Species Plantarum. The International Code for Botanical Nomenclature now uses the same starting date, but names of Gasteromycetes used by Christian Hendrik Persoon in his Synopsis Methodica Fungorum (1801) are sanctioned and automatically replace earlier names. Since Persoon used the specific epithet ruber, the correct name for the species is Clathrus ruber. Several historical names of the fungus are now synonyms: Clathrus flavescens, named by Persoon in 1801;[9] Clathrus cancellatus by Joseph Pitton de Tournefort and published by Elias Fries in 1823;[10] Clathrus nicaeensis, published by Jean-Baptiste Barla in 1879;[11] and Clathrus ruber var. flavescens, published by Livio Quadraccia and Dario Lunghini in 1990.[12][13]

Clathrus ruber is the type species of the genus Clathrus, and is part of the group of Clathrus species known as the Laternoid series. Common features uniting this group include the vertical arms of the receptacle (fruit body) that are not joined together at the base, and the spongy structure of the receptacle.[14] According to a molecular analysis published in 2006, out of the about 40 Phallales species used in the study, C. ruber is most closely related to Aseroe rubra, Clathrus archeri, Laternea triscapa, and Clathrus chrysomycelinus.[3]

The generic name Clathrus is derived from Ancient Greek κλειθρον or "lattice", and the specific epithet is Latin ruber, meaning "red".[15] The mushroom is commonly known as the "basket stinkhorn",[16] the "lattice stinkhorn",[17] or the "red cage".[18] It was known to the locals of the Adriatic hinterland in the former Yugoslavia as veštičije srce or vještičino srce, meaning "witch's heart".[19]


Description

Before the volva opens, the fruiting body is egg-shaped to roughly spherical, up to 6 cm (2.4 in) in diameter, with a gelatinous interior up to 3 mm (0.1 in) thick. White to grayish in color, it is initially smooth, but develops a network of polygonal marks on the surface prior to opening as the internal structures expand and stretch the peridium taut.[20] The fruit body, or receptacle, bursts the egg open as it expands (a process that can take as little as a few hours),[6] and leaves the remains of the peridium as a cup or volva surrounding the base.[20] The receptacle ranges in color from red to bright pink to pale orange, and it is often lighter in color approaching the base. The color appears to be dependent upon the temperature and humidity of the environment.[21] The receptacle consists of a spongy network of "arms" interlaced to make meshes of unequal size. At the top of the receptacle, the arms are up to 1.5 cm (0.6 in) thick, but they taper down to smaller widths near the base. A cross-section of the arm reveals it to be spongy, and made up of one wide inner tube and two indistinct rows of tubes towards the outside. The outer surface of the receptacle is ribbed or wrinkled.[20] There are between 80 and 120 mesh holes in the receptacle.[22] The unusual shape of the receptacle has inspired some creative comparisons: David Arora likened it to a whiffleball,[21] while the German Mycological Society—who named C. ruber the 2011 "Mushroom of the Year"—described it as "like an alien from a science fiction horror film".[23]

A considerable variation in height has been reported for the receptacle, ranging from 8 to 20 cm (3.1 to 7.9 in) tall.[6] The base of the fruit bodies are attached to the substrate by rhizomorphs (thickened cords of mycelia). The dark olive-green to olive-brown, foul-smelling sticky gleba covers the inner surface of the receptacle, except near the base. The odor—described as resembling rotting meat[24][25]—attracts flies, other insects, and, in one report, a scarab beetle (Scarabaeus sacer)[26] that help disperse the spores.[21][27] The putrid odor—and people's reaction to it—have been well documented. In 1862 Mordecai Cubitt Cooke wrote "it is recorded of a botanist who gathered one for the purpose of drying it for his herbarium, that he was compelled by the stench to rise during the night and cast the offender out the window."[28] American mycologist David Arora called the odor "the vilest of any stinkhorn".[21] The receptacle collapses about 24 hours after its initial eruption from the egg.[6]

The spores are elongated, smooth, and have dimensions of 4–6 by 1.5–2 µm.[20] Scanning electron microscopy has revealed that C. ruber (in addition to several other Phallales species) has a hilar scar—a small indentation in the surface of the spore where it was previously connected to the basidium via the sterigma.[29] The basidia (spore-bearing cells) are six-spored.[30]


Similar species

Clathrus ruber may be distinguished from the closely related tropical species C. crispus by the absence of the corrugated rims which surround each mesh of the C. crispus fruit body.[31] The phylogenetically close species C. chrysomycelinus has a yellow receptacle with arms that are structurally simpler, and its gleba is concentrated on specialized "glebifers" located at the lattice intersections. It is known only from Venezuela to southern Brazil.[20] Clathrus columnatus has a fruit body with two to five long vertical orange or red spongy columns, joined together at the apex.[32]


Edibility and folklore

Although edibility for C. ruber has not been officially documented,[33] its foul smell would dissuade most individuals from consuming it. In general, stinkhorn mushrooms are considered edible when still in the egg stage, and are even considered delicacies in some parts of Europe and Asia, where they are pickled raw and sold in markets as "devil's eggs".[21] However, an 1854 report provides a cautionary tale to those considering consuming the mature fruit body. Dr. F. Peyre Porcher, of Charleston, South Carolina, described an account of poisoning caused by the mushroom:

    "A young person having eaten a bit of it, after six hours suffered from a painful tension of the lower stomach, and violent convulsions. He lost the use of his speech, and fell into a state of stupor, which lasted for forty-eight hours. After taking an emetic he threw up a fragment of the mushroom, with two worms, and mucus, tinged with blood. Milk, oil, and emollient fomentations, were then employed with success."[34]

British mycologist Donald Dring, in his 1980 monograph on the Clathraceae family, wrote that C. ruber was not regarded highly in southern European folklore. He mentions a case of poisoning following its ingestion, reported by Barla in 1858, and notes that Ciro Pollini reported finding it growing on a human skull in a tomb in a deserted church.[20] According to John Ramsbottom, Gascons consider the mushroom a cause of cancer;[35] they will usually bury specimens they find.[6] In other parts of France it has been reputed to produce skin rashes or cause convulsions.[35]


Ecology, habitat, and distribution

Like most of the species of the Phallales order, Clathrus ruber is saprobic—a decomposer of wood and plant organic matter—and is commonly found fruiting in mulch beds.[36] The fungus grows alone or clustered together near woody debris, in lawns, gardens, and cultivated soil.[33]

Clathrus ruber was originally described by Micheli from Italy. It is considered native to southern and central continental Europe, as well as Macaronesia (the Azores[20] and the Canary Islands[37]), western Turkey,[38] North Africa (Algeria), and western Asia (Iran).[20] The fungus is rare in central Europe,[19] and is listed in the Red data book of Ukraine.[39]

The fungus has probably been introduced elsewhere, often because of the use of imported mulch used in gardening and landscaping.[33] It may have extended its range northwards into the British Isles or been introduced in the nineteenth century.[40] It now has a mainly southerly distribution in England and has been recorded from Cornwall,[41] Devon,[42] Dorset, Somerset,[20] the Isle of Wight,[43] Hampshire, Sussex, Surrey, and Middlesex. In Scotland, it has been recorded from Argyll. It is also known from Wales, the Channel Islands,[20] and Ireland.[44] The fungus also occurs in the United States (California, Florida, Georgia, Virginia, North Carolina, and New York),[45] Canada, Mexico, and Australasia.[46] The species was also reported from South America (Argentina).[47] In China, it has been collected from Guangdong, Sichuan, Guizhou, and Tibet.[22] Records from Japan[48] are referable to Clathrus kusanoi; records from the Caribbean are probably of C. crispus.[20]


Biochemistry

Like other stinkhorn fungi, C. ruber bioaccumulates the element manganese. It has been postulated that this element plays a role in the enzymatic breakdown of the gleba with simultaneous formation of odorous compounds. Compounds like dimethyl sulfide, aldehydes, and amines—which contribute to the disagreeable odor of the gleba—are produced by the enzymatic decarboxylation of keto acids and amino acids, but the enzymes will only work in the presence of manganese.[6] A chemical analysis of the elemental composition of the gelatinous outer layer, the embryonic receptacle and the gleba showed the gelatinous layer to be richest in potassium, calcium, manganese, and iron ions. Calcium2+ stabilizes the polysaccharide gel, protecting the embryonic receptacle from drying out during the growth of the egg. Potassium is required for the gelatinous layer to retain its osmotic pressure and retain water; high concentrations of the element are needed to support the rapid growth of the receptacle. The high concentration of elements suggests that the gelatinous layer has a "placenta-like" function—serving as a reservoir from which the receptacle may draw upon as it rapidly expands.[6]

Pigments responsible for the orange to red colors of the mature fruit bodies have been identified as carotenes, predominantly lycopene and beta-carotene—the same compounds responsible for the red and orange colors of tomatoes and carrots, respectively. Lycopene is also the main pigment in the closely related fungus Clathrus archeri, while beta-carotene is the predominant pigment in the Phallaceae species Mutinus caninus, M. ravenelii, and M. elegans.[49]


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Clathrus ruber es una especie saprófita de hongos de la familia de las falláceas. A menudo se lo conoce como "jaula roja", en referencia a su cuerpo que toma la forma de una pelota oval con un enrejado de ramas entrelazadas. Cubierto con una membrana mucosa en sus superficies internas, esta especie no-comestible posee un olor fétido, descripto como de carne en descomposición,1 con lo que atrae moscas y otros insectos que le ayudan a dispersar sus esporas.2


Descripción

Antes de que se abra la volva, el cuerpo posee una forma de huevo con un interior gelatinoso, y un color blancuzco. Luego de que se abre, se convierte en un receptáculo rojo o anaranjado que consiste en una malla esponjosa. Se ha observado una significativa variación en la altura de esta especie, que alcanza entre 8 a 20 cm.3 La gleba oscura y de olor fétido recubre la superficie interior del receptáculo y la zona basal del receptáculo se encuentra rodeada de una volva blanca con una central mycelial cord. Las esporas son alargadas, suaves, sus dimensiones son 5–6 x 1.7–2 µm.

Esta especie se diferencia de la especie tropical Clathrus crispin por la falta de laminillas corrugadas que rodean cada reticulado del esporocarpio del C. crispin.4


Hábitat

Este fungus crece aislado o en grupos en proximidad de residuos de madera, en praderas de césped, jardines, y tierra cultivada.5
Comestibilidad

Si bien no se ha documentado en forma oficial si el C. ruber es o no comestible,6 su olor fétido disuade a la mayoría de las personas de consumirlo.

En 1854 el Dr. F. Peyre Porcher, de Charleston, Carolina del Sur, Estados Unidos escribió un relato sobre un envenenamiento con esta especie:

    "Luego de que una persona joven comió un poco de este hongo, a las seis horas se quejó de fuertes dolores en el bajo estómago, y tuvo convulsiones violentas. Perdió el habla, y cayó en un estado de sopor, que duró 48 horas. Luego de administrale un hemetico devolvio arrojando un fragmento del hongo, con dos lombrices, y mucus, con trazas de sangre. Luego se utilizaron con éxito fomentos con leche, aceite y emolientes."7


Taxonomía

A menudo esta especie ha sido descrita por autores norteamericanos como C. cancellatus Linnaeus, basándose en el Código de Nomenclatura Botánica Norteamericano, el cual está basado en el Species plantarum, 1753 de Linnaeus. De acuerdo al International Code for Botanical Nomenclature, en 1801 se comenzó con la nomenclatura de los Gasteromicetos, y la Synopsis methodica fungorum de Christian Hendrik Persoon fue publicada en 1801. Por lo tanto el epíteto específico correcto de esta especie es Clathrus ruber.
Distribución

El C. ruber se encuentra en Estados Unidos, Australasia,8 el sureste y suroeste de Europa,9 Islas Canarias,10 Islas del Caribe, las Azores, Canadá, México, Japón,11 France,12 y el Reino Unido (Isla de Wight,13 Cornwall,14 Devon,15 ) e Irlanda.16 En Estados Unidos se ha informado sobre el hallazgo de C. ruber en California, Florida, Georgia, Virginia, Carolina del Norte y Nueva York. También se lo ha encontrado en Córdoba , República Argentina. C. ruber aparece en el libro rojo de Ucrania.17


Bioquímica

El C. ruber al igual que lo que sucede en otros hongos con olores desagradables bioacumula el elemento manganeso. Se cree que este elemento juega algún rol en la descomposición enzimática de la gleba con la formación en este proceso de compuestos con olor.3 Se han realizado análisis de elementos de la capa exterior gelatinosa, el receptáculo embriónico y la gleba, y la capa gelatinosa, los mismos indican importantes contenidos de potasio, calcio, manganeso y hierro. El calcio estabiliza el gel polisacárido que protege el carpoforo embriónico (the stalk of the esporocarpo) de que se seque durante el desarrollo del huevo. Las concentraciones elevadas de otros elementos (comparadas con las del carpoforo en crecimiento) sugiere que la capa gelatinosa posee una función similar a la de una placenta.3

Los pigmentos responsables de las coloraciones naranjas y rojas de los frutos maduros son carotenos, predominantemente licopeno y beta caroteno.18 19


Complete en:
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