Protomycena
Encyclopedia
Protomycena is an extinct monotypic
genus
of gilled
fungus
in the Mycenaceae
family, of order Agaricales
. At present it contains the single species
Protomycena electra, known from a single specimen collected in an amber
mine in the Cordillera Septentrional
area of the Dominican Republic
. The fruit body
of the fungus has a convex cap
that is 5 mm (0.196850393700787 in) in diameter, with distantly spaced gills on the underside. The curved stipe
is smooth and cylindrical, measuring 0.75 mm (0.0295275590551181 in) thick by 10 mm (0.393700787401575 in) long, and lacks a ring
. It resembles extant (currently living) species of the genus Mycena
. Protomycena is one of only five known agaric fungus species known in the fossil record and the second to be described from Dominican amber.
specimen, a single fruit body
(mushroom
) currently residing in the private collection owned by Ettore Morone of Turin, Italy. The specimen was collected in one of the amber mines in the Cordillera Septentrional
area of the island of Hispaniola
, in the Dominican Republic
. The amber is believed to date from the Miocene
Burdigalian
stage, about 20 to 16 million years before the present. It was first studied by a group of researchers consisting of David Hibbett
and Michael Donoghue from Harvard University
, with David Grimaldi
of the American Museum of Natural History
. Hibbett and colleagues published their 1997 type description in the American Journal of Botany
. The generic name Protomycena is derived from a combination of the Latin
proto meaning "first", and "Mycena
", a modern genus that it resembles. The specific epithet electra was coined by the authors from the Latin for "amber", in reference to the mode of preservation.
When it was reported, Protomycena electra was the third species of fossil agaric fungus to be described. The two species Coprinites dominicana
and Aureofungus yaniguaensis
are also known from the amber mines of the Dominican Republic, while the fourth species Archaeomarasmius leggeti
is from the older, Cretaceous
age New Jersey Amber. With the 2007 publication of a fifth extinct agaric species, Palaeoagaracites antiquus
, the minimum age for the Agaricales
order was pushed back to the Albian
(approximately 100 Ma).
is 5 mm (0.196850393700787 in) in diameter and has a convex shape, sporting a raised central region (an umbo
). The pale flesh
appears yellowish in the amber, and is smooth and glossy, changing to striate and slightly translucent towards the margin. The pileus margin is striated and slightly flared. The gills on the underside of the pileus are broadly attached (adnate) to the top of the stipe
, and distantly spaced—between six and eight gills extend completely from the pileus margin to the stipe
. These full-length gills are anastomosed
with lamellulae (short gills which do not reach the edge of the stipe from the pileus margin) of varying lengths. The pileus is centered on the curved stipe, which is smooth and cylindrical, measuring 0.75 mm (0.0295275590551181 in) thick by 10 mm (0.393700787401575 in) long. The stipe lacks a ring
and rhizoids. The mushroom is preserved with a small liquid and gas-filled bubble, possibly originating from the mushroom itself, which indicates the amber to be very solid and well-sealed.
In Hibbett and colleagues' 1997 publication, Protomycena was placed in the subfamily Myceneae, which at the time was considered part of the Tricholomataceae
family; Mycena is currently classified in the Mycenaceae
family. The placement was based solely on the visible structures, or macromorphology
of the fruit body. Many of the features which are typically used to classify species into fungal families
and subfamilies are based on microscopic features not visible or preserved in the amber specimen. Consequently, the assignment to Mycena is provisional (the authors also note certain similarities with extant members of Marasmius
), and the describing authors leave open the option of treating the genus placement as incertae sedis
(uncertain placement) within the Agaricales
. Protomycena is distinct from other amber-preserved mushroom taxa such as Coprinites
, in the grooved surface of its pileus and its anastomosing gills.
Monotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...
genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of gilled
Agaricales
The fungal order Agaricales, also known as gilled mushrooms , or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species, along with five extinct genera known only from the fossil record...
fungus
Fungus
A fungus is a member of a large 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, Fungi, which is separate from plants, animals, and bacteria...
in the Mycenaceae
Mycenaceae
The Mycenaceae are a family of fungi in the order Agaricales. According to the Dictionary of the Fungi , the family contains 10 genera and 705 species. This is one of several families that were separated from the Tricholomataceae as a result of phylogenetic analyses...
family, of order Agaricales
Agaricales
The fungal order Agaricales, also known as gilled mushrooms , or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species, along with five extinct genera known only from the fossil record...
. At present it contains the single species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...
Protomycena electra, known from a single specimen collected in an amber
Amber
Amber is fossilized tree resin , which has been appreciated for its color and natural beauty since Neolithic times. Amber is used as an ingredient in perfumes, as a healing agent in folk medicine, and as jewelry. There are five classes of amber, defined on the basis of their chemical constituents...
mine in the Cordillera Septentrional
Cordillera Septentrional
The Cordillera Septentrional runs parallel to the north coast of the Dominican Republic, with extensions to the northwest, the Tortuga island, and to the southeast, the Samaná Peninsula where it becomes the Sierra de Samaná. Its highest mountain is Diego de Ocampo, close to Santiago de los...
area of the Dominican Republic
Dominican Republic
The Dominican Republic is a nation on the island of La Hispaniola, part of the Greater Antilles archipelago in the Caribbean region. The western third of the island is occupied by the nation of Haiti, making Hispaniola one of two Caribbean islands that are shared by two countries...
. The fruit body
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
of the fungus has a convex cap
Pileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
that is 5 mm (0.196850393700787 in) in diameter, with distantly spaced gills on the underside. The curved stipe
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
is smooth and cylindrical, measuring 0.75 mm (0.0295275590551181 in) thick by 10 mm (0.393700787401575 in) long, and lacks a ring
Annulus (mycology)
An annulus is the ring like structure sometimes found on the stipe of some species of mushrooms. The annulus represents the remaining part of the partial veil, after it has ruptured to expose the gills or other spore-producing surface. An annulus may be thick and membranous, or it may be cobweb-like...
. It resembles extant (currently living) species of the genus Mycena
Mycena
Mycena is a large genus of small saprotrophic mushrooms that are rarely more than a few centimeters in width. They are characterized by a white spore print, a small conical or bell-shaped cap, and a thin fragile stem. Most are gray or brown, but a few species have brighter colors. Most have a...
. Protomycena is one of only five known agaric fungus species known in the fossil record and the second to be described from Dominican amber.
Discovery and classification
The genus is known only from the holotypeHolotype
A holotype is a single physical example of an organism, known to have been used when the species was formally described. It is either the single such physical example or one of several such, but explicitly designated as the holotype...
specimen, a single fruit body
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...
(mushroom
Mushroom
A mushroom is the fleshy, spore-bearing fruiting body of a fungus, typically produced above ground on soil or on its food source. The standard for the name "mushroom" is the cultivated white button mushroom, Agaricus bisporus; hence the word "mushroom" is most often applied to those fungi that...
) currently residing in the private collection owned by Ettore Morone of Turin, Italy. The specimen was collected in one of the amber mines in the Cordillera Septentrional
Cordillera Septentrional
The Cordillera Septentrional runs parallel to the north coast of the Dominican Republic, with extensions to the northwest, the Tortuga island, and to the southeast, the Samaná Peninsula where it becomes the Sierra de Samaná. Its highest mountain is Diego de Ocampo, close to Santiago de los...
area of the island of Hispaniola
Hispaniola
Hispaniola is a major island in the Caribbean, containing the two sovereign states of the Dominican Republic and Haiti. The island is located between the islands of Cuba to the west and Puerto Rico to the east, within the hurricane belt...
, in the Dominican Republic
Dominican Republic
The Dominican Republic is a nation on the island of La Hispaniola, part of the Greater Antilles archipelago in the Caribbean region. The western third of the island is occupied by the nation of Haiti, making Hispaniola one of two Caribbean islands that are shared by two countries...
. The amber is believed to date from the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
Burdigalian
Burdigalian
The Burdigalian is, in the geologic timescale, an age or stage in the early Miocene. It spans the time between 20.43 ± 0.05 Ma and 15.97 ± 0.05 Ma...
stage, about 20 to 16 million years before the present. It was first studied by a group of researchers consisting of David Hibbett
David S. Hibbett
David Hibbett is an associate professor in biology at Clark University. He is considered one of today's leading researchers "in the analysis of fungal relationships through DNA analysis." At Clark he concentrates his lab work in evolutionary biology and ecology of Fungi.He spent 1991 as a Science...
and Michael Donoghue from Harvard University
Harvard University
Harvard University is a private Ivy League university located in Cambridge, Massachusetts, United States, established in 1636 by the Massachusetts legislature. Harvard is the oldest institution of higher learning in the United States and the first corporation chartered in the country...
, with David Grimaldi
David Grimaldi
David A. Grimaldi is an entomologist and Curator of Invertebrate Zoology at the American Museum of Natural History in New York. He received his graduate training at Cornell University, where he earned his doctorate in Entomology in 1986. Dr. Grimaldi is an authority in many fields of insect...
of the American Museum of Natural History
American Museum of Natural History
The American Museum of Natural History , located on the Upper West Side of Manhattan in New York City, United States, is one of the largest and most celebrated museums in the world...
. Hibbett and colleagues published their 1997 type description in the American Journal of Botany
American Journal of Botany
The American Journal of Botany is a peer-reviewed scientific journal which includes research papers on all aspects of plant biology. It is published by the Botanical Society of America on a monthly basis since 1914....
. The generic name Protomycena is derived from a combination of the Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...
proto meaning "first", and "Mycena
Mycena
Mycena is a large genus of small saprotrophic mushrooms that are rarely more than a few centimeters in width. They are characterized by a white spore print, a small conical or bell-shaped cap, and a thin fragile stem. Most are gray or brown, but a few species have brighter colors. Most have a...
", a modern genus that it resembles. The specific epithet electra was coined by the authors from the Latin for "amber", in reference to the mode of preservation.
When it was reported, Protomycena electra was the third species of fossil agaric fungus to be described. The two species Coprinites dominicana
Coprinites
Coprinites is an extinct monotypic genus of gilled fungus in the Agaricales family Agaricaceae. At present it contains the single species Coprinites dominicana....
and Aureofungus yaniguaensis
Aureofungus
Aureofungus is an extinct monotypic genus of gilled fungus in the order Agaricales. At present it contains the single species Aureofungus yaniguaensis....
are also known from the amber mines of the Dominican Republic, while the fourth species Archaeomarasmius leggeti
Archaeomarasmius
Archaeomarasmius is an extinct genus of gilled fungus in the Agaricales family Tricholomataceae, containing the single species Archaeomarasmius leggetti. It is known from two fruit bodies recovered from amber, one consisting of a complete cap with a broken stem, the other consisting of a fragment...
is from the older, Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
age New Jersey Amber. With the 2007 publication of a fifth extinct agaric species, Palaeoagaracites antiquus
Palaeoagaracites
Palaeoagaracites is an extinct monotypic genus of gilled fungus in the order Agaricales. At present it contains the single species Palaeoagaracites antiquus....
, the minimum age for the Agaricales
Agaricales
The fungal order Agaricales, also known as gilled mushrooms , or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species, along with five extinct genera known only from the fossil record...
order was pushed back to the Albian
Albian
The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous epoch/series. Its approximate time range is 112.0 ± 1.0 Ma to 99.6 ± 0.9 Ma...
(approximately 100 Ma).
Description
The holotype of Protomycena is a single fruit body without any associated structures, preserved in a piece of clear light yellow polished amber approximately 4.5 – wide. The pileusPileus (mycology)
The pileus is the technical name for the cap, or cap-like part, of a basidiocarp or ascocarp that supports a spore-bearing surface, the hymenium. The hymenium may consist of lamellae, tubes, or teeth, on the underside of the pileus...
is 5 mm (0.196850393700787 in) in diameter and has a convex shape, sporting a raised central region (an umbo
Umbo (mycology)
thumb|right|[[Cantharellula umbonata]] has an umbo.thumb|right|The cap of [[Psilocybe makarorae]] is acutely papillate.An umbo is a raised area in the center of a mushroom cap. Caps that possess this feature are called umbonate. Umbos that are sharply pointed are called acute, while those that are...
). The pale flesh
Trama (mycology)
In mycology trama is a term for the inner, fleshy portion of a mushroom's basidiocarp, or fruit body. It is distinct from the outer layer of tissue, known as the pileipellis or cuticle, and from the spore-bearing tissue layer known as the hymenium....
appears yellowish in the amber, and is smooth and glossy, changing to striate and slightly translucent towards the margin. The pileus margin is striated and slightly flared. The gills on the underside of the pileus are broadly attached (adnate) to the top of the stipe
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
, and distantly spaced—between six and eight gills extend completely from the pileus margin to the stipe
Stipe (mycology)
thumb|150px|right|Diagram of a [[basidiomycete]] stipe with an [[annulus |annulus]] and [[volva |volva]]In mycology a stipe refers to the stem or stalk-like feature supporting the cap of a mushroom. Like all tissues of the mushroom other than the hymenium, the stipe is composed of sterile hyphal...
. These full-length gills are anastomosed
Anastomosis
An anastomosis is the reconnection of two streams that previously branched out, such as blood vessels or leaf veins. The term is used in medicine, biology, mycology and geology....
with lamellulae (short gills which do not reach the edge of the stipe from the pileus margin) of varying lengths. The pileus is centered on the curved stipe, which is smooth and cylindrical, measuring 0.75 mm (0.0295275590551181 in) thick by 10 mm (0.393700787401575 in) long. The stipe lacks a ring
Annulus (mycology)
An annulus is the ring like structure sometimes found on the stipe of some species of mushrooms. The annulus represents the remaining part of the partial veil, after it has ruptured to expose the gills or other spore-producing surface. An annulus may be thick and membranous, or it may be cobweb-like...
and rhizoids. The mushroom is preserved with a small liquid and gas-filled bubble, possibly originating from the mushroom itself, which indicates the amber to be very solid and well-sealed.
In Hibbett and colleagues' 1997 publication, Protomycena was placed in the subfamily Myceneae, which at the time was considered part of the Tricholomataceae
Tricholomataceae
The Tricholomataceae are a large family of mushrooms within the Agaricales. A classic "wastebasket taxon", the Tricholomataceae is inclusive of any white-, yellow-, or pink-spored genera in the Agaricales not already classified as belonging to the Amanitaceae, Lepiotaceae, Hygrophoraceae,...
family; Mycena is currently classified in the Mycenaceae
Mycenaceae
The Mycenaceae are a family of fungi in the order Agaricales. According to the Dictionary of the Fungi , the family contains 10 genera and 705 species. This is one of several families that were separated from the Tricholomataceae as a result of phylogenetic analyses...
family. The placement was based solely on the visible structures, or macromorphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
of the fruit body. Many of the features which are typically used to classify species into fungal families
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
and subfamilies are based on microscopic features not visible or preserved in the amber specimen. Consequently, the assignment to Mycena is provisional (the authors also note certain similarities with extant members of Marasmius
Marasmius
Marasmius is a genus of mushrooms, in the family Marasmiaceae. It contains about 500 species of agarics, of which a few, such as Marasmius oreades, are edible. However, most members of this genus are small, unimpressive brown mushrooms...
), and the describing authors leave open the option of treating the genus placement as incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...
(uncertain placement) within the Agaricales
Agaricales
The fungal order Agaricales, also known as gilled mushrooms , or euagarics, contains some of the most familiar types of mushrooms. The order has 33 extant families, 413 genera, and over 13000 described species, along with five extinct genera known only from the fossil record...
. Protomycena is distinct from other amber-preserved mushroom taxa such as Coprinites
Coprinites
Coprinites is an extinct monotypic genus of gilled fungus in the Agaricales family Agaricaceae. At present it contains the single species Coprinites dominicana....
, in the grooved surface of its pileus and its anastomosing gills.
External links
- Botanical Society of America Image of amber specimen