''Entropezites'' is an

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

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 ypermycoparasitic

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 order

Hypocreales The Hypocreales are an order of fungi within the class Sordariomycetes. In 2008, it was estimated that it contained some 237 genera, and 2647 species in seven families. Since then, a considerable number of further taxa have been identified, in ...

. At present it contains the single

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 ...

''Entropezites patricii''. The genus is solely known from the

Lower Cretaceous Lower may refer to: *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is situated about five miles south west of Dursley, eight ...

, Upper Albian stage (about 100 Ma), Burmese amber deposits in

Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...

. ''Mycetophagites'' is one of only two known instances of hyperfungal species known in the fossil record, and is the oldest to be described.

History and classification

The genus is known only from the single

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

, number "''AB-368''",

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 parasitizing a single partial fruiting body specimen. When described the

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 ...

is part of in the private collection of Ron Buckley of

Florence, Kentucky Florence is a home rule-class city in Boone County, Kentucky, United States. Florence is the second largest city located in Northern Kentucky, after Covington, and part of the Greater Cincinnati Metropolitan Area. The population was 31,946 at ...

USA The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country Continental United States, primarily located in North America. It consists of 50 U.S. state, states, a Washington, D.C., ...

. The collection has been sold and is now owned by Deniz Erin of Istanbul, Turkey.Fossil page of Ron Buckley
Retrieved 7 December 2010 ''AB-368'' was collected from one of the amber mines in the

Hukawng Valley The Hukawng Valley ( my, ဟူးကောင်းချိုင့်ဝှမ်း; also spelt Hukaung Valley) is an isolated valley in Myanmar, roughly in area. It is located in Tanaing Township in the Myitkyina District of Kachin State i ...

area southwest of Maingkhwan, Kachin Region, Northern

. It was first studied by a pair of researchers led by Dr George Poinar from

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 ...

who worked with Ron Buckley. Poinar and Buckley published their 2007

type description A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have be ...

in ''

Mycological Research ''Fungal Biology'' is a scientific journal that publishes peer-reviewed papers on all aspects of basic and applied research of the fungi, including lichens, yeasts, oomycetes, and slime moulds. A publication of the British Mycological Society, it w ...

'', journal of The British Mycological Society. The genus has been assigned the MycoBank number ''MB501250'', with the species being assigned number ''MB501251''. The generic epithet ''Entropezites'' is Greek in derivation and is a combination of the words ''entrope'' meaning "to turn inward" and ''ites'' which means "pertaining to" referencing the hypermycoparasitic nature of the species. When published, ''Entropezites patricii'', its host '' Mycetophagites atrebora'' and ''M. atreboras host '' Palaeoagaracites antiquus'' were the first known instance of hyperparasitism on

mycoparasitism A mycoparasite is an organism with the ability to parasitize fungi. Mycoparasites might be biotrophic or necrotrophic, depending on the type of interaction with their host. Types of mycoparasitic organisms Myco-heterotrophy Various plants may ...

to be described in the fossil record, and the oldest. The fossil shows that this type of fungal parasitic relationship had been established by the Albian, 100 million years ago. An earlier instance of mycoparasitism is known from the extinct species '' Palaeoserenomyces allenbyensis

and '' Cryptodidymosphaerites, Cryptodidymosphaerites princetonensis

described in 1998 from

chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a prec ...

s found in

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, ...

Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...

Description

The holotype of ''Entropezites'' consists of

e which form a complex and extensive intergrowth with the '' Mycetophagites atrebora'' host hyphae. The fungi are preserved in a rectangular piece of yellow amber approximately by by . The pileus is in diameter and possesses a convex shape with the flesh a bluish gray color and hairy. The hyphae are very small, ranging from 1 to 2.5

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 ...

in diameter and

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 ...

. The nonseptate hyphae are branched, intergrown with and penetrating the host hyphae. ''Entropezites'' hyphae penetrate host areas of necrotic tissue. The specimen sports a number of simple, hyaline conidiophores 7–10 μm in length which are borne upright from the hyphae. Each of the conidiophores is borne singly or as sparse clusters and are upright or almost upright. The single-celled condia are borne in chains on the conidiophore tips. As with the other organs of ''Entropezites'', the spherical to cylindrical condia are hyaline in coloration. Also present are possible

zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding struc ...

s ranging from 15 to 30 μm in diameter. ''Entropezites'' presents the oldest evidence of fungal hyperparasitism by other fungi in the fossil record. The fossil displays a complex interrelationship between three different fungal genera. The preserved '' Palaeoagaracites antiquus'' cap is host to both a mycoparasitic fungus and a hypermycoparasitic fungus. The surface of the gilled fungus ''Palaeoagaracites'' specimen hosts the '' Mycetophagites atrebora''

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 mycelia of ''Mycetophagites'' are found across the surface of the ''P. antiquus'' pileus, and the hyphae penetrate into the ''P. antiquus'' tissues themselves, forming necrotic areas. ''Mycetophagites'' is in turn host to a hypermycoparasitic necrotrophic fungus species ''Entropezites patricii''. Hyphae of ''Entropezites'' are preserved, penetrating the ''Mycetophagites'' hyphae forming areas of decomposing tissues. ''Entropezites'' also displays a range of growth stages for probable

s. The combined distinguishable characters of ''Entropezites'' were not enough for Poinar and Buckley to place the genus further than Hypocreales '' incertae sedis''. Where the hyphae of ''Entropezites'' penetrate into the ''Mycetophagites'' hyphae, distinct areas of necrotic tissue appear to be present. The

necrotrophic 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 t ...

nature of the interaction is similar to the modern genera '' Penicillium'', ''

Schizophyllum ''Schizophyllum'' is a genus of fungi in the family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purp ...

'' and ''

Trichoderma ''Trichoderma'' is a genus of fungi in the family Hypocreaceae that is present in all soils, where they are the most prevalent culturable fungi. Many species in this genus can be characterized as opportunistic avirulent plant symbionts. This ref ...

''. However the extensive intergrowth of the host and parasite hyphae is distinct from those found in the modern genera.

References

{{Taxonbar, from=Q5380774 †Entropezites †Entropezites Parasitic fungi Prehistoric fungi Fossil taxa described in 2007 Cretaceous fungi Natural history of Myanmar Burmese amber Fossils of Myanmar Taxa named by George Poinar Jr.