Russulaceae

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Russulaceae
Russula emetica 117475.jpg
Russula emetica
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Basidiomycota
Class: Agaricomycetes
Order: Russulales
Family: Russulaceae
Lotsy (1907) [1]
Type genus
Russula
Pers. (1796)
Genera

1including Arcangeliella , Gastrolactarius , and Zelleromyces
2including Cystangium , Gymnomyces, Elasmomyces, Martellia, and Macowanites

Contents

Synonyms [2] [3]

The Russulaceae are a diverse family of fungi in the order Russulales, with roughly 1,900 known species and a worldwide distribution. They comprise the brittlegills and the milk-caps, well-known mushroom-forming fungi that include some edible species. These gilled mushrooms are characterised by the brittle flesh of their fruitbodies.

In addition to these typical agaricoid forms, the family contains species with fruitbodies that are laterally striped (pleurotoid), closed (secotioid or gasteroid), or crust-like (corticioid). Molecular phylogenetics has demonstrated close affinities between species with very different fruitbody types and has discovered new, distinct lineages.

An important group of root-symbiotic ectomycorrhizal fungi in forests and shrublands around the world includes Lactifluus , Multifurca , Russula , and Lactarius . The crust-forming genera Boidinia , Gloeopeniophorella , and Pseudoxenasma , all wood-decay fungi, have basal positions in the family.

Systematics and taxonomy

The family Russulaceae was first validly named in 1907 by Dutch botanist Johannes Paulus Lotsy, [4] who included three genera: Russula , Lactarius , and Russulina (now considered a synonym of Russula). He emphasised features such as the granular flesh, thick gills, spiny spores, and milky hyphae and rounded cells (sphaerocytes). [1] A prior usage of "Russulariées" by French mycologist Ernst Roze in 1876 [5] is not considered a valid publication, since the proper Latin termination for the family rank specified in article 18.4 of the nomenclature code was not used. [4] [6]

Synonyms of Russulaceae include: Ernst Albert Gäumann's Lactariaceae (1926), Fernand Moreau's Asterosporaceae (1953), [2] and David Pegler and Thomas Young's Elasmomycetaceae (1979). The latter family was proposed to contain species with statismosporic (non-forcibly discharged) and symmetric spores, including the gasteroid genera Elasmomyces , Gymnomyces , Martellia , and Zelleromyces . Calonge and Martín reduced the Elasmomycetaceae to synonymy with the Russulaceae when molecular analysis confirmed the close genetic relationship between the gasteroid and agaricoid genera. [3]

Placement of the family

Historically, the gilled mushrooms of the family Russulaceae were classified with other gilled species in the order Agaricales, [7] but microscopical studies of spore and fruitbody flesh features raised the possibility that they were more closely related with certain "lower fungi" presenting nongilled, crust-like fruitbodies. [2] [8] [9] The use of molecular phylogenetics confirmed that these morphologically diverse fungi form a distinct lineage, first termed the "russuloid clade" [10] [11] and today classified as order Russulales in the class Agaricomycetes. [12] The family's sister group within the order appears to be the crust-like Gloeocystidiellaceae. [13]

Internal systematics

Lactarius

Multifurca

Russula

Lactifluus

Boidinia

Gloeopeniophorella

Pseudoxenasma

Phylogeny of the Russulaceae; dashed lines indicate uncertain placement. [10] [13] [14] [15]

A 2008 molecular phylogenetic study clarified the relationships among the mushroom-forming species of the family. [15] The authors demonstrated the existence of four distinct lineages of gilled mushrooms, which led to the description of Multifurca as a new genus separated from Russula [15] and the segregation of Lactifluus from Lactarius. [16] [17]

Genera with closed fruitbodies within the family are form taxa instead of natural groups: Arcangeliella, Gastrolactarius, and Zelleromyces are phylogenetically part of Lactarius, while Cystangium, Elasmomyces, Gymnomyces, Macowanites, and Martellia belong to Russula. [3] [16] Nevertheless, some of these genus names are still in use, as many of the concerned species have not yet formally been synonymised with Lactarius or Russula. [18]

The crust-like genera Boidinia , Gloeopeniophorella , and Pseudoxenasma , formerly placed in the Corticiaceae or Gloeocystidiellaceae, are now classified in the Russulaceae and basal to the clade of mushroom-forming species described above. [13] [18] Studies have so far failed to clearly circumscribe and place these genera within the family. [10] [13] [14] Boidinia in its current extent is polyphyletic, with some species not falling into the Russulaceae. [10]

Species diversity

Altogether, the Russulaceae comprise around 1,900 accepted species. [18] Russula is by far the largest genus with c. 1100 species, Lactarius has c. 550, Lactifluus c. 120, Boidinia 13, Multifurca 6, Gloeopeniophorella 6, and Pseudoxenasma 1 species. [18] [19] Closed-fruitbody species not yet synonymised with Lactarius or Russula (see above) account for some 150 species. [18]

New species in the Russulaceae continue to be described from various regions, such as the US, [20] Guyana, [21] Brazil, [22] Patagonia, [23] Togo, [24] Sri Lanka, [25] or Thailand. [25] It has been estimated that the real number of Russula species in North America alone (currently around 400 described) might be as high as 2000. [26] Cryptic species may increase true diversity: some morphologically well-defined species, especially in Lactifluus, have been shown to actually encompass several phylogenetic species. [27] [28] [29]

Description

Macroscopic characteristics

Milchlinge-Fleisch.jpg
Lactifluus vellereus (Russulaceae)
Pilzfleisch.jpeg
Suillus variegatus (Suillaceae)
Comparison of brittle flesh texture in Russulaceae and the more fibrous flesh found in most other fungi

Three major types of fruitbodies occur in the Russulaceae: agaricoid and pleurotoid forms with a cap, gills, and a stipe; forms with closed ( gasteroid ) or partially closed ( secotioid ) fruitbodies, and corticioid , crust-like forms.

The agaricoid species in Lactarius, Lactifluus, Multifurca, and Russula are readily distinguished from other gilled mushrooms by the consistency of their flesh, which is granular, brittle and breaks easily, somewhat like a piece of chalk. [1] [30] Russulaceae never have a volva, [30] but a partial veil can be found in some tropical species. [31] [32] Gills are adnate to decurrent, and the colour of the spore print ranges from white to ochre or orange [15] [33] (with the brown-spored Lactarius chromospermus as an exception [34] ).

Caps can be dull to very colourful, the latter especially in Russula; [33] their size ranges from 17 mm diameter or less in Russula campinensis [35] to 30 cm (12 in) in Lactifluus vellereus . [33] Concentrically ringed (zonate) caps occur in all Multifurca [15] and several Lactarius species. [33] Laterally striped (pleurotoid) fruitbodies exist in some, mainly tropical Lactifluus and Russula species. [35] [36] [37] [38] Taste is a distinguishing characteristic in many species, from mild to very acrid. [33] A conspicuous feature of the "milk-caps" in Lactarius, Lactifluus, and Multifurca furcata is the latex or "milk" their fruitbodies exude when bruised. [15] [30]

The secotioid and gasteroid species in Lactarius and Russula are derived from agaricoid forms. [15] [39] Secotioid species still have a stipe but the cap does not open fully, while in gasteroid species, fruitbodies are completely closed and the stipe is reduced; in both cases, the spore-bearing structure is made up of convoluted gills that are more or less crowded and anastomosed. [39] These closed-fruitbody species represent a continuum of secotioid to gasteroid, above-ground to below-ground fruitbodies, with spores forcibly discharged or not. [25] [39] [40] Secotioid or gasteroid Lactarius exude latex just like their agaricoid relatives. [3] [25] [40]

The corticioid species of Boidinia, Gloeopeniophorella, and Pseudoxenasma develop crust-like fruitbodies with a smooth, porous, or flaky surface and grow on tree logs or dead branches. [10] [41] [42] [43]

Morphological variety of Russulaceae
*These species belong phylogenetically to Lactarius.

Microscopic characteristics

Spores of Lactarius rubidus, showing ornamentation with a blue, amyloid stain reaction that is typical for Russulaceae Lactarius rubidus spores 1000x.JPG
Spores of Lactarius rubidus , showing ornamentation with a blue, amyloid stain reaction that is typical for Russulaceae

All Russulaceae, including the corticioid species, are characterised by spherical to elliptic basidiospores with a faint to very distinct (e.g. warty, spiny, or crested) ornamentation that stains bluish-black with Melzer's reagent (an amyloid stain reaction). [10] [30] Basidia (spore-bearing cells) are usually club-shaped and four-spored. [44] Russulaceae species do not have clamp connections. [45]

Characteristic cells with an oily content ( gloeocystidia ) are found in the hymenium. In Russulaceae, these show a positive colour reaction when treated with sulfoaldehydes (sulfovanillin is mostly used). [10] [30] They are also present in the hyphal sheath of ectomycorrhizal roots colonised by Russulaceae. [14]

The feature responsible for the brittle fruitbody structure in the mushroom-forming species are globular cells, called sphaerocytes or sphaerocysts, that compose the flesh (trama) alongside the usual hyphae. [30] Sometimes, these cells are clustered, and the position and arrangement of these clusters differs among genera. [30]

Another particular trama cell type are lactiferous hyphae (also lactifers). These are hyphae carrying the "milk" or "latex" exuded by the milk-caps; they react positively with sulfoaldehydes, form an abundantly branched system in the trama and end as pseudocystidia in the hymenium. [30] In general, only Lactarius, Lactifluus and Multifurca furcata possess lactifers. [15] In Russula, similar hyphae can sometimes be observed in the trama, but these are not as abundantly branched as real lactifers and do not extend into the hymenium as pseudocystidia. [30] This traditional distinction line between the "milk-caps" and Russula is however less evident in some tropical species presenting intermediate states. [36]

Genera distinction

Lactarius torminosus (3).JPG
Lactarius torminosus
Multifurca sp 85586.jpg
Unidentified Multifurca species
Zonate caps are a feature of several species of Russulaceae in Lactarius and Multifurca.

Some characteristics of the mushroom-forming genera (marked with * below) can be less obvious or absent in tropical species. [15] [36] Distinguishing between Lactarius and Lactifluus based on morphology alone is quite difficult, as clear synapomorphies for both genera have yet to be identified. [17] Most field guides treat the two genera together, often because Lactifluus is not yet recognised as a separate genus. [33] [46]

Distribution

The Russulaceae as a whole have a worldwide distribution, but patterns differ among genera. Russula is the most widespread, found in North, [47] [48] Central [49] [50] and South America, [32] [51] Europe, [33] temperate [52] [53] and tropical Asia, [54] [55] Africa, [56] and Australasia. [36] [57] [58] It is the only Russulaceae genus that occurs in the Nothofagus zone of temperate South America. [59]

Lactarius is mainly known from the north temperate zone, but some species also occur in tropical Asia and Africa. [17] Lactifluus has a more tropical distribution than Lactarius, with most species known from tropical Africa, Asia, South America, and Australasia, but some also occurring in the north temperate zone. [17] Multifurca is the rarest among the four mushroom genera, known only from some punctual records in North and Central America, Asia, and Australasia. [15] [19]

Species of Lactarius, Lactifluus, and Russula have repeatedly been introduced with trees outside their native range: An overview article lists introductions in Chile, Argentina, Uruguay, Brazil, the US, Great Britain, the Faroe Islands, South Africa, China, Thailand, and New Zealand. [60]

Among the corticioid genera, Pseudoxenasma is only known from Europe. [61] In contrast, Boidinia species have been found in Europe, [62] Taiwan, [63] and Japan, [41] and Gloeopeniophorella species in North America, [64] South America, [43] [65] Europe, [66] West Africa, [65] Taiwan, [63] Australia, [43] and New Zealand. [43]

Ecology

Ectomycorrhizal symbiosis

Evening shadows on tundra.jpg
Arctic tundra, Canada
Flickr - ggallice - Miombo woodland.jpg
Miombo woodland, Malawi
Ectomycorrhizal Russulaceae are found in a wide variety of habitats ranging from high latitudes to the tropics.

The genera Lactarius, Lactifluus, Multifurca and Russula form a mutualistic ectomycorrhizal root symbiosis with trees and shrubs, exchanging mineral nutrients for photosynthetic sugar. They are one of several fungal lineages that have evolved such a lifestyle and are sometimes referred to as the "/russula-lactarius" clade in the scientific literature. [67] Worldwide, they are one of the most frequently encountered lineages on ectomycorrhizal roots. [68] While some tropical species were initially believed to be parasitic, the observation that species fruiting on tree trunks do form ectomycorrhiza in tropical Guyana supports the view of an exclusively symbiotic lineage. [35]

Associations are known with several plant families. In the Northern Hemisphere, these are essentially the well-known ectomycorrhizal trees and shrubs in the Betulaceae, Fagaceae, Pinaceae and Salicaceae, [33] [46] but in arctic and alpine habitats, Russulaceae also associate with Bistorta vivipara (Polygonaceae), [69] Kobresia (Cyperaceae), [70] and Dryas octopetala (Rosaceae), [71] ectomycorrhizal plants untypic in their respective families. In the tropics, known plant partners include Dipterocarpaceae, [55] [72] Fabaceae, [35] [72] Nyctaginaceae, [73] [74] Phyllanthaceae, [72] Polygonaceae ( Coccoloba ), [74] Sarcolaenaceae, [75] and the gymnosperm Gnetum gnemon , [76] and in the Southern Hemisphere, Nothofagaceae, [58] [59] [77] Myrtaceae ( Eucalyptus [57] and Leptospermum [58] [77] ), and Rhamnaceae ( Pomaderris ). [78] Some Russulaceae are quite specialised in their ectomycorrhizal symbiosis, such as Lactarius and Russula species that only grow with Cistus shrubs in the Mediterranean basin. [79]

The different plant partners are reflected in the wide variety of habitats worldwide. [80] Ectomycorrhizal Russulaceae have been observed in arctic and alpine tundra, [69] boreal and alpine forest, [52] [81] north temperate forest, [33] [46] mires, [82] mediterranean forests and scrub (maquis), [79] [83] miombo woodland, [56] tropical lowland rainforest, [55] [74] tropical cloud forest, [84] tropical dry forest, [85] Australian eucalypt woodlands, [86] and south temperate forests. [59] [77] [78] Where they are introduced, they typically grow in plantations of their native host species, e.g. with pine in South Africa, [87] Eucalyptus in Thailand, [88] or birch in New Zealand. [77]

Other types of mycorrhiza

Root tips of the myco-heterotrophic plant Monotropa uniflora, with a mycorrhizal sheath formed by Russula brevipes Uniflora-root.jpg
Root tips of the myco-heterotrophic plant Monotropa uniflora , with a mycorrhizal sheath formed by Russula brevipes

Some of the ectomycorrhizal Russulaceae are also involved in other types of root symbioses with plants.

A mutualistic association similar to ectomycorrhiza but with some hyphae penetrating into the plant root cells, termed arbutoid mycorrhiza , [89] is formed by Russulaceae with shrubs of the genera Arbutus [83] and Arctostaphylos , [90] both in subfamily Arbutoideae of the Ericaceae. [83] [90]

Some Russulaceae are associated with myco-heterotrophic plants of the Ericaceae subfamily Monotropoideae, forming monotropoid mycorrhiza . [91] [92] This is an epiparasitic relationship, where the heterotrophic plant ultimately derives its carbon from the primary, ectomycorrhizal plant partner of the fungus. [92] The association is often very specific, with the heterotrophic plants only associating with selected fungus partners, including Russulaceae. [91] [93]

Russulaceae are also an important group of orchid mycorrhizal fungi. [94] This symbiosis is mutualistic in the case of green orchids, [89] but a partly or fully epiparasitic relationship in the case of myco-heterotrophic [95] [96] and mixotrophic [97] orchids, respectively. In some cases, the association with Russulaceae is, as in monotropoid mycorrhiza, very specific: the Mediterranean orchid Limodorum abortivum predominantly associates with Russula delica and closely related species; [97] in Corallorhiza maculata , different genotypes of the same species have distinct Russula partners. [95]

Wood decay species

The corticioid species in Boidinia, Gloeopeniophorella, and Pseudoxenasma are saprotrophic, wood-degrading fungi that develop on dead wood. [10] Their early-branching positions in the phylogeny suggests this has been the ancestral trophic mode of the Russulaceae, and that the mycorrhizal lifestyle (see above) evolved later. [10] The saprotrophic nature of these species has been questioned, based on the observation that other inconspicuous, crust-forming fungi are ectomycorrhizal; [14] a subsequent author reaffirms nevertheless that "[n]one of the corticioid species in the family shows any sign of mycorrhizal activity." [13]

Hypogeous fruiting

Hypogeous fruitbodies, or fruitbodies developing below ground, occur in Lactarius and Russula and have previously been considered as distinct genera (see Systematics and taxonomy: Internal systematics). As such species are especially diverse in some warm and dry regions, e.g. in Spain, [3] California, [98] or Australia, [57] below-ground fruiting has been interpreted as an adaptation to drought. [39] However, hypogeous Russulaceae are also known from cold temperate regions [23] [99] and tropical rainforest. [25] The fact that hypogeous species in the Russulaceae do not form their own lineages but are scattered in Russula or Lactarius shows that this type of fruiting evolved several times. [25] It is believed that these changes are evolutionarily quite recent. [25]

Parasites

Asterophora parasitica grows on rotting Russulaceae fruitbodies. 2011-09-27 Asterophora parasitica (Bull. ex Pers.) Singer 171045 crop.jpg
Asterophora parasitica grows on rotting Russulaceae fruitbodies.

Russulaceae fruitbodies are subject to parasitisation by other fungi. The genus Asterophora develops on old fruitbodies of the mushroom species in the family, [46] as does Dendrocollybia racemosa on at least Russula crassotunicata . [100] Fruitbodies of Lactifluus or Russula species otherwise hot-tasting and unpalatable are regarded as choice edibles in North America when infected by the "lobster mushroom" Hypomyces lactifluorum . [46] Heterotrophic plants, including orchids or monotropoids, also parasitise ectomycorrhizal Russulaceae and their plant partners – see above, Other types of mycorrhiza.

Threats and conservation

As with most fungi, [101] little information is available on the threat of extinction for Russulaceae species, and they have not been assessed in the International Union for the Conservation of Nature's Red List. [102] However, national lists contain some species of Lactarius, Lactifluus and Russula, indicating that they have small populations and are endangered, e.g. in Great Britain, [103] Switzerland, [104] the Czech Republic, [105] and New Zealand. [106]

Although data on Russulaceae themselves are scarce, more is known about the habitats they occur in, especially for the ectomycorrhizal species which depend on their host plants: Several of these habitats are affected by loss or degradation, such as peatlands, [107] Mediterranean forests and scrub [108] or tropical African dry woodland. [109] Similarly, dead wood, the habitat of the corticioid Russulaceae, is rare in many exploited forests and needs special management. [110]

Recent studies have found some traditional Russulaceae species to comprise several cryptic species (see Systematics and taxonomy: Species diversity). This may imply that distribution range and population size for each of such distinct species are smaller than previously thought. [111]

Edibility

Lactarius indigo and another Lactarius species for sale on a market in Guatemala 081117 hongos.JPG
Lactarius indigo and another Lactarius species for sale on a market in Guatemala

Several species of Lactarius, Lactifluus and Russula are valued as excellent edible mushrooms. This is the case for example for the north temperate species Lactarius deliciosus , Lactifluus volemus , or Russula vesca , and other species are popular in other parts of the world, e.g. Lactarius indigo in Mexico, or Lactifluus edulis in tropical Africa. [112] Some species, like Russula vesca, can even be eaten raw. [113] The brittle texture of Russula fruitbodies makes them different from other mushrooms and is not appreciated by some. [114]

Several species have a hot to very acrid taste and can cause gastrointestinal symptoms. [115] Despite this, such species are eaten in some regions, e.g. Lactarius torminosus in Finland [116] or Russia. [117] Often, they are parboiled or pickled to make them palatable, [118] and sometimes, they are used as spice, for example Russula emetica in Eastern Europe. [119] Some species are however truly poisonous: the East Asian and North American Russula subnigricans causes rhabdomyolysis and is potentially lethal, [120] and Lactarius turpis from Eurasia contains a mutagenic substance. [121]

Cultivation of edible Russulaceae, as in other ectomycorrhizal fungi, is challenging, since the presence of host trees is required. In spite of this difficulty, the European Lactarius deliciosus has been successfully grown in "mushroom orchards" in New Zealand. [122]

Chemistry

Lactarius indigo contains a striking blue pigment. Lactarius indigo 7283.jpg
Lactarius indigo contains a striking blue pigment.

Fruitbodies of Russulaceae have been the subject of natural product research, and different classes of organic compounds have been isolated from them.

Aroma compounds are responsible for the particular odour or taste in some species, e.g. sotolon in the fenugreek-smelling Lactarius helvus , [123] or the similar quabalactone III in Lactarius rubidus which causes a maple syrup-like odour in dried specimens. [124] Pigments have been isolated from brightly coloured species, e.g. (7-isopropenyl-4-methylazulen-1-yl)methyl stearate from the blue Lactarius indigo [125] or russulaflavidin and a derivative from the yellow Russula flavida . [126] Some Russula species contain pigmented pteridine derivatives called russupteridines that are not found in the milk-caps. [127] Sesquiterpenes are characteristic secondary metabolites of many Russulaceae, especially milk-caps which have been quite intensively studied. [128] [129] They are thought to be responsible for the hot taste in many species and may have deterrent, antifeeding functions in nature. [128]

Other metabolites isolated from different species include dibenzonaphtyridinone alkaloids , [128] prenylated phenols , [128] benzofurans , [128] chromenes , [128] natural rubber ( polyisoprene ), [130] sterols, [131] and the sugar alcohol volemitol. [132] Among toxic substances, Lactarius turpis contains the mutagenic alkaloid necatorin, [121] and the small compound cycloprop-2-ene carboxylic acid has been identified as the toxic agent in Russula subnigricans. [133] Some secondary metabolites showed antibiotic properties in laboratory tests. [128] An ethanolic extract of Russula delica was antibacterial, [134] and a lectin from Russula rosea showed antitumor activity. [135]

Related Research Articles

<span class="mw-page-title-main">Russulales</span> Order of fungi

The Russulales are an order of the Agaricomycetes,. According to the Dictionary of the Fungi, the order consists of 12 families, 80 genera, and 1767 species. According to Species Fungorum, the order contains 13 families, 117 genera, and 3,060 species.

<span class="mw-page-title-main">Agaricales</span> Order of mushrooms

The Agaricales are an order of fungi in the division Basidiomycota. As originally conceived, the order contained all the agarics, but subsequent research has shown that not all agarics are closely related and some belong in other orders, such as the Russulales and Boletales. Conversely, DNA research has also shown that many non-agarics, including some of the clavarioid fungi and gasteroid fungi belong within the Agaricales. The order has 46 extant families, more than 400 genera, and over 25,000 described species, along with six extinct genera known only from the fossil record. Species in the Agaricales range from the familiar Agaricus bisporus and the deadly Amanita virosa to the coral-like Clavaria zollingeri and bracket-like Fistulina hepatica.

<i>Lactarius</i> Genus of fungi

Lactarius is a genus of mushroom-producing, ectomycorrhizal fungi, containing several edible species. The species of the genus, commonly known as milk-caps, are characterized by the milky fluid ("latex") they exude when cut or damaged. Like the closely related genus Russula, their flesh has a distinctive brittle consistency. It is a large genus with over 500 known species, mainly distributed in the Northern hemisphere. Recently, the genus Lactifluus has been separated from Lactarius based on molecular phylogenetic evidence.

<i>Lactarius deterrimus</i> Species of fungus

Lactarius deterrimus, also known as false saffron milkcap or orange milkcap, is a species of fungus in the family Russulaceae. The fungus produces medium-sized fruit bodies (mushrooms) with orangish caps up to 12 centimetres wide that develop green spots in old age or if injured. Its orange-coloured latex stains maroon within 30 minutes. Lactarius deterrimus is a mycorrhizal fungus that associates with Norway spruce and bearberry. The species is distributed in Europe, but has also found in parts of Asia. A visually similar species in the United States and Mexico is not closely related to the European species. Fruit bodies appear between late June and November, usually in spruce forests. Although the fungus is edible—like all Lactarius mushrooms from the section Deliciosi—its taste is often bitter, and it is not highly valued. The fruit bodies are used as source of food for the larvae of several insect species. Lactarius deterrimus can be distinguished from similar Lactarius species by difference in the mycorrhizal host or latex colour.

<i>Russula</i> Genus of fungi

Russula is a very large genus composed of around 750 worldwide species of ectomycorrhizal mushrooms. They are typically common, fairly large, and brightly colored – making them one of the most recognizable genera among mycologists and mushroom collectors. Their distinguishing characteristics include usually brightly coloured caps, a white to dark yellow spore print, brittle, attached gills, an absence of latex, and absence of partial veil or volva tissue on the stem. Microscopically, the genus is characterised by the amyloid ornamented spores and flesh (trama) composed of spherocysts. Members of the related genus Lactarius have similar characteristics but emit a milky latex when their gills are broken. The genus was described by Christian Hendrik Persoon in 1796.

<span class="mw-page-title-main">Bondarzewiaceae</span> Family of fungi

The Bondarzewiaceae are a family of fungi in the order Russulales. The type species for both its genus and the family as a whole, Bondarzewia montana, closely resembles members of Polyporales, but has ornamented spores like those of Lactarius or Russula. This characteristic suggested the relationship between physically dissimilar species that eventually led to the restructuring of Russulales using molecular phylogeny. According to the Dictionary of the Fungi, the family contains 8 genera and 48 species.

<i>Lactarius torminosus</i> Fungus in the family Russulaceae from North Africa, northern Asia, Europe, and North America

Lactarius torminosus, commonly known as the woolly milkcap or the bearded milkcap, is a large agaric fungus. A common and widely distributed species, it is found in North Africa, northern Asia, Europe, and North America. It was first described scientifically by Jacob Christian Schäffer in 1774 as an Agaricus, and later transferred to the genus Lactarius in 1821 by Samuel Frederick Gray. A variety, L. torminosus var. nordmanensis, is known from the United States, Canada, and Switzerland. L. torminosus officially became the type species of Lactarius in 2011 after molecular studies prompted the taxonomic reshuffling of species between several Russulaceae genera.

<span class="mw-page-title-main">Milk-cap</span> Index of fungi with the same common name

Milk-cap is a common name that refers to mushroom-forming fungi of the genera Lactarius, Lactifluus, and Multifurca, all in the family Russulaceae. The common and eponymous feature of their fruitbodies is the latex ("milk") they exude when cut or bruised. Mushrooms with typical milk-cap characteristics are said to have a lactarioid habit. Some of them are edible.

<i>Lactifluus piperatus</i> Species of fungus

Lactifluus piperatus, commonly known as the blancaccio, is a semi-edible basidiomycete fungus of the genus Lactifluus. Despite being edible, it is not recommended by some because of its poor taste, though can be used as seasoning when dried. The fruiting body is a creamy-white mushroom which is funnel-shaped when mature, with exceptionally crowded gills. It bleeds a whitish peppery-tasting milk when cut. Widely distributed across Europe and eastern North America, Lactifluus piperatus has been accidentally introduced to Australia. Mycorrhizal, it forms a symbiotic relationship with various species of deciduous tree, including beech, and hazel, and fruiting bodies are found on the forest floor in deciduous woodland.

<i>Lactifluus volemus</i> Species of edible fungus in the family Russulaceae widely distributed in the northern hemisphere

Lactifluus volemus, formerly known as Lactarius volemus, and commonly known as the weeping milk cap or bradley, is a species of fungus in the family Russulaceae. It is widely distributed in the northern hemisphere, in temperate regions of Europe, North America and Asia as well as some subtropical and tropical regions of Central America and Asia. A mycorrhizal fungus, its fruit bodies grow on the ground at the base of various species of trees from summer to autumn, either individually or in groups. It is valued as an edible mushroom, and is sold in markets in Asia. Several other Lactifluus mushrooms resemble L. volemus, such as the closely related edible species L. corrugis, but these can be distinguished by differences in distribution, visible morphology, and microscopic characteristics. L. volemus produces a white spore print and has roughly spherical spores about 7–8 micrometres in diameter.

<i>Lactarius vietus</i> Species of fungus

Lactarius vietus is a species of fungus in the family Russulaceae, first described by Elias Magnus Fries. It produces moderately sized and brittle mushrooms, which grow on the forest floor or on rotting wood. The flattened-convex cap can vary in shape, sometimes forming the shape of a wide funnel. It is typically grey, but the colour varies. The species has crowded, light-coloured gills, which produce white milk. The spore print is typically whitish, but also varies considerably. The mushrooms typically have a strong, acrid taste and have been described as inedible, but other authors have described them as consumable after boiling. L. vietus feeds by forming an ectomycorrhizal relationship with surrounding trees, and it favours birch. It grows in autumn months and is fairly common in Europe, North America and eastern Asia.

<i>Multifurca</i> Genus of fungi

Multifurca is a rare genus of ectomycorrhizal fungi in the family Russulaceae. It was described in 2008, after molecular phylogenetic study had shown that it forms a monophyletic lineage within the family, sister to Lactarius. The genus contains six species known from the United States, Mexico, Costa Rica, India, China, Thailand, Australia, and New Caledonia, but so far has not been reported from Europe, Africa, or South America. Four of those species were formerly classified as Russula section Ochricompactae, and Multifurca furcata was originally described as a Lactarius species.

<i>Zelleromyces</i> Genus of fungi

Zelleromyces is a genus of fungi in the family Russulaceae. It was first described by mycologists Rolf Singer and Alexander H. Smith in 1960 to contain hypogeous (underground) fungi with gasteroid fruit bodies that "bleed" latex when they are cut.

<span class="mw-page-title-main">Lactifluus deceptivus</span> Species of fungus

Lactifluus deceptivus, commonly known as the deceiving milkcap, is a common species of fungus in the family Russulaceae. It is found throughout eastern North America on the ground in coniferous forests near hemlock or deciduous forests near oak, and in oak-dominated forests of Costa Rica. It produces large mushrooms with funnel-shaped caps reaching up to 25 cm (9.8 in) in diameter, on top of hard white stems that may reach 4–10 cm (1.6–3.9 in) long and up to 3 cm (1.2 in) thick. The gills are closely spaced together and yellowish-cream in color. When young, the cap is white in all parts, but the depressed center becomes dull brownish in age and breaks up into scales. The edge of the cap has a roll of cottony tissue that collapses as the cap expands. The surface of the stem—especially near the base—has a velvety texture. The mushroom "bleeds" a milky white acrid latex when it is cut or injured. Similar milk-cap species with which L. deceptivus might be confused include Lactifluus piperatus, L. pseudodeceptivus, L. caeruleitinctus, L. subvellereus, Lactarius arcuatus and Lactarius parvulus.

<i>Lactifluus</i> Genus of fungi

Lactifluus is one of three genera of mushroom-forming fungi containing species commonly named "milk-caps", the others being Lactarius and Multifurca. It has been separated from Lactarius based on molecular phylogenetic evidence but is very similar to that genus. There are roughly 150 known Lactifluus species, which have a mainly tropical distribution but are also found in the north temperate zone and Australasia. Some of them are edible mushrooms.

Russula campinensis is a neotropical species in the genus Russula. This species is highly different from most other known Russula species in that it has very small (2–17 mm), pleurotoid fruitbodies that develop on tree trunks in up to 2 m elevation from the forest floor. It was described as Lactarius campinensis from Brazil (Amazonas) by Rolf Singer. T.W. Henkel, M.C. Aime and S.L. Miller later found this species in tropical rainforest of the Pakaraima Mountains, Guyana, and showed with molecular phylogenetic methods that it belongs to Russula. Although forming fruitbodies on wood, R. campinensis is a root-symbiotic ectomycorrhizal species and has been observed in Guyana associated with Dicymbe altsonii and D. jenzenii trees in the legume subfamily Caesalpinioideae.

<i>Lactifluus clarkeae</i> Species of fungus

Lactifluus clarkeae, formerly known as Lactarius clarkeae, is a species of mushroom-forming fungus in the family Russulaceae. It is found in Australia and New Zealand in mycorrhizal association with species of Nothofagus and the family Myrtaceae.

Dr Teresa Lebel is a taxonomist and ecologist who works on fungi, with a particular interest in subterranean truffle-like fungi and their mushroom, bolete, bracket or cup relatives.

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