Palaeoserenomyces Allenbyensis
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''Paleoserenomyces'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce 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 "unisp ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of pleosporale
fungus A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
of uncertain family placement. When described it contained the single
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
''Paleoserenomyces allenbyensis''. The genus is solely known from the
Early Eocene 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 ...
,
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 ...
. ''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 ''Uhlia'' is an extinct genus of Coryphoideae, coryphoid Arecaceae, palm containing a monotypic, single species ''Uhlia allenbyensis''. The species is known from permineralized remains recovered from the Princeton Chert in British Columbia, Canad ...
'', 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 ot ...
''
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 ''Uhlia'' is an extinct genus of Coryphoideae, coryphoid Arecaceae, palm containing a monotypic, single species ''Uhlia allenbyensis''. The species is known from permineralized remains recovered from the Princeton Chert in British Columbia, Canad ...
'' fossils. The peels were made with
hydrofluoric acid Hydrofluoric acid is a solution of hydrogen fluoride (HF) in water. Solutions of HF are colorless, acidic and highly corrosive. A common concentration is 49% (48–52%) but there are also stronger solutions (e.g. 70%) and pure HF has a boiling p ...
and mounted in Eukitt mounting material, than affixed to microscope slides using double sided tape. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 (biology), isotype ...
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, the first premier of Alberta, and Henry Marshall Tory, t ...
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 ''Serenomyces'' is a genus of fungi within the family Phaeochoraceae. Species As accepted by Species Fungorum ''Index Fungorum'' is an international project to index all formal names (Binomial nomenclature, scientific names) in the fungus ...
'', 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 def ...
, plus the prefix ''paleo-'' denoting its status as a fossil. They chose the
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''allenbyensis'' as a reference to the mining
ghost town A ghost town, deserted city, extinct town, or abandoned city is an abandoned settlement, usually one that contains substantial visible remaining buildings and infrastructure such as roads. A town often becomes a ghost town because the economi ...
of
Allenby Allenby is a surname of English origin. Notable people with the surname include: *Edmund Allenby, 1st Viscount Allenby (1861–1936), British Army field-marshal ** Named after Edmund Allenby: *** Viscount Allenby, a title in the Peerage of the Unit ...
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 (biology), order of Perithecium, perithecial sac fungi containing mostly foliar parasites. This order lacks reliable morphological characters making taxonomic placement of genera difficult. There is controversy amo ...
a placement which is used by some fungal databases while other databases only place the genus to the division
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 (: locules) or loculus (; : loculi) 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'' usually refers to a chamber within an ovary ...
s formed in a layer within the stromata. Each locule has an
aperture In optics, the aperture of an optical system (including a system consisting of a single lens) is the hole or opening that primarily limits light propagated through the system. More specifically, the entrance pupil as the front side image o ...
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 o ...
. 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 is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
,
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
, which comprises an anatomically preserved flora of Eocene Epoch (geology), Epoch 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 ...
on the east bank of the Similkameen River, south of the town of Princeton, British Columbia. Notable in conjunction with the coal seams of the Allenby Formation are sections of chert 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 fens 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 ''Lagerstätten'' in British Columbia, with the Canadian Penticton Group at the international border and the Klondike Mountain Formation of Republic, Washington in northern Ferry County, Washington, Ferry County 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 ''Uhlia'' is an extinct genus of Coryphoideae, coryphoid Arecaceae, palm containing a monotypic, single species ''Uhlia allenbyensis''. The species is known from permineralized remains recovered from the Princeton Chert in British Columbia, Canad ...
'' palm leaves resulting in areas considered to be similar to modern "tar spot" infections on leaves. Within the tissues of ''P. allenbeyensis'', the hyperparasite, 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 Ascocarp#Perithecium, stromata Hymenium, locules. 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 (biology), order of Perithecium, perithecial sac fungi containing mostly foliar parasites. This order lacks reliable morphological characters making taxonomic placement of genera difficult. There is controversy amo ...
existed from at least .


Paleoenvironment

The Princeton chert preserves an aquatic system with silica rich slow moving waters which was likely a peat–fen ecosystem. While other fossil producing areas of the Allenby Formation are likely the product of deep water deposition and diatomaceous , diatomite 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 habitat, mesic Microthermal, upper microthermal to mesothermal, 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 of Eastern North America and Eastern Asia. Based on the fossil Biota (plant), biotas the lakes were higher and cooler than the coeval coastal forests preserved in the Puget Group and Chuckanut Formation of Western Washington, which are described as Tropical forest, lowland tropical forest ecosystems. Estimates of the elevation, paleoelevation range between higher than the coastal forests. This is consistent with the paleoelevation estimates for the lake systems, which range between , which is similar to the modern elevation , but higher. Estimates of the Climate, mean annual temperature have been derived from Paleothermometer#CLAMP (Climate leaf analysis multivariate program), climate leaf analysis multivariate program (CLAMP) analysis and Paleothermometer#Leaf margin analysis, leaf margin analysis (LMA) the Princeton paleoflora. The CLAMP results after multiple linear regressions for Princeton's gave a , and the LMA returned a mean annual temperature of . This is lower than the mean annual temperature estimates given for the coastal Puget Group, which is estimated to have been between . The bioclimatic analysis for Princeton suggest mean annual precipitation amount of . The warm temperate uplands floras of the Allenby Formation and greater highlands in association with downfaulted Lacustrine basin, lacustrine basins and active volcanism are noted to have no exact modern equivalents. This is due to the more seasonally equitable conditions of the Early Eocene, resulting in much lower seasonal temperature shifts. The highlands, however, have been compared to the upland ecological islands in the Virunga Mountains within the Albertine Rift of the East African Rift, African rift valley.


References

{{Taxonbar, from=Q115821265 Phyllachorales, † Eocene life of North America Prehistoric life genera Paleogene fungi Allenby Formation Fossil taxa described in 1998 Endoparasites