Xerocomellus Zelleri
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''Xerocomellus zelleri'', commonly known as Zeller's bolete, is an
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
species of
mushroom A mushroom or toadstool is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground, on soil, or on its food source. ''Toadstool'' generally denotes one poisonous to humans. The standard for the name "mushroom" is t ...
in the family
Boletaceae The Boletaceae are a family of mushroom-forming fungi, primarily characterised by small pores on the spore-bearing hymenial surface (at the underside of the mushroom), instead of gills as are found in most agarics. Nearly as widely distributed ...
. First described scientifically by American mycologist
William Alphonso Murrill William Alphonso Murrill (October 13, 1869 – December 25, 1957) was an American mycologist, known for his contributions to the knowledge of the Agaricales and Polyporaceae. In 1904, he became the assistant Curator at the New York Botanical Ga ...
in 1912, the species has been juggled by various authors to several
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 ...
, including ''
Boletus ''Boletus'' is a genus of mushroom-producing fungi, comprising over 100 species. The genus ''Boletus'' was originally broadly defined and described by Carl Linnaeus in 1753, essentially containing all fungi with hymenial pores instead of gil ...
'', ''
Boletellus ''Boletellus'' is a genus of fungi in the family Boletaceae. The genus has a widespread distribution, especially in subtropical regions, and contains about 50 species. The genus was first described by American mycologist William Alphonso Murrill i ...
'', and ''
Xerocomus ''Xerocomus'' is a genus of poroid fungi related to ''Boletus''. Many mycologists did not originally recognize the distinction between the two genera and placed ''Xerocomus'' taxa in genus ''Boletus''. However, several molecular phylogenetic stu ...
''. Found solely in western North America from
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
south to
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, the fruit bodies are distinguished by their dark reddish brown to nearly black
caps Caps are flat headgear. Caps or CAPS may also refer to: Science and technology Computing * CESG Assisted Products Service, provided by the U.K. Government Communications Headquarters * Composite Application Platform Suite, by Java Caps, a Java ...
with uneven surfaces, the yellow pores on the underside of the caps, and the red-streaked yellow stems. The fungus grows in summer and autumn on the ground, often in
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 ...
forests or on their margins. The development of the
fruit bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
is gymnocarpic, meaning that the
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 ...
appears and develops to maturity in an exposed state, not enclosed by any protective membrane.


Taxonomy

''Xerocomellus zelleri'' was first described by American mycologist
William Alphonso Murrill William Alphonso Murrill (October 13, 1869 – December 25, 1957) was an American mycologist, known for his contributions to the knowledge of the Agaricales and Polyporaceae. In 1904, he became the assistant Curator at the New York Botanical Ga ...
in 1912, based on specimens he found on the campus of the
University of Washington The University of Washington (UW, simply Washington, or informally U-Dub) is a public research university in Seattle, Washington. Founded in 1861, Washington is one of the oldest universities on the West Coast; it was established in Seattle a ...
. Murrill named it ''Ceriomyces zelleri'' before switching the genus later that year to ''
Boletus ''Boletus'' is a genus of mushroom-producing fungi, comprising over 100 species. The genus ''Boletus'' was originally broadly defined and described by Carl Linnaeus in 1753, essentially containing all fungi with hymenial pores instead of gil ...
''. In 1944,
Walter Henry Snell Walter Henry "Doc" Snell (May 19, 1889 – July 23, 1980) was a pinch-hitter/catcher in Major League Baseball who played briefly for the Boston Red Sox during the season. Following this brief baseball career he became a successful mycologist ...
thought the taxon would be more appropriate in the genus ''
Xerocomus ''Xerocomus'' is a genus of poroid fungi related to ''Boletus''. Many mycologists did not originally recognize the distinction between the two genera and placed ''Xerocomus'' taxa in genus ''Boletus''. However, several molecular phylogenetic stu ...
''. In 1959, mycologists
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 ...
, Snell and Esther A. Dick transferred the species to ''
Boletellus ''Boletellus'' is a genus of fungi in the family Boletaceae. The genus has a widespread distribution, especially in subtropical regions, and contains about 50 species. The genus was first described by American mycologist William Alphonso Murrill i ...
'', explaining that the microstructure of the trama and the faint ornamentation of the spores were inconsistent with placement in ''Xerocomus''. American mycologist
Harry D. Thiers Harry Delbert Thiers (January 22, 1919 in Fort McKavett, Texas – August 8, 2000 in Ohio) was an American mycologist who studied and named many fungi native to North America, particularly California. Thiers taught mycology at San Francisco St ...
, in his 1976
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 ...
on North American boletes, claimed that he failed to consistently find ornamentation on the spores of material he collected, and preferred to retain the species in ''Boletus''. In 2011, it was moved to the genus ''Xerocomellus. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''zelleri'' was chosen by Murrill to honor Professor
Sanford Myron Zeller Sanford Myron Zeller (19 October 1885 – 4 November 1948) was an American mycologist. Born in Coldwater, Michigan, Zeller was educated at Lawrence College in Wisconsin, then Greenville College in Illinois, from which he received a Bachelor of S ...
, mycologist at
Oregon State University Oregon State University (OSU) is a public land-grant, research university in Corvallis, Oregon. OSU offers more than 200 undergraduate-degree programs along with a variety of graduate and doctoral degrees. It has the 10th largest engineering co ...
. Zeller accompanied Murrill in his
Seattle Seattle ( ) is a seaport city on the West Coast of the United States. It is the seat of King County, Washington. With a 2020 population of 737,015, it is the largest city in both the state of Washington and the Pacific Northwest regio ...
expedition, and discovered the first specimens of the mushroom.


Description

The
cap A cap is a flat headgear, usually with a visor. Caps have crowns that fit very close to the head. They made their first appearance as early as 3200 BC. Caps typically have a visor, or no brim at all. They are popular in casual and informal se ...
is typically between in diameter, initially convex but flattening somewhat in maturity. It is fleshy, with an uneven velvety surface, and dark brown to nearly black; the margin of the cap is a pale cream color. Young specimens are covered by a grayish bloom. The tubes that comprise the undersurface of the cap (the
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 ...
) are up to long and angular, yellow, becoming dirty yellow and finally greenish-yellow; there are 1–2 pores per millimeter on the hymenium surface. They do not change color when bruised, although they may turn slightly brownish when exposed to the air for a time. The flesh is yellow to dirty yellow, up to thick, and inconsistently bruises blue when cut or broken. The
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
is up to tall, thick, and swollen toward the base. The stem surface is red or yellowish with red lines, often white or yellow at the base, and solid (that is, not hollow), with fibrous flesh; in maturity the stem ages to yellowish-red to dark red. The
spore print 300px, Making a spore print of the mushroom ''Volvariella volvacea'' shown in composite: (photo lower half) mushroom cap laid on white and dark paper; (photo upper half) cap removed after 24 hours showing pinkish-tan spore print. A 3.5-centimeter ...
is olive-brown; one source notes that creating a spore print may result in "a lot of yellow juice on the paper". The
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 ...
are
ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a surface that may be defined as the ...
in shape, smooth, and have dimensions of 12–16 by 4–6 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
, although occasionally there will be some "giant spores" with lengths of up to 24 µm. The
basidia A basidium () is a microscopic sporangium (a spore-producing structure) found on the hymenophore of fruiting bodies of basidiomycete fungi which are also called tertiary mycelium, developed from secondary mycelium. Tertiary mycelium is highly-c ...
, the spore-bearing cells, are 26–35 by 9.5–12 µm, and four-spored. The
cystidia A cystidium (plural cystidia) is a relatively large cell found on the sporocarp of a basidiomycete (for example, on the surface of a mushroom gill), often between clusters of basidia. Since cystidia have highly varied and distinct shapes that ar ...
are roughly cylindrical and thin-walled, with dimensions of 38–77 by 5.5–14.8 µm. There are no
clamp connection A clamp connection is a hook-like structure formed by growing hyphal cells of certain fungi. It is a characteristic feature of Basidiomycetes fungi. It is created to ensure that each cell, or segment of hypha separated by septa (cross walls), rece ...
s present in the
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e. The fruit body tissue stains a greenish color when a drop of
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous was ...
solution is applied.


Edibility

''Xerocomellus zelleri'' is an
edible An edible item is any item that is safe for humans to eat. "Edible" is differentiated from "eatable" because it does not indicate how an item tastes, only whether it is fit to be eaten. Nonpoisonous items found in nature – such as some mushroo ...
species, although care should be taken to ensure that specimens collected for consumption are free of fly larvae. In his book ''100 Edible Mushrooms'', Michael Kuo gave the mushroom an edibility rating of "mediocre". There is no distinguishable odor, and the taste is alternately described as pleasant, mild, or "slightly acidic". The original species description noted that the texture was "slightly
mucilaginous Mucilage is a thick, gluey substance produced by nearly all plants and some microorganisms. These microorganisms include protists which use it for their locomotion. The direction of their movement is always opposite to that of the secretion of m ...
". The mushroom is suitable for preserving or drying, or as a "filler" to add bulk to a dish. It is harvested and sold commercially in local markets in
British Columbia, Canada British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, for ...
.


Similar species

The red-cracked bolete ('' Xerocomellus chrysenteron'') has an olive-brown cap that cracks, exposing flesh that ages to pinkish red. '' Boletellus chrysenteroides'', found only in eastern North America, has a velvety to smooth, dark reddish brown, cracked cap with pale exposed flesh. Also similar is '' Boletus mirabilis''.


Fruit body development

In 1914, Zeller published a study of the development of the mushroom, made possible by the prolific fruiting of the fungus in Seattle in the fall of 1912. Development was studied by examining thin sections of tissues in different stages of development, and the differentiation of tissues and structures followed by using histological stains. The growth form of ''Xerocomellus zelleri'' is called gymnocarpic, meaning that the hymenium appears and develops to maturity in an exposed state, not enclosed by any protective membrane. In this type of development, the cap is formed from hyphae at the top of the stem and subsequently expands by growth along the margins; the hymenium forms later beneath the cap in a direction away from the center. The mushrooms originate as minute fruit bodies (called "pins" due to their shape) from a yellow mycelium that forms a mat and tends to engulf pine needles. The pins, typically 1–2 mm in diameter, lengthen vertically until they are roughly three or four times longer than they are thick. Until this point, the fruit body is a homogenous mass of tissue. It differentiates simultaneously into cap and stem along a cleavage plane (an axis along which any cell division occurs) from the outside inward, which gives rise to deep furrow encircling the fruit body. The hymenium is formed in the roof of this furrow, growing inward and upward from the outside edge. The cap develops from the upper section of this division, the stem from the lower.


Habitat and distribution

This species grows solitarily or in small groups on the ground or in forest duff in mature
coniferous forest 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 exta ...
s, occasionally abundant on grassy edges of the forest, rarely on badly decayed conifer logs. It is an
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 ...
mushroom, meaning that the fungal hyphae form sheaths around the rootlets of certain trees, exchanging nutrients with them in a mutualistic relationship. The fungus associates with
alder Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...
, poplar and other
hardwood Hardwood is wood from dicot trees. These are usually found in broad-leaved temperate and tropical forests. In temperate and boreal latitudes they are mostly deciduous, but in tropics and subtropics mostly evergreen. Hardwood (which comes from ...
s, and has been shown in laboratory culture to form ectomycorrhizae with Western Hemlock (''
Tsuga heterophylla ''Tsuga heterophylla'', the western hemlock or western hemlock-spruce, is a species of hemlock native to the west coast of North America, with its northwestern limit on the Kenai Peninsula, Alaska, and its southeastern limit in northern Sonoma ...
''). However, the fungus may have
saprobic 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 ...
tendencies, as it has been noted to grow under
California Redwood ''Sequoia sempervirens'' ()''Sunset Western Garden Book,'' 1995:606–607 is the sole living species of the genus '' Sequoia'' in the cypress family Cupressaceae (formerly treated in Taxodiaceae). Common names include coast redwood, coastal ...
(sometimes in the rotted wood of old trunks), a tree not known to form mycorrizhae. It is known to form long
rhizomorphs 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 function ...
(aggregations of hyphae that resemble roots), and has been noted to be more abundant in sites with buried wood than without. In British Columbia, it occurs from summer to early winter, although it also appears infrequently in early spring. In California, the mushroom often fruits after the rainy period in autumn through to March or April. The dark coloring of the cap make this species difficult to notice, "unless a glimpse of the yellow hymenium is obtained". Fruit bodies are eaten by the
American shrew-mole The American shrew mole (''Neurotrichus gibbsii'') is the smallest species of mole. It is the only living member of the genus ''Neurotrichus'' and the tribe Neurotrichini. It is also known as Gibb's shrew mole and least shrew mole. It is not cl ...
. ''Xerocomellus zelleri'' is distributed in North America in the
Pacific Northwest The Pacific Northwest (sometimes Cascadia, or simply abbreviated as PNW) is a geographic region in western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though ...
south to California and Mexico. In Mexico, it has been reported in high-altitude
cloud forest A cloud forest, also called a water forest, primas forest, or tropical montane cloud forest (TMCF), is a generally tropical or subtropical, evergreen, montane, moist forest characterized by a persistent, frequent or seasonal low-level cloud c ...
s of
Mexican Beech ''Fagus mexicana'', the Mexican beech or haya, is a species of beech endemic to northeastern and central Mexico, where it occurs from Nuevo León, Tamaulipas south to Hidalgo, Veracruz and Puebla. It is restricted to Tropical montane cloud for ...
(''Fagus mexicana''), a rare and endangered habitat. It has also been reported from
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa people, ...
, but this may be based on a misidentification.


Chemistry

''Xerocomellus zelleri'' has been shown to contain the
phenethylamine Phenethylamine (PEA) is an organic compound, natural monoamine alkaloid, and trace amine, which acts as a central nervous system stimulant in humans. In the brain, phenethylamine regulates monoamine neurotransmission by binding to trace amin ...
alkaloid compounds
tyramine Tyramine ( ) (also spelled tyramin), also known under several other names, is a naturally occurring trace amine derived from the amino acid tyrosine. Tyramine acts as a catecholamine releasing agent. Notably, it is unable to cross the blood-b ...
, ''N''-methyltyramine, and
hordenine Hordenine is an alkaloid of the phenethylamine class that occurs naturally in a variety of plants, taking its name from one of the most common, barley (''Hordeum'' species). Chemically, hordenine is the ''N''-methyl derivative (chemistry), derivat ...
, although the chemotaxonomic significance of this is not clear.


See also

*
List of North American boletes __NOTOC__ This is a list of bolete species found in North America. Bolding of the species name, and an asterisk (*) following indicate the species is the type species of that genus. ''Aureoboletus'' *'' Aureoboletus auriporus'' *'' Aureoboletus ...


References


External links

* *
Photographs
at
Mushroom Observer Mushroom Observer is a collaborative mycology website started by Nathan Wilson in 2006. Reproduced on thMykoWeb website. Its purpose is to "record observations about mushrooms, help people identify mushrooms they aren't familiar with, and expand th ...
{{Taxonbar, from=Q1943850 Boletaceae Edible fungi Fungi described in 1912 Fungi of North America Taxa named by William Alphonso Murrill