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''Collybia tuberosa'', commonly known as the lentil shanklet or the appleseed coincap, is an inedible
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
fungus 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 th ...
in the family
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 ...
, and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
of the genus ''
Collybia ''Collybia'' (in the strict sense) is a genus of mushrooms in the family Tricholomataceae. The genus has a widespread but rare distribution in northern temperate areas, and contains three species that grow on the decomposing remains of other mus ...
''. Like the two other members of its genus, it lives on the decomposing remains of other fleshy
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 ...
s. The fungus produces small whitish
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 ...
with
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 ...
up to wide held by thin stems up to long. On the underside of the cap are closely spaced white
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
that are broadly attached to the stem. At the base of the stem, embedded in the substrate is a small reddish-brown
sclerotium A sclerotium (; (), is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant until favo ...
that somewhat resembles an apple seed. The appearance of the sclerotium distinguishes it from the other two species of ''Collybia'', which are otherwise very similar in overall appearance. ''C. tuberosa'' is found in Europe, North America, and Japan, growing in dense clusters on species of ''
Lactarius ''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 ...
'' and ''
Russula ''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 mushr ...
'',
bolete {{refimprove, date=July 2020 A bolete is a type of mushroom, or fungal fruiting body. It can be identified thanks to a unique mushroom cap. The cap is clearly different from the stem. On the underside of the cap there is usually a spongy surfa ...
s,
hydnum ''Hydnum'' is a genus of fungi in the family Hydnaceae. They are notable for their unusual spore-bearing structures of teeth rather than gills. The best known are the edible species ''Hydnum repandum'' and '' H. rufescens''. There are no known tox ...
s, and
polypore Polypores are a group of fungi that form large fruiting bodies with pores or tubes on the underside (see Delimitation for exceptions). They are a morphological group of basidiomycetes-like gilled mushrooms and hydnoid fungi, and not all polypor ...
s.


Taxonomy, phylogeny, and naming

The species was first described under the name ''Agaricus tuberosus'' by the French naturalist Jean Bulliard in the 6th volume of his ''Herbier de la France'' (1786).
Christian Hendrik Persoon Christiaan Hendrik Persoon (1 February 1761 – 16 November 1836) was a German mycologist who made additions to Linnaeus' mushroom taxonomy. Early life Persoon was born in South Africa at the Cape of Good Hope, the third child of an imm ...
called it ''Agaricus amanitae'' subsp. ''tuberosus'' in his 1799 publication ''Observationes Mycologicae'', while
Samuel Frederick Gray Samuel Frederick Gray (10 December 1766 – 12 April 1828) was a British botanist, mycologist, and pharmacologist. He was the father of the zoologists John Edward Gray and George Robert Gray. Background He was the son of Samuel Gray, a London s ...
referred it to ''
Gymnopus ''Gymnopus'' is a genus of fungus in the family Omphalotaceae. The genus has a widespread, cosmopolitan distribution and contains about 300 species. History and classification The type species for the Gymnopus, '' Gymnopus fusipes,'' dates ba ...
'' in 1821. It was transferred to ''
Collybia ''Collybia'' (in the strict sense) is a genus of mushrooms in the family Tricholomataceae. The genus has a widespread but rare distribution in northern temperate areas, and contains three species that grow on the decomposing remains of other mus ...
'' by
Paul Kummer Paul Kummer (22 August 1834 – 6 December 1912) was a minister, teacher, and scientist in Zerbst, Germany, known chiefly for his contribution to mycological nomenclature. Earlier classification of agarics by pioneering fungal taxonomist Elias Ma ...
in 1886. The species has also been called ''Microcollybia tuberata'' in a 1979 publication by Joanne Lennox, but the genus ''Microcollybia'' has since been folded into ''Collybia''. Additional taxonomic synonyms include ''Marasmius sclerotipes'' Bres. 1881, ''Chamaeceras sclerotipes'' (Bres.) Kuntze 1898, and ''Collybia sclerotipes'' (Bres.) S.Ito 1950. It has also been named Collybia Acervata later.
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 ...
analysis reported in 2001 used
RNA sequence 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 ...
s to establish that ''C. tuberosa'' forms a
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 ...
group with '' C. cookei'' and '' C. cirrhata''; this finding was later corroborated in a 2006 publication. 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 ...
''tuberosa'' is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word for "tuberous". The mushroom is commonly known as the "lentil shanklet", or the "appleseed coincap". Samuel Gray called it the "tuberous naked-foot" in his 1821 ''Natural Arrangement of British plants''.


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 ...
of ''C. tuberosa'' ranges in shape from obtusely convex to cushion-shaped with a margin curved inward when young, to flattened in age, with margin curved downward to straight. The cap sometimes has a shallow depression in the center, or a shallow umbo. Its diameter is small, reaching a maximum of . The cap surface is dry to moist, smooth to covered with fine soft hairs, and somewhat
hygrophanous The adjective hygrophanous refers to the color change of mushroom tissue (especially the pileus surface) as it loses or absorbs water, which causes the pileipellis to become more transparent when wet and opaque when dry. When identifying hygrop ...
—changing color depending on the level of hydration. Sometimes the cap margin is pleated or grooved. The center of the cap is pinkish-
buff Buff or BUFF may refer to: People * Buff (surname), a list of people * Buff (nickname), a list of people * Johnny Buff, ring name of American world champion boxer John Lisky (1888–1955) * Buff Bagwell, a ring name of American professional ...
but whitish around the margin, and it becomes whitish overall as it matures. The
flesh Flesh is any aggregation of soft tissues of an organism. Various multicellular organisms have soft tissues that may be called "flesh". In mammals, including humans, ''flesh'' encompasses muscle Skeletal muscles (commonly referred to as mu ...
is thin, and colored whitish to light buff. The mushroom has no distinctive taste or odor, and is considered inedible though nonpoisonous. The
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
are
adnate Adnate may refer to: * Adnation, in botany, the fusion of two or more whorls of a flower * Adnate, in mycology, a classification of lamellae (gills) * Conjoined twins Conjoined twins – sometimes popularly referred to as Siamese twins – are ...
(bluntly fused to the stem), becoming subdecurrent with age (running slightly down the length of the stem). The gill spacing is close to subdistant, and the individual gills are whitish to pinkish-buff, thin, narrow to moderately broad, and have straight edges. 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 long by , and roughly equal in width throughout its length. It is slender and thread-like, flexible and pliant, with a dry surface. The top of the stem is covered with scales or a fine whitish powder, while the lower portion has hairs ranging from delicate to coarse. The color of the stem is generally whitish to pinkish-buff, but it darkens after it has been handled. The stem interior is
pith Pith, or medulla, is a tissue in 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 ...
y, and becomes hollow with age. The stems originate from a dark reddish-brown
sclerotium A sclerotium (; (), is a compact mass of hardened fungal mycelium containing food reserves. One role of sclerotia is to survive environmental extremes. In some higher fungi such as ergot, sclerotia become detached and remain dormant until favo ...
of variable shape, typically measuring by . The surface of the sclerotium is initially smooth, but later becomes wrinkled or furrowed; its interior is solid and white. It is often compared to an apple seed in appearance. Typically, several sclerotia are connected by thin strands of
mycelia Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates in ...
. The sclerotium is a resting structure that allows to fungus to overwinter in its host. In 1915,
William 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 ...
reported the sclerotia of ''C. tuberosa'' to be
bioluminescent Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorganisms including ...
. 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 white. Individual
spore 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 ...
s are smooth,
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 ...
to tear-shaped in profile, obovoid to ellipsoid or cylindric in face or back view, with dimensions of 4.2–6.2 by 2.8–3.5 μm. They are
inamyloid In mycology a tissue or feature is said to be amyloid if it has a positive amyloid reaction when subjected to a crude chemical test using iodine as an ingredient of either Melzer's reagent or Lugol's solution, producing a blue to blue-black stain ...
and acyanophilous (non-reactive to
staining Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in the ...
with
Melzer's reagent Melzer's reagent (also known as Melzer's iodine reagent, Melzer's solution or informally as Melzer's) is a chemical reagent used by mycologists to assist with the identification of fungi, and by phytopathologists for fungi that are plant pathogens ...
and
Methyl blue Methyl blue is a chemical compound with the molecular formula C37H27N3Na2O9S3. It is used as a stain in histology, and stains collagen blue in tissue sections. It can be used in some differential staining techniques such as Mallory's connect ...
, respectively). 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 ...
(spore-bearing cells in 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 club-shaped to cylindric and 15.4–21 by 3.5–5 μm. The cheilocystidia (
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 ...
on the gill edge) are scattered to infrequent, inconspicuous, and 17.5–31.5 μm long. Their shape ranges from a contorted cylinder to roughly club-shaped to irregularly diverticulate (with short offshoots approximately at right angles to the main stem). There are no pleurocystidia (cystidia on the gill face). The gill tissue is made of interwoven hyphae that are non-reactive to Melzer's reagent. These hyphae are smooth and thin-walled, measuring 2.8–6.4 μm in diameter. The cap tissue is made of hyphae that is interwoven below the center of the cap, radially oriented over the gills, and inamyloid. These hyphae are smooth, thin-walled, and 2.8–7 μm in diameter. The
cap cuticle The pileipellis is the uppermost layer of hyphae in the pileus of a fungal fruit body. It covers the trama, the fleshy tissue of the fruit body. The pileipellis is more or less synonymous with the cuticle, but the cuticle generally describes th ...
is a thin layer of smooth thin-walled hyphae that are more or less radially oriented, bent-over, cylindric and somewhat gelatinous, measuring 2–5 μm in diameter; they are occasionally diverticulate. The cuticle of the stem is made of a layer of parallel, vertically oriented smooth, thin-walled hyphae that are 2–4.2 μm in diameter, pale yellowish brown in alkali mounting solution. The stem has moderately thin-walled and smooth cystidia that are resemble
flexuous {{Short pages monitor