The Hygrophoraceae are a

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

fungi A fungus ( : fungi or funguses) is any member of the 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, separately from ...

in the order

Agaricales The fungal order Agaricales, also known as gilled mushrooms (for their distinctive gills) or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13,000 described species, alo ...

. Originally conceived as containing white-spored, thick-gilled

agarics An agaric () is a type of fungus fruiting body characterized by the presence of a pileus (cap) that is clearly differentiated from the stipe (stalk), with lamellae (gills) on the underside of the pileus. In the UK, agarics are called "mushrooms ...

(gilled mushrooms), including ''

Hygrophorus ''Hygrophorus'' is a genus of agarics (gilled mushrooms) in the family Hygrophoraceae. Called "woodwaxes" in the UK or "waxy caps" (together with ''Hygrocybe'' species) in North America, basidiocarps (fruit bodies) are typically fleshy, often wi ...

'' and ''

Hygrocybe ''Hygrocybe'' is a genus of agarics (gilled fungi) in the family (biology), family Hygrophoraceae. Called waxcaps in English (sometimes waxy caps in North America), basidiocarps (fruit bodies) are often brightly coloured and have dry to waxy pile ...

'' species (the waxcaps or waxy caps), DNA evidence has extended the limits of the family, so it now contains not only agarics, but also

basidiolichen Basidiolichens are lichenized members of the Basidiomycota, a much smaller group of lichens than the far more common ascolichens in the Ascomycota. In arctic, alpine, and temperate forests, the most common basidiolichens are in the agaric genus ...

s and

corticioid fungi The corticioid fungi are a group of fungi in the Basidiomycota typically having effused, smooth basidiocarps (fruit bodies) that are formed on the undersides of dead tree trunks or branches. They are sometimes colloquially called crust fungi or pa ...

. Species are thus diverse and are variously

ectomycorrhizal An ectomycorrhiza (from Greek ἐκτός ', "outside", μύκης ', "fungus", and ῥίζα ', "root"; pl. ectomycorrhizas or ectomycorrhizae, abbreviated EcM) is a form of symbiotic relationship that occurs between a fungal symbiont, or mycobi ...

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.mosses Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and horn ...

, or

saprotrophic Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi (f ...

. The family contains 25

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 ...

and over 600 species. None is of any great economic importance, though fruit bodies of some ''Hygrocybe'' and ''Hygrophorus'' species are considered edible and may be collected for sale in local markets.

Taxonomy

History

The family Hygrophoraceae was first proposed by Dutch botanist

Johannes Paulus Lotsy Johannes Paulus Lotsy or Jan Paulus Lotsy (11 April 1867 – 17 November 1931) was a Dutch botanist, specializing in evolution and heredity. He promoted the idea of evolution being driven by hybridization. Career Lotsy was born into a wealth ...

(1907) to accommodate

with thick, waxy

lamellae Lamella (plural lamellae) means a small plate or flake in Latin, and in English may refer to: Biology * Lamella (mycology), a papery rib beneath a mushroom cap * Lamella (botany) * Lamella (surface anatomy), a plate-like structure in an animal * ...

(gills) and white

spores 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 ...

. Lotsy's concept of the family included not only the waxcap-related genera ''

'', ''

'', ''Camarophyllus'' (= ''Hygrophorus''), and ''Godfrinia'' (= ''Hygrocybe''), but also ''

Gomphidius ''Gomphidius'' is a genus of mushrooms, commonly known as spike-caps, that are members of the Boletales ( suborder Suillineae), or pored fungi. They appear to have gill-like structures which resemble those of agarics, however the similarity is s ...

'' (despite its blackish spores) and ''Nyctalis'' (= '' Asterophora''). Not all subsequent authors accepted the Hygrophoraceae;

Carleton Rea Carleton Rea (7 May 1861 – 26 June 1946) was an English mycologist, botanist, and naturalist. Background and education Carleton Rea was born in Worcester, the son of the City Coroner. He was educated at The King's School and Magdalen Colle ...

(1922), for example, continued to place these genera within a widely defined

Agaricaceae The Agaricaceae are a family of basidiomycete fungi and include the genus ''Agaricus'', as well as basidiomycetes previously classified in the families Tulostomataceae, Lepiotaceae, and Lycoperdaceae. Taxonomy The family Agaricaceae was publishe ...

. In his major and influential revision of the Agaricales, however,

Rolf Singer Rolf Singer (June 23, 1906 – January 18, 1994) was a Germany, German-born mycologist and one of the most important Taxonomy (biology), taxonomists of gilled mushrooms (agarics) in the 20th century. After receiving his Ph.D. at the University ...

(1951) did accept the Hygrophoraceae, omitting ''Gomphidius'' and ''Nyctalis'', but including '' Neohygrophorus''. Singer's circumscription, with a few later additions, was followed by most authors until the 1990s. Thus the 1995 edition of the ''Dictionary of the Fungi'' listed '' Austroomphaliaster'', ''Bertrandia'' (=''Hygrocybe''), '' Camarophyllopsis'', '' Cuphophyllus'', ''

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''. The genus ''Porpolomopsis'' is closely related, and the species in it ...

'', ''

Hygroaster ''Hygroaster'' is a genus of mushroom-forming fungi in the family Hygrophoraceae. The genus was described by mycologist Rolf Singer in 1955. See also *List of Agaricales genera This is a list of mushroom-forming fungi genera in the order Agari ...

'', ''

'', ''Hygrotrama'' (= '' Camarophyllopsis''), ''Neohygrophorus'' (='' Pseudoomphalina''), and ''Pseudohygrocybe'' (=''Hygrocybe'') as genera of the Hygrophoraceae.

Cornelis Bas Dr. Cornelis (Kees) Bas (1928 – February 10, 2013) was a Dutch mycologist. Dr. Bas was born in Rotterdam and graduated in Biology at Leiden University in 1954. In 1953, he began working at the National Herbarium of the Netherlands, as cura ...

(1990), however, did not consider the group distinct, placing the hygrophoroid genera within the

Tricholomataceae The Tricholomataceae are a large family of mushrooms within the Agaricales. Originally a classic "wastebasket taxon", the family included any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to e.g. the ...

, a disposition followed by the next (2001) edition of the ''Dictionary of the Fungi''. In contrast,

Marcel Bon Marcel Bon (17 March 1925 – 11 May 2014)http://fmbds.org/wp-content/uploads/2014/06/CAFAM-2014-CR-complet.pdf was one of France's best known field mycologists. He was born in Picardy in 1925 and came to mycology through general botany, and ph ...

(1990) believed the Hygrophoraceae were so distinct, he placed the family in its own separate order, the Hygrophorales.

Current status

Recent

molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...

research, based on

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

analysis of

DNA sequences A nucleic acid sequence is a succession of bases signified by a series of a set of five different letters that indicate the order of nucleotides forming alleles within a DNA (using GACT) or RNA (GACU) molecule. By convention, sequences are usua ...

, suggests the Hygrophoraceae are distinct from the Tricholomataceae and are

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

(and hence a natural grouping). The genera '' Camarophyllopsis'' and '' Neohygrophorus'', however, do not belong within the family, but several other agaric and non-agaric genera do. The agaric genera include ''

Ampulloclitocybe ''Ampulloclitocybe'' is a genus of three species of fungi with a widespread distribution. See also *List of Agaricales genera This is a list of mushroom-forming fungi genera in the order Agaricales. Genera * See also *List of Agaricales famil ...

'', '' Cantharellula'', and ''

Lichenomphalia ''Lichenomphalia'' is both a basidiolichen and an agaric genus. Most of the species have inconspicuous lichenized thalli that consist of scattered, small, loose, nearly microscopic green balls or foliose small flakes containing single-celled gree ...

'', as well as the partly agaric, partly

cyphelloid The cyphelloid fungi are a group of fungi in the Basidiomycota that have disc-, tube-, or cup-shaped basidiocarps (fruit bodies), resembling species of discomycetes (or "cup fungi") in the Ascomycota. They were originally referred to the genus '' ...

genus '' Arrhenia''. The non-agaric genera include the

corticioid The corticioid fungi are a group of fungi in the Basidiomycota typically having effused, smooth basidiocarps (fruit bodies) that are formed on the undersides of dead tree trunks or branches. They are sometimes colloquially called crust fungi or pa ...

Eonema ''Eonema'' is a fungal genus in the family Hygrophoraceae. It is monotypic, consisting of the single corticioid species ''Eonema pyriforme''. This fungus was previously classified in the genus '' Athelia'' until molecular analysis demonstrated th ...

'' (formerly placed in '' Athelia'') and '' Cyphellostereum'', as well as the shelf-like

genera ''

Acantholichen ''Acantholichen'' is a fungal genus in the family Hygrophoraceae. The genus was circumscribed by Norwegian lichenologist Per Magnus Jørgensen to contain the type, and at that time, only species, the basidiolichen ''Acantholichen pannarioides'' ...

'', '' Cora'', '' Corella'' and ''

Dictyonema ''Dictyonema'' is a genus of mainly tropical basidiolichens in the family Hygrophoraceae. The ''Dictyonema'' symbiosis Most lichens are a symbiosis between an ascomycete fungus and a photosynthetic green alga. However, a small percentage of ...

''. As a result, the Hygrophoraceae as currently understood have no known morphological features in common that define them (

synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

Habitat, nutrition, and distribution

The majority of species in the Hygrophoraceae are ground-dwelling, though a few (such as '' Chrysomphalina'' species) occur on wood, or on mosses ('' Arrhenia'' species), or herbaceous stems ('' Eonema pyriforme''). Most are found in woodland, though (in Europe at least) ''

'' species are typical of

waxcap grassland Waxcap grassland is short-sward, nutrient-poor grassland that supports a rich assemblage of larger fungi, particularly waxcaps (species of '' Hygrocybe'' and related genera), characteristic of such habitats. Waxcap grasslands occur principally in ...

s. Species are nutritionally diverse. ''

'' species are

, typically forming associations with the roots of living trees. ''

'' species are now believed to be moss associates, as are some or all species of '' Arrhenia'' and '' Cantharellula''. Three genera, ''

'', ''

'', and '' Lichenomphalina'', are

s, forming associations with

algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...

and

cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...

. A few genera, such as ''

'' and ''

'', may be

. Members of the Hygrophoraceae are distributed worldwide, from the tropics to the subpolar regions. Over 400 species have been described to date.

Economic usage

Fruit bodies of some ''

'' and ''

'' species are edible and widely collected, sometimes being offered for sale in local markets. Examples of wild mushrooms collected and sold include '' Hygrophorus russula'', '' H. purpurascens'', '' H. chrysodon'', and '' H. hypothejus'' in Mexico, and '' H. eburneus'' and '' H. latitabundus'' in the

Spanish Pyrenees The Pyrenees (; es, Pirineos ; french: Pyrénées ; ca, Pirineu ; eu, Pirinioak ; oc, Pirenèus ; an, Pirineus) is a mountain range straddling the border of France and Spain. It extends nearly from its union with the Cantabrian Mountains to ...

. '' Hygrophorus gliocyclus'' was used as food by the St'at'imc and

Nlaka'pamux The Nlaka'pamux or Nlakapamuk ( ; ), also previously known as the ''Thompson'', ''Thompson River Salish'', ''Thompson Salish'', ''Thompson River Indians'' or ''Thompson River people'', and historically as the ''Klackarpun'', ''Haukamaugh'', ''Kni ...

people of Canada.Dugan (2011), p. 88. None is cultivated commercially. File:2010-08-11 Hygrophorus erubescens 99991.jpg, '' Hygrophorus erubescens'' File:Dictyonema pavonium.jpg, '' Dictyonema pavonium'' File:Arrheniasphatulata2.jpg, '' Arrhenia spathulata'' File:Humidicutis lewelliniae 1 smaller.jpg, '' Humidicutis lewelliniae''

References

Cited literature

* * {{Taxonbar, from=Q596145 Agaricales families Lichen families Taxa named by Johannes Paulus Lotsy Taxa described in 1907