Paleoserenomyces
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''Paleoserenomyces'' 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 pleosporale
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 ...
of uncertain family placement. When described it contained 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 ...
''Paleoserenomyces allenbyensis''. The genus is solely known from the
Early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian i ...
,
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
aged,
Princeton Chert The Princeton Chert is a fossil locality in British Columbia, Canada, which comprises an anatomically preserved flora of Eocene Epoch age, with rich species abundance and diversity. It is located in exposures of the Allenby Formation on the east ...
deposit of the
Allenby Formation The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an abun ...
. ''Palaeoserenomyces'' is one of only three described fossil fungus species found in the Princeton Chert, being a tar spot like parasite of the fossil palm '' Uhlia allenbyensis'', and is host for the
hyperparasite A hyperparasite, also known as a metaparasite, is a parasite whose host, often an insect, is also a parasite, often specifically a parasitoid. Hyperparasites are found mainly among the wasp-waisted Apocrita within the Hymenoptera, and in two othe ...
''
Cryptodidymosphaerites princetonensis ''Cryptodidymosphaerites'' is an extinct monotypic genus of pleosporale fungus of uncertain family placement. When described it contained the single species ''Cryptodidymosphaerites princetonensis''. The genus is solely known from the Early Eoc ...
''.


History and classification

The type series of fossils were first identified in serial thin section cellulose acetate peels of '' Uhlia allenbyensis'' fossils. The peels were made with
hydrofluoric acid Hydrofluoric acid is a Solution (chemistry), solution of hydrogen fluoride (HF) in water. Solutions of HF are colourless, acidic and highly Corrosive substance, corrosive. It is used to make most fluorine-containing compounds; examples include th ...
and mounted in Eukitt mounting material, than affixed to microscope slides using double sided tape. The
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 ...
and
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
microscope slides were accessioned into the
University of Alberta The University of Alberta, also known as U of A or UAlberta, is a public research university located in Edmonton, Alberta, Canada. It was founded in 1908 by Alexander Cameron Rutherford,"A Gentleman of Strathcona – Alexander Cameron Rutherfor ...
palaeobotanical collections and the formal description of the new genus and species was published by Randolph Currah, Ruth Stockey, and Ben LePage (1998) in the journal ''
Mycologia ''Mycologia'' is a peer-reviewed scientific journal that publishes papers on all aspects of the fungi, including lichens. It first appeared as a bimonthly journal in January 1909, published by the New York Botanical Garden under the editorship of W ...
''. The researchers formed the genus name as combination of the name '' Serenomyces'', a modern genus of phaeochoraceous
sac fungi Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defi ...
, plus the prefix ''paleo-'' denoting its status as a fossil. They chose 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 ...
''allenbyensis'' as a reference to the mining
ghost town Ghost Town(s) or Ghosttown may refer to: * Ghost town, a town that has been abandoned Film and television * Ghost Town (1936 film), ''Ghost Town'' (1936 film), an American Western film by Harry L. Fraser * Ghost Town (1956 film), ''Ghost Town'' ...
of Allenby in Canada that is also the namesake of the type locality's parent formation. The family affiliation of ''P. allenbyensis'' is undetermined, with Currah, Stockey, and LePage (1998) only assigning to the order
Phyllachorales Phyllachorales is a small order of perithecial sac fungi containing mostly foliar parasites. This order lacks reliable morphological characters making taxonomic placement of genera difficult. There is controversy among mycologists as to the bou ...
a placement which is used by some fungal databases while other databases only place the genus to the
division Division or divider may refer to: Mathematics *Division (mathematics), the inverse of multiplication *Division algorithm, a method for computing the result of mathematical division Military *Division (military), a formation typically consisting ...
Ascomycota Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
.


Description

''Paleoserenomyces allenbyensis'' stromata are darker in coloration than the ''Uhlia allenbyensis'' host cells, from which the infections can be present on either the upper or lower leaf surface. The individual stomata are around thick and up to long. Stromata cell walls comprise an inner layer of column shaped cells, round in cross-section, and an outer layer of infected leaf epidermal tissue. They are polyloculate, with the around by -thick
locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...
s formed in a layer within the stromata. Each locule has an
aperture In optics, an aperture is a hole or an opening through which light travels. More specifically, the aperture and focal length of an optical system determine the cone angle of a bundle of rays that come to a focus in the image plane. An opt ...
formed on the tip of a round umbo, and the locule walls are formed of two to four layers of
hyphae 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 ...
. None of the described fossil material had any sexual or asexual cells present, possibly dues to parasitization of the locules by ''
Cryptodidymosphaerites princetonensis ''Cryptodidymosphaerites'' is an extinct monotypic genus of pleosporale fungus of uncertain family placement. When described it contained the single species ''Cryptodidymosphaerites princetonensis''. The genus is solely known from the Early Eoc ...
''.


Distribution

''Paleoserenomyces'' is known exclusively from the
Princeton Chert The Princeton Chert is a fossil locality in British Columbia, Canada, which comprises an anatomically preserved flora of Eocene Epoch age, with rich species abundance and diversity. It is located in exposures of the Allenby Formation on the east ...
, a fossil locality 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 ...
, which comprises an anatomically preserved flora of
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...
age, with rich species abundance and diversity. The chert is located in exposures of the
Allenby Formation The Allenby formation is a sedimentary rock formation in British Columbia which was deposited during the Ypresian stage of the Early Eocene. It consists of conglomerates, sandstones with interbedded shales and coal. The shales contain an abun ...
on the east bank of the
Similkameen River The Similkameen River runs through southern British Columbia, Canada, eventually discharging into the Okanagan River near Oroville, Washington, in the United States. Through the Okanagan River, it drains to the Columbia River. The river is said ...
, south of the town of
Princeton, British Columbia Princeton (originally Vermilion Forks) is a town in the Similkameen region of southern British Columbia, Canada."The rich history of Princeton or how Vermilion Forks made it on the map...", Princeton 2008 Visitors Guide, p. 4. It lies just east o ...
. Notable in conjunction with the coal seams of the Allenby Formation are sections of
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 ...
which formed during silica-rich periods. The rapid cyclical changes from coal to chert and back are not noted in any other fossil locality in the world. An estimated 49 coal-chert cycles are known, though the exact conditions for this process are not well understood. Silica-rich volcanic episodes in the region during deposition would have been needed for formation of the cherts, while slowly moving waters and gently subsiding terrains would be needed for the peats and
fen A fen is a type of peat-accumulating wetland fed by mineral-rich ground or surface water. It is one of the main types of wetlands along with marshes, swamps, and bogs. Bogs and fens, both peat-forming ecosystems, are also known as mires. T ...
s to accumulate. Rates of organic deposition in swamps have been estimated at approximately in modern temperate climates, this suggests the time needed for each chert layer would be at least 100 years or more, with the full sequence of cycles taking place over no more than 15,000 years. The Allenby Formation is one of the southern-most of the
Eocene Okanagan Highlands The Eocene Okanagan Highlands or Eocene Okanogan Highlands are a series of Early Eocene geological formations which span a transect of British Columbia, Canada, and Washington state, United States. Known for a highly diverse and detailed Paleoflo ...
'' Lagerstätten'' in British Columbia, with the Canadian
Penticton Group Penticton ( ) is a city in the Okanagan Valley of the Southern Interior of British Columbia, Canada, situated between Okanagan and Skaha lakes. In the 2016 Canadian Census, its population was 33,761, while its census agglomeration population w ...
at the international border and the
Klondike Mountain Formation The Klondike Mountain Formation is an Early Eocene (Ypresian) geological formation located in the northeast central area of Washington state. The formation, named for the type location designated in 1962, Klondike Mountain north of Republic, Wash ...
of
Republic, Washington Republic is a city in Ferry County, Washington, United States. The population was 1,073 at the 2010 census, a 12.5% increase over the 2000 census. It is the county seat of Ferry County. It was the largest mining camp in the Republic Mining Distr ...
in northern
Ferry County Ferry County is a County (United States), county located on the northern border of the U.S. state of Washington (state), Washington. As of the 2020 United States census, 2020 census, the population was 7,178, making it the fourth-least populous ...
to the south. The highlands, including the Allenby Formation, have been described as one of the "Great Canadian ''Lagerstätten''" based on the diversity, quality and unique nature of the biotas that are preserved. The Eocene Okanagan Highlands temperate-subtropical biome preserved across a large transect of lakes recorded many of the earliest appearances of modern genera, while also documenting the last stands of ancient lines.


Paleoecology

''Paleoserenomyces allenbeyensis'' was a parasitic fungus, invading the surface cells of '' Uhlia allenbyensis'' palm leaves resulting in areas considered to be similar to modern " tar spot" infections on leaves. Within the tissues of ''P. allenbeyensis'', the hyperparasitic fungus, ''
Cryptodidymosphaerites princetonensis ''Cryptodidymosphaerites'' is an extinct monotypic genus of pleosporale fungus of uncertain family placement. When described it contained the single species ''Cryptodidymosphaerites princetonensis''. The genus is solely known from the Early Eoc ...
'' would invade cells and develop within the
stromata The ''Stromata'' ( el, Στρώματα), a mistake for ''Stromateis'' (Στρωματεῖς, "Patchwork," i.e., ''Miscellanies''), attributed to Clement of Alexandria (c. 150 – c. 215), is the third of a trilogy of works regarding the Christ ...
locules A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...
. The layered parasitic symbiosis of the palm and fungus suggests the relation between certain palms and tar spot forming
Phyllachorales Phyllachorales is a small order of perithecial sac fungi containing mostly foliar parasites. This order lacks reliable morphological characters making taxonomic placement of genera difficult. There is controversy among mycologists as to the bou ...
existed from at least .


Paleoenvironment

The Princeton chert preserves an aquatic system with silica rich slow moving waters which was likely a
peat Peat (), also known as turf (), is an accumulation of partially decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, moors, or muskegs. The peatland ecosystem covers and is the most efficien ...
–fen ecosystem. While other fossil producing areas of the Allenby Formation are likely the product of deep water deposition and
diatomite Diatomaceous earth (), diatomite (), or kieselgur/kieselguhr is a naturally occurring, soft, siliceous sedimentary rock that can be crumbled into a fine white to off-white powder. It has a particle size ranging from more than 3  μm to le ...
sedimentation, the chert layers originate from shallow waters, as evidenced by plant and animal fossils. The Okanagan Highland sites, such as the Princeton chert represent upland lake systems that were surrounded by a warm temperate ecosystem with nearby volcanism. The highlands likely had a mesic upper microthermal to lower mesothermal climate, in which winter temperatures rarely dropped low enough for snow, and which were seasonably equitable. The Okanagan Highlands paleoforest surrounding the lakes have been described as precursors to the modern
temperate broadleaf and mixed forests Temperate broadleaf and mixed forest is a temperate climate terrestrial habitat type defined by the World Wide Fund for Nature, with broadleaf tree ecoregions, and with conifer and broadleaf tree mixed coniferous forest ecoregions. These fo ...
of Eastern North America and Eastern Asia. Based on the fossil biotas the lakes were higher and cooler than the
coeval {{Short pages monitor