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''Hydnellum'' 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
tooth fungi The hydnoid fungi are a group of fungi in the Basidiomycota with basidiocarps (fruit bodies) producing spores on pendant, tooth-like or spine-like projections. They are colloquially called tooth fungi. Originally such fungi were referred to the g ...
in the family
Bankeraceae The ''Bankeraceae'' are a family of fungi in the order Thelephorales. Taxa are terrestrial, and ectomycorrhizal with plant species in families such as Pinaceae or Fagaceae. The family was circumscribed by Marinus Anton Donk in 1961. According ...
(order
Thelephorales The Thelephorales are an order (biology), order of fungi in the class (biology), class Agaricomycetes. The order includes corticioid fungi, corticioid and hydnoid fungi, together with a few polypores and clavarioid fungi, clavarioid species. Most ...
). Widely distributed in the Northern Hemisphere, the genus contains around 40 species. The fruitbodies of its members grow by slowly enveloping nearby bits of grass and vegetation. There is great variability in the form of ''Hydnellum'' fruitbodies, which are greatly influenced by environmental conditions such as rainfall and humidity, drying winds, and temperature. They are too tough and woody to eat comfortably. Several species have become the focus of increasing
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and managem ...
concern following widespread declines in abundance. ''Hydnellum'' species produce
pigments A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
that have been used to
dye A dye is a colored substance that chemically bonds to the substrate to which it is being applied. This distinguishes dyes from pigments which do not chemically bind to the material they color. Dye is generally applied in an aqueous solution an ...
textiles Textile is an umbrella term that includes various fiber-based materials, including fibers, yarns, filaments, threads, different fabric types, etc. At first, the word "textiles" only referred to woven fabrics. However, weaving is not the ...
. Several chemical compounds—some with unique
biological activity In pharmacology, biological activity or pharmacological activity describes the beneficial or adverse effects of a drug on living matter. When a drug is a complex chemical mixture, this activity is exerted by the substance's active ingredient or ...
—have been isolated and identified from the genus. One of the better-known species is the unusual pinkish-white ''
Hydnellum peckii ''Hydnellum peckii'' is a fungus in the genus '' Hydnellum'' of the family Bankeraceae. It is a hydnoid species, producing spores on the surface of vertical spines or tooth-like projections that hang from the undersurface of the fruit bodies. ...
'', also known as "strawberries and cream" or as the "bleeding tooth fungus" due to the red droplets that appear on the pinkish or whitish fruitbodies. Another species, '' H. suaveolens'', has a strong odor of
anise Anise (; '), also called aniseed or rarely anix is a flowering plant in the family Apiaceae native to Eurasia. The flavor and aroma of its seeds have similarities with some other spices and herbs, such as star anise, fennel, licorice, and ta ...
or
peppermint Peppermint (''Mentha'' × ''piperita'') is a hybrid species of mint, a cross between watermint and spearmint. Indigenous to Europe and the Middle East, the plant is now widely spread and cultivated in many regions of the world.Euro+Med Plantbas ...
.


Taxonomy

''Hydnellum'' was
circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...
by Finnish mycologist
Petter Adolf Karsten Petter Adolf Karsten (16 February 1834 – 22 March 1917) was a Finnish mycologist, the foremost expert on the fungi of Finland in his day, and known in consequence as the "father of Finnish mycology". Karsten was born in Merimasku near Turku, st ...
in 1879 with what was then known as '' Hydnum suaveolens'' as 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 ...
. Before then, fungi with spines (
hydnoid fungi The hydnoid fungi are a group of fungi in the Basidiomycota with basidiocarps (fruit bodies) producing spores on pendant, tooth-like or spine-like projections. They are colloquially called tooth fungi. Originally such fungi were referred to the g ...
) had been grouped in ''
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 ...
'' by
Elias 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 an ...
in his 1821 work ''Systema mycologicum''. Karsten defined ''Hydnellum'' as having fruitbodies with a corky or leathery, tough
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 ...
, and a centrally attached stipe.
Synonyms A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''Hydnellum'' include ''Calodon'' (Karsten, 1881), and ''Phaeodon'' (
Joseph Schröter Joseph Schröter (14 March 1837 – 12 December 1894) was a noted German mycologist, doctor and scientist. He wrote several books and texts, and discovered and described many species of flora and fungi. He also spent around fifteen years, from 187 ...
, 1888). ''Hydnellum'' is classified in the family
Bankeraceae The ''Bankeraceae'' are a family of fungi in the order Thelephorales. Taxa are terrestrial, and ectomycorrhizal with plant species in families such as Pinaceae or Fagaceae. The family was circumscribed by Marinus Anton Donk in 1961. According ...
, which was circumscribed by
Marinus Anton Donk Marinus Anton Donk (14 August 1908 – 2 September 1972) was a Dutch mycologist. He specialized in the taxonomy and nomenclature of mushrooms. Rolf Singer wrote in his obituary that he was "one of the most outstanding figures of contemporary myc ...
in 1961. The genus was not in Donk's original family concept, which included only '' Bankera'' and ''
Phellodon ''Phellodon'' is a genus of tooth fungi in the family Bankeraceae. Species have small- to medium-sized fruitbodies with white spines on the underside from which spores are released. All ''Phellodon'' have a short stalk or stipe, and so the g ...
'', genera whose species produce
hyaline A hyaline substance is one with a glassy appearance. The word is derived from el, ὑάλινος, translit=hyálinos, lit=transparent, and el, ὕαλος, translit=hýalos, lit=crystal, glass, label=none. Histopathology Hyaline cartilage is ...
(translucent), and ornamented
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 ...
. Donk left ''Hydnellum'' in the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
Hydnelleae of the family
Thelephoraceae The Thelephoraceae are a family of fungi in the order Thelephorales. This grouping of mushrooms is commonly known as the "leathery earthfans". Genera The family includes the following eight genera: * '' Amaurodon'' * '' Hypochnus'' * '' Lenzit ...
, along with ''
Sarcodon ''Sarcodon'' is a genus of fungi in the family Bankeraceae, which is part of the order Thelephorales The Thelephorales are an order of fungi in the class Agaricomycetes. The order includes corticioid and hydnoid fungi, together with a few ...
'' and '' Hydnodon''. In 1981, however, Walter Jülich emended Donk's concept of the Bankeraceae, adding hydnoid genera that produced brown, lobed spores—''Hydnellum'' and ''Sarcodon''. The name originates from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
word ''ύδνο'' (=''hydnum'') meaning "spongy plant" or "fungus". The
British Mycological Society The British Mycological Society is a learned society established in 1896 to promote the study of fungi. Formation The British Mycological Society (BMS) was formed by the combined efforts of two local societies: the Woolhope Naturalists' Field ...
, in their recommended list of
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
s for fungi in the United Kingdom, name ''Hydnellum'' fungi in the form "descriptor word" plus "tooth", such as "gold tooth" ('' H. auratile''), "zoned tooth" ('' H. concrescens''), and "velvet tooth" ('' H. spongiosipes'').


Phylogeny

Phylogenetic studies have shown that, in its current circumscription, the genus ''Hydnellum'' is polyphyletic and only distantly related to the white-spored genera '' Bankera'' and ''
Phellodon ''Phellodon'' is a genus of tooth fungi in the family Bankeraceae. Species have small- to medium-sized fruitbodies with white spines on the underside from which spores are released. All ''Phellodon'' have a short stalk or stipe, and so the g ...
'', currently grouped together in the family Bankeraceae. Analyses of the ITS region by Parfitt and colleagues and Ainsworth and colleagues, have further revealed multiple cryptic lineages within the genus. Most of the cryptic species belong to the '' H. concrescens''/'' H. scrobiculatum'' species complex, resulting in frequent misinterpretations of these old and widely applied names. At the moment, the number of phylogenetically distinct lineages confirmed by molecular sequencing exceeds the number of described taxa in the genus, making further taxonomic revisions inevitable in the future.


Morphology

''Hydnellum'' fruitbodies have caps and stipes, often with indeterminant growth forms, that may grow in spurts and decay over several weeks. Neighboring fruitbodies can coalesce, forming intricately intertwined caps and partially fused stipes. 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 ...
has a zoned appearance and is fibrous when fresh, but becomes hard and woody when dry. Zones in the flesh reflect differences in growth during periods of low daytime and high nighttime humidity, and give a fairly accurate record of daily growth. The spines are crowded closely together and are typically
decurrent ''Decurrent'' (sometimes decurring) is a term used in botany and mycology to describe plant or fungal parts that extend downward. In botany, the term is most often applied to leaf blades that partly wrap or have wings around the stem or petio ...
(extending down the length of the stipe). Fruitbodies may display a variety of colors, from white to yellow, olive green, shades of orange, light brown, or dark brown in age.
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 ...
of ''Hydnellum'' are almost spherical to oblong and tuberculate, and are brown in
mass Mass is an intrinsic property of a body. It was traditionally believed to be related to the quantity of matter in a physical body, until the discovery of the atom and particle physics. It was found that different atoms and different elementar ...
. 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) are narrowly club-shaped and usually four-spored; there are no
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 ...
in ''Hydnellum''. Three types of
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 are found in 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 ...
of ''Hydnellum'': generative hyphae (thin-walled, not inflated); skeletal hyphae (thick-walled and narrow); and thin-walled gloeoplerous-like hyphae, which
stain A stain is a discoloration that can be clearly distinguished from the surface, material, or medium it is found upon. They are caused by the chemical or physical interaction of two dissimilar materials. Accidental staining may make materials app ...
with
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 ...
. In conditions of high humidity, several species can form striking colored drops on the actively growing caps: red drops in ''H. peckii'', ''H. diabolus'', ''H. ferrugineum'', and ''H. cruentum'', yellow drops in ''H. caeruleum'', and coffee-colored drops in ''H. mirabile''.Harrison (1961)
p. 15.
/ref> The common names of ''H. peckii'' reflect its appearance: "strawberries and cream" and "bleeding tooth fungus". Some ''Hydnellum'' species have a mealy odor (e.g. ''H. mirabile'' and ''H. pineticola'') similar to freshly ground flour. ''H. zonatum'' smells like
melilot ''Melilotus'', known as melilot, sweet clover, and kumoniga (from the Cumans),Bulgarian Folk Customs, Mercia MacDermott, pg 27 is a genus in the family Fabaceae (the same family that also includes the ''Trifolium'' clovers). Members are known a ...
, while ''H. suaveolens'' has a sweet odor resembling
anise Anise (; '), also called aniseed or rarely anix is a flowering plant in the family Apiaceae native to Eurasia. The flavor and aroma of its seeds have similarities with some other spices and herbs, such as star anise, fennel, licorice, and ta ...
or
peppermint Peppermint (''Mentha'' × ''piperita'') is a hybrid species of mint, a cross between watermint and spearmint. Indigenous to Europe and the Middle East, the plant is now widely spread and cultivated in many regions of the world.Euro+Med Plantbas ...
. All are too tough and woody to be
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 ...
, and many have an acrid taste anyway. Differences between ''Hydnellum'' species tend to be more distinguishable in younger specimens. Fruitbody development is greatly influenced by environmental factors such as levels of rainfall, drying winds, and temperature. The blue tooth (''H. caeruleum''), for example, develops a deeper blue color when it grows during cooler autumn weather. Optimal growth occurs during periods of frequent light rains and high humidity; if the habitat dries out, growth will stop, but may resume after further precipitation. This intermittent growth affects the fruitbodies of different species to variable extents, leading to large variations in form, surface texture, and color. The morphological variability of fruitbodies and the dependence of their appearance upon environmental conditions has made ''Hydnellum'' a difficult group to study. Canadian mycologist Kenneth A. Harrison, who described several new species from North America, noted " e remarkable longevity of individual
sporophore {{Short pages monitor