Rhizopogon Subgenus Villosuli
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''Rhizopogon'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
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 ...
basidiomycetes Basidiomycota () is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya (often referred to as the "higher fungi") within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basi ...
in the family
Rhizopogonaceae ''Rhizopogonaceae'' are a family of fungi in the order Boletales. The family, first named and described by botanists Ernst Albert Gäumann and Carroll William Dodge in 1928, contains 2 genera and 151 species. The genus ''Fevansia'', formerly thou ...
. Species form hypogeous sporocarps commonly referred to as "
false truffle A false truffle or a hymenogastrale is any species of fungus that has underground fruiting bodies that produce basidiocarps resembling the true truffles of genus ''Tuber Tubers are a type of enlarged structure used as storage organs for nutri ...
s". The general morphological characters of ''Rhizopogon'' sporocarps are a
simplex In geometry, a simplex (plural: simplexes or simplices) is a generalization of the notion of a triangle or tetrahedron to arbitrary dimensions. The simplex is so-named because it represents the simplest possible polytope in any given dimension. ...
or duplex peridium surrounding a loculate gleba that lacks a columnella.
Basidiospores A basidiospore is a reproductive spore produced by Basidiomycete fungi, a grouping that includes mushrooms, shelf fungi, rusts, and smuts. Basidiospores typically each contain one haploid nucleus that is the product of meiosis, and they are pro ...
are produced upon basidia that are borne within the fungal
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some ...
that coats the interior surface of gleba locules. The peridium is often adorned with thick mycelial cords, also known as
rhizomorph Mycelial cords are linear aggregations of parallel-oriented hyphae. The mature cords are composed of wide, empty vessel hyphae surrounded by narrower sheathing hyphae. Cords may look similar to plant roots, and also frequently have similar functio ...
s, that attach the sporocarp to the surrounding
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (locomotion), the surface over which an organism lo ...
. The scientific name ''Rhizopogon'' is Greek for 'root' (Rhiz-) 'beard' (-pogon) and this name was given in reference to the rhizomorphs found on sporocarps of many species. ''Rhizopogon'' species are primarily found in ectomycorrhizal association with trees in the family Pinaceae and are especially common
symbionts Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
of
pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...
,
fir Firs (''Abies'') are a genus of 48–56 species of evergreen coniferous trees in the family (biology), family Pinaceae. They are found on mountains throughout much of North America, North and Central America, Europe, Asia, and North Africa. The ...
, and
Douglas fir The Douglas fir (''Pseudotsuga menziesii'') is an evergreen conifer species in the pine family, Pinaceae. It is native to western North America and is also known as Douglas-fir, Douglas spruce, Oregon pine, and Columbian pine. There are three va ...
trees. Through their ectomycorrhizal relationships ''Rhizopogon'' are thought to play an important role in the ecology of
coniferous forests Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All extant ...
. Recent micromorphological and
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
study has established that ''Rhizopogon'' is a member of the
Boletales The Boletales are an order of Agaricomycetes containing over 1300 species with a diverse array of fruiting body types. The boletes are the best known members of this group, and until recently, the Boletales were thought to only contain boletes. T ...
, closely related to '' Suillus''.


Taxonomy and diversity


Historical classification

The genus ''Rhizopogon'' occurs throughout the natural and introduced ranges of family Pinaceae trees. Though this range covers much of the northern temperate zones, the diversity of ''Rhizopogon'' species is well characterized only in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
and
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
. There are currently over 150 recognized species of ''Rhizopogon''. The morphology of ''Rhizopogon'' species is highly cryptic and characters vary greatly throughout sporocarp maturity. This has led to the description of multiple species from various developmental stages of a single fungus. The genus ''Rhizopogon'' was first described from Europe by
Elias Magnus Fries Elias Magnus Fries (15 August 1794 – 8 February 1878) was a Swedish mycologist and botanist. Career Fries was born at Femsjö (Hylte Municipality), Småland, the son of the pastor there. He attended school in Växjö. He acquired ...
in 1817. The North American
monograph A monograph is a specialist work of writing (in contrast to reference works) or exhibition on a single subject or an aspect of a subject, often by a single author or artist, and usually on a scholarly subject. In library cataloging, ''monograph ...
was produced by Alexander H. Smith in 1966 with second author credits given
posthumously Posthumous may refer to: * Posthumous award - an award, prize or medal granted after the recipient's death * Posthumous publication – material published after the author's death * ''Posthumous'' (album), by Warne Marsh, 1987 * ''Posthumous'' (E ...
to Sanford Myron Zeller due to his contributions to the study of the genus. A European monograph of ''Rhizopogon'' has also been published. In the recent past, molecular phylogenetic methods have allowed the revision of the taxonomic concepts of the genus ''Rhizopogon''


Modern classification

Modern taxonomic concepts of the genus ''Rhizopogon'' recognize five subgenera of ''Rhizopogon''. These are subgenus ''
Rhizopogon ''Rhizopogon'' is a genus of ectomycorrhizal basidiomycetes in the family Rhizopogonaceae. Species form hypogeous sporocarps commonly referred to as "false truffles". The general morphological characters of ''Rhizopogon'' sporocarps are a simpl ...
'', subgenus '' Versicolores'', subgenus '' Villosuli'', subgenus '' Amylopogon'', and subgenus '' Roseoli''.


Ecology


Mammalian diet and spore dispersal

''Rhizopogon'' species have been established as a common component in the diet of many small mammals as well as deer in Western North America. The viability of ''Rhizopogon'' spores is maintained and may even be increased after mammalian gut passage, making mammals an important
dispersal vector A dispersal vector is an agent of biological dispersal that moves a dispersal unit, or organism, away from its birth population to another location or population in which the individual will reproduce. These dispersal units can range from pollen t ...
for ''Rhizopogon''.


Disturbance ecology

''Rhizopogon'' species are common members of the fungal communities that colonize the roots of trees during seedling establishment and persist into old growth stands. ''Rhizopogon'' spores are long lived in soil and the spores of some species can persist for at least four years with an increase in viability over time. ''Rhizopogon'' seems to be especially common upon the roots of establishing tree seedlings following disturbance such as fire or logging. ''Rhizopogon'' are also abundant colonizers of pot cultivated and field cultivated conifer seedlings growing in soil from conifer stands that lacked observations of ''Rhizopogon'' upon the roots of mature trees. These finding suggest that ''Rhizopogon'' species are an important factor in the recovery of conifer forests following disturbance.


Invasive facilitator

''Rhizopogon'' species have been shown to have a global distribution in the Homogenocene. The enzymes exuded from some species within the subgenus amylopogon is essential in activating seed germination in some species of Monotropoideae, such as '' Pterspora andromedeae'' . This makes Rhizopogon an obligatory host to species like ''P. andromedeae.'' The exoenzymatic activity also confers higher competitive advantages to host species, mainly within the genus ''Pinus,'' by helping to break down nutrients within the soil. The presence of Rhizopogon in soil facilitates ''Pinus'' as an invasive species. This exoenzymatic activity is nitrogen limited. In the case of ''R. amylopogon'' parasitized by ''P. andromedeae'' the nitrogen cost of exoenzymatic production is in part paid for by bacteria within the family Burkholderiaceae that is hosted by ''P. andromedeae''


Species

*'' Rhizopogon albidus'' *'' Rhizopogon ater'' *''
Rhizopogon amylopogon Rhizopogon amylopogon is a sub-genus of ''Rhizopogon'' containing seven species. * '' R. brunsii'' * '' R. arctostaphyli'' * '' R. kretzerae'' * '' R. salebrosus'' * '' R. pinyonensis'' * ''R. ellenae'' * '' R. subpurpurascens'' Sub-genus ''Amy ...
'' *'' Rhizopogon atroviolaceus'' *'' Rhizopogon brunneniger'' *'' Rhizopogon ellenae'' *'' Rhizopogon evadens'' *''
Rhizopogon fulvigleba ''Rhizopogon'' is a genus of ectomycorrhizal basidiomycetes in the family Rhizopogonaceae. Species form hypogeous sporocarps commonly referred to as "false truffles". The general morphological characters of ''Rhizopogon'' sporocarps are a simpl ...
'' *'' Rhizopogon fuscorubens'' *'' Rhizopogon hawkerae'' *'' Rhizopogon luteolus'' *'' Rhizopogon nigrescens'' *''
Rhizopogon occidentalis ''Rhizopogon occidentalis'' is an ectomycorrhizal fungus in the family Rhizopogonaceae of the Basidiomycota. It occurs most commonly in western North America in association with two-needle and three-needle pine hosts. They are false truffles with ...
'' *'' Rhizopogon ochraceorubens'' *''
Rhizopogon parksii ''Rhizopogon'' is a genus of ectomycorrhizal basidiomycetes in the family Rhizopogonaceae. Species form hypogeous sporocarps commonly referred to as "false truffles". The general morphological characters of ''Rhizopogon'' sporocarps are a simp ...
'' *'' Rhizopogon parvisporus '' *'' Rhizopogon pedicellus'' *''
Rhizopogon roseolus ''Rhizopogon roseolus'', shōro (Japanese: 松露/ショウロ), is an ectomycorrhizal fungus, considered a delicacy in east Asia and Japan and used as a soil inoculant in agriculture and horticulture. Morphology The fruiting bodies are appr ...
'' *''
Rhizopogon salebrosus ''Rhizopogon salebrosus'' is a mushroom species within the ''Rhizopogon'' sub-genus '' Amylopogon''. ''R.'' ''salebrosus'' is a monotropoid mycorrhiza that is of vital importance to the ecology of conifer forests, especially in the Pacific North ...
'' *'' Rhizopogon subareolatus'' *'' Rhizopogon subaustralis'' *'' Rhizopogon subcaerulescens'' *'' Rhizopogon subpurpurascens'' *'' Rhizopogon subsalmonius'' *'' Rhizopogon succosus'' *'' Rhizopogon togasawariana'' *''
Rhizopogon truncatus ''Rhizopogon truncatus'' is an ectomycorrhizal fungus in the family Rhizopogonaceae ''Rhizopogonaceae'' are a family of fungi in the order Boletales. The family, first named and described by botanists Ernst Albert Gäumann and Carroll William ...
'' *'' Rhizopogon vesiculosus'' *'' Rhizopogon villosulus'' *''
Rhizopogon vinicolor ''Rhizopogon vinicolor'' is a species complex of ectomycorrhizal fungus which forms a mutualistic relationship with the Douglas-fir (''Pseudotsuga'' spp.). The species was first described scientifically by American mycologist Alexander H. Smith ...
'' *'' Rhizopogon vulgaris''


Ethnomycology Ethnomycology is the study of the historical uses and sociological impact of fungi and can be considered a subfield of ethnobotany or ethnobiology. Although in theory the term includes fungi used for such purposes as tinder, medicine (medicinal mu ...


Forestry

The first intentional use of ''Rhizopogon'' species in forestry occurred in the early part of the 20th century when '' Rhizopogon luteolus'' was deliberately introduced into ''
Pinus radiata ''Pinus radiata'' ( syn. ''Pinus insignis''), the Monterey pine, insignis pine or radiata pine, is a species of pine native to the Central Coast of California and Mexico (Guadalupe Island and Cedros island). It is an evergreen conifer in the fa ...
'' plantations in
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
after it was observed to improve tree growth. Since that time, ''Rhizopogon'' species have been widely studied as a component of managed forests. ''Rhizopogon'' species have been noted as common members of the ectomycorrhizal community colonizing tree roots of pine and Douglas-fir timber plantations. Naturally occurring ''Rhizopogon roseolus'' (''=rubescens'') spores have been shown to out-compete the spores of other ectomycorrhizal fungi in pine plantations even when competing spores were directly inoculated onto seedlings. The survival rate and performance of pine and Douglas-fir plantation seedlings are increased after inoculation with ''Rhizopogon'' species.


Gastronomy

Though this genera is considered edible, most members are not held in high culinary esteem. A notable exception is ''
Rhizopogon roseolus ''Rhizopogon roseolus'', shōro (Japanese: 松露/ショウロ), is an ectomycorrhizal fungus, considered a delicacy in east Asia and Japan and used as a soil inoculant in agriculture and horticulture. Morphology The fruiting bodies are appr ...
'' (''=rubescens'') which is considered a delicacy in
Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...
where it is traditionally known as shōro. Techniques for the commercial cultivation of this fungus in pine plantations have been developed and applied with successful results in Japan and
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
.


References


External links


Index FungorumA preliminary account of the North American species of ''Rhizopogon''
by Alexander H. Smith and S. M. Zeller, 1966. (Full text of monograph.) {{Taxonbar, from=Q874176 Rhizopogonaceae Boletales genera