Hygrocybe
Encyclopedia
Hygrocybe is a genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 of agarics (gilled fungi) in the family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Hygrophoraceae
Hygrophoraceae
The Hygrophoraceae are a family of fungi in the order Agaricales. Originally conceived as containing white-spored, thick-gilled agarics , including Hygrophorus and Hygrocybe species , DNA evidence has extended the limits of the family, so that it now contains not only agarics but also...

. Called "waxcaps" in English (sometimes "waxy caps" in North America), basidiocarp
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...

s (fruit bodies) are often brightly coloured and have waxy to slimy 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...

s, white 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...

s, and smooth, ringless stems
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...

. In Europe they are characteristic of old, unimproved grasslands (termed waxcap grasslands
Waxcap grassland
Waxcap grassland is a term used to describe short-sward, nutrient-poor grasslands supporting rich assemblages of larger fungi, particularly waxcaps , characteristic of such habitats. Waxcap grasslands occur principally in Europe, where they are declining as a result of agricultural practices...

) which are a declining habitat, making many Hygrocybe species of conservation concern. Elsewhere they are more typically found in woodlands. Most are ground-dwelling and all are believed to be moss associates. Around 150 species are currently recognized worldwide. Fruit bodies of several Hygrocybe species are considered edible and are sometimes offered for sale in local markets.

History

Hygrocybe was first published in 1821 by Swedish mycologist Elias Magnus Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

 as a subsection of Agaricus
Agaricus
Agaricus is a large and important genus of mushrooms containing both edible and poisonous species, with possibly over 300 members worldwide...

and in 1871 was raised to the rank of genus by Kummer
Paul Kummer
Paul Kummer was a priest, teacher, and scientist in Zerbst, Germany, known chiefly for his contribution to mycological nomenclature. Earlier classification of agarics by pioneering fungal taxonomist Elias Magnus Fries designated only a very small number of genera, with most species falling into...

. In several papers, Karsten
Petter Adolf Karsten
Petter Adolf Karsten was a Finnish mycologist, the foremost expert on the fungi of Finland in his day, and known in consequence as the "father of Finnish mycology"....

 and Murrill
William Murrill
William Alphonso Murrill was an American mycologist, known for his contributions to the knowledge of the Agaricales and Polyporaceae.- Education :...

 used the name Hydrocybe, but this is now taken as an orthographic variant of Hygrocybe. The generic name is derived from the Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 ῦγρὁς (= moist) + κυβη (= head).

Despite its comparatively early publication, the genus Hygrocybe was not widely accepted until the 1970s, most previous authors treating it as a synonym of Hygrophorus
Hygrophorus
Hygrophorus is a genus of agarics in the family Hygrophoraceae. Called "woodwaxes" in the UK or "waxy caps" in North America, basidiocarps are typically fleshy, often with slimy caps and lamellae that are broadly attached to decurrent...

, a related genus of ectomycorrhizal agarics.

Hygrocybe itself has been split into subgenera, several of which – notably Cuphophyllus (= Camarophyllus sensu Singer
Rolf Singer
Rolf Singer was a German-born mycologist and one of the most important taxonomists of gilled mushrooms in the 20th century....

, non Fries
Elias Magnus Fries
-External links:*, Authors of fungal names, Mushroom, the Journal of Wild Mushrooming.*...

) – have subsequently been raised to generic rank. As of 2011, however, most standard authorities place these split genera in synonymy with Hygrocybe. Some species, such as the mauve splitting waxcap (Humidicutis lewelliniae
Humidicutis lewelliniae
Humidicutis lewelliniae, commonly known as the mauve splitting wax-cap, is a gilled fungus of the waxcap family found in wet forests of eastern Australia and New Zealand. The small mauve- or lilac-coloured mushrooms are fairly common and appear in moss or leaf litter on the forest floor in autumn,...

), have been described in the small genus Humidicutis
Humidicutis
Humidicutis is a small genus of brightly coloured agarics, the majority of which are found in Eastern Australia. They were previously described as members of Hygrocybe, such as the Mauve splitting wax-cap Humidicutis lewelliniae of eastern Australia and Malaysia.The generic name derives from the...

.

Current status

Recent molecular research, based on cladistic analysis of DNA sequences, suggests that Hygrocybe as currently understood is paraphyletic and does not form a single clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 within the Hygrophoraceae. As a result, it has been suggested that at least the genus Cuphophyllus (comprising Hygrocybe pratensis and its allies) should be recognized and removed from Hygrocybe sensu stricto, together with the genus Gliophorus
Gliophorus
Gliophorus is a genus of agaric fungi in the family Hygrophoraceae. Gliophorus species belong to a group known as waxcaps in English, sometimes also waxy caps in North America or waxgills in New Zealand. The genus was described by Czech mycologist Josef Herink in 1958, but is currently synonymized...

(comprising Hygrocybe psittacina
Hygrocybe psittacina
Hygrocybe psittacina, commonly known as the Parrot Toadstool or Parrot Waxcap, is a colourful member of the genus Hygrocybe, the waxcaps, found across Northern Europe. Originally described as Hygrophorus psittacinus, there have been at least two subspecies described, H. psittacina var. psittacina...

and its allies). As yet, however, only a few species of Hygrocybe have been sequenced and though these suggestions have been accepted in principle (e.g. by Boertmann, 2010), they have not yet been widely implemented.

Description

Fruit bodies of Hygrocybe species are all agaricoid
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...

, most (but not all) having smooth to slightly scaly caps that are convex to conical and waxy to slimy when damp. Many (but not all) are brightly coloured in shades of red, orange, or yellow – less commonly pink or green. Where present, the gills beneath the cap are often equally coloured and usually distant, thick, and waxy. One atypical South American species, Hygrocybe aphylla, lacks gills. The stems of Hygrocybe species lack a ring
Annulus (mycology)
An annulus is the ring like structure sometimes found on the stipe of some species of mushrooms. The annulus represents the remaining part of the partial veil, after it has ruptured to expose the gills or other spore-producing surface. An annulus may be thick and membranous, or it may be cobweb-like...

. The spore print
Spore print
thumb|300px|right|Making a spore print of the mushroom Volvariella volvacea shown in composite: mushroom cap laid on white and dark paper; cap removed after 24 hours showing pinkish-tan spore print...

 is white. Fruit bodies of some species, notably Hygrocybe conica
Hygrocybe conica
Hygrocybe conica, commonly known as the witch's hat, conical wax cap or conical slimy cap, is a colourful member of the genus Hygrocybe , found across northern Europe and North America...

, blacken with age or when bruised. Microscopically, Hygrocybe species lack true cystidia and have comparatively large, smooth, inamyloid
Amyloid (mycology)
In mycology the term amyloid refers to a crude chemical test using iodine in either Melzer's reagent or Lugol's solution, to produce a black to blue-black positive reaction. It is called amyloid because starch gives a similar reaction, and that reaction for starch is also called an amyloid reaction...

 basidiospores.

Habitat, nutrition, and distribution

Most species of Hygrocybe are ground-dwelling, though a few (such as Hygrocybe mexicana and H. rosea) are only known from mossy tree trunks or logs. In Europe, species are typical of unimproved (nutrient-poor), short-sward grasslands, often termed "waxcap grasslands
Waxcap grassland
Waxcap grassland is a term used to describe short-sward, nutrient-poor grasslands supporting rich assemblages of larger fungi, particularly waxcaps , characteristic of such habitats. Waxcap grasslands occur principally in Europe, where they are declining as a result of agricultural practices...

", but elsewhere they are more commonly found in woodland.

Their nutritional status has long been debated, but recent research shows that they are neither mycorrhizal nor saprotrophic. It seems they may be symbiotically
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 associated with mosses, as suggested by several earlier authors.

Species are distributed worldwide, from the tropics to the sub-polar regions. Around 150 have been described to date.

Conservation

In Europe, waxcap grasslands and their associated fungi are of conservation concern, since unimproved grasslands (formerly commonplace) have declined dramatically as a result of changes in agricultural practice. As a result, by 1993, 89% of European Hygrocybe species appeared on one or more national red lists of threatened fungi. In several countries, action has been taken to conserve waxcap grasslands and some of the rarer Hygrocybe species. In the United Kingdom, some grasslands have gained a measure of legal protection as Sites of Special Scientific Interest because of their waxcap interest.

Economic usage

Fruit bodies of one of the commoner European waxcap species, Hygrocybe pratensis, are edible and widely collected, sometimes being offered for sale in local markets. Because Hygrocybe species cannot be maintained in culture, none is cultivated commercially. Fruit bodies of a few additional species are considered edible in eastern Europe, south-east Asia, and Central America and are collected and consumed locally.

Literature

No comprehensive monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...

 of the genus has yet been published. In Europe, however, species of Hygrocybe have been illustrated and described in a standard English-language guide by Boertmann (2010) and also (together with Hygrophorus) in an Italian guide by Candusso (1997). European species have also been covered, more briefly, in descriptive French keys by Bon (1990). Dutch species were illustrated and described by Arnolds (1990). No equivalent modern guides have been published for North America, the most recent being by Hesler
Lexemuel Ray Hesler
-External links:* Finding Aid for the Lexemuel Ray Hesler Collection, 1899-1982...

 & Smith
Alexander H. Smith
Alexander Hanchett Smith was an American mycologist known for his extensive contributions to the taxonomy and phylogeny of the higher fungi, especially the agarics.-Early life:...

 (1963). There is, however, a guide to Californian species by Largent (1985). In Australia, Hygrocybe species have been illustrated and described by Young (2005) and in New Zealand by Horak (1990).

External links

  • WaxCap Website, University of Wales, Aberystwyth
    University of Wales, Aberystwyth
    Aberystwyth University is a university located in Aberystwyth, Wales. Aberystwyth was a founding Member Institution of the former federal University of Wales. As of late 2006, the university had over 12,000 students spread across seventeen academic departments.The university was founded in 1872 as...

    .
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