Sporormiella

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Sporormiella
Sporormiella vexans.png
Sporormiella vexans asci exiting fungi
Scientific classification
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Genus:
Sporormiella

Ellis & Everh.
Type species
Sporormiella nigropurpurea
Ellis & Everh.

Sporormiella is a genus of fungi in the phylum Ascomycota whose species can be found worldwide, including the Arctic [1] . It grows primarily on dung but also can be found in soil and plant debris [2] . The exact number of species is debated and can range from 60 [3] to 80 [4] in total depending on the source. A majority of these species are coprophilous [5] , however, there are a few that are endophytes (S. minimoides) [6] and saprobic [7] .

Their lifecycle is thought to require herbivorous digestion, via wild or domestic herbivores, in order for spores to properly germinate although this is still under review [8] . After being consumed and passed through the digestive tract, their fruitbodies utilize herbivorous dung as a substrate to reproduce via asci [5] . This genus is characterized by their dark, olive-brown, 4-celled spores with a defined germ slit that are contained within a gelatinous sheath that they are forcibly ejected from and stick to nearby vegetation where they will hopefully be eaten and repeat their lifecycle [9] .

This association with herbivorous animals has allowed this fungus to be utilized in a paleontological context as a proxy for megafauna presence and abundance in the late Quaternary period [5] . Research has primarily focused on the use of this proxy to track the decline of megafauna in North America in the Late Pleistocene as spores can be present in high abundances within areas with large assemblages of large grazing animals are present [10] . It is hypothesized that these groups of large animals will produce more dung and thus, more Sporormiella will be present and wash into water basins where they can be later detected in lake sediment cores; although the potential biases of using this proxy, such as spore abundances being higher at shorelines closer to grazing activity, are being discovered and discussed the more this proxy is used [11] .

Taxonomy

The genus was originally described by Ellis & Everh. in 1892 with the type species Sporormiella nigropurpurea. It was originally described separately from the genus Sporormia based on the morphological feature of a dark fungal stroma; a stroma is generally defined as a dense mass of hyphae that acts a precursor to reproductive fungal structures [12] . In 1972, Ahmed & Cain published a paper that refuted the presence of a stroma (although the dung surface was blackened) and triggered an investigation in the validity of the genus' establishment [5] . During this investigation, the paper mentions the brief split of the genus Sporormia into two genera: Sporormiopsis & Sporormia as proposed in 1944, but this decision was nullified when Sporormiopsis was determined to be synonymous with Sporormiella [3] . This brief split resulted in conflicting names for Sporormiella minima (current name) which was the type species, Sporormiopsis minima [5] . Due to this brief naming issue, the species, minima, is still known under three genera: Preussia, Sporormia, & Sporormiella [13] .

The exact number of species in the Sporormiella genus is debated and varies between different databases. Ahmed & Cain's paper created a dichotomous key for 66 described species which include detailed illustrations of all except five non-coprophilous species [5] . The Global Biodiversity Information Facility only lists 61 species in their database that utilizes data from the Species Fungorium Project based in the Royal Botanic Gardens Kew [2] . Meanwhile, Ainsworth & Bisby's Dictionary of The Fungi lists 80 species in the genus [4] .

Morphology

Microscopic image of Sporormiella sexual morph a) single ascospore cell b) single four celled ascospore c) pseudothecia Sporormiella.png
Microscopic image of Sporormiella sexual morph a) single ascospore cell b) single four celled ascospore c) pseudothecia

In Sporormiella, the sexual stage (easiest to observe) consists of small, dark brown, and glabrous (smooth and without ornamentation) to hairy pseudothesia that are unilocular (single cavity) and contain bitunicate asci [7] . Pseudothesium are a double-walled fungal structure that are the sites of spore production but lack well-organized hymenium (spore-bearing surfaces) as seen in perithesia. Asci are translucent, cylindrical to clavate, and are gelatinous [7] . They contain eight, dark to olive-brown ascospores that segmented into four cells and have a germ slit [5] that are ejected from the asci and pseudothecium upon maturity and the spore will stick to nearby vegetation to be eaten and digested [9] .

The asexual stage is characterized by septate hyphae and exists primarily within herbivorous digestion tracts [7] .

Ecology

Sporormiella is a genus of coprophilous fungi that can be found worldwide, including the Arctic [1] . As many of the species within this genus utilize dung as a substrate and are present within herbivorous intestinal tracts, it can potentially be carried long distances [9] . It is important to note they are not restricted to a specific taxonomic group and therefore can be consumed by a variety of organisms besides mammals including birds, reptiles, etc which allows this genus to potentially travel long distances [15] . This generalist lifestyle has allowed these fungi to persist for thousands of years as part of the nutrient cycling system and has become of interest to scientists to indicate the presence and abundance of herbivores over time [10] . Currently, there are calls for studies of potential obligate associations between Sporormiella species and certain herbivores; for instance, they are heavily associated with mammoths, if a species was discovered to be a specialist with mammoths, it could provide a stronger proxy in studies and procure a better understanding of Sporormiella's impacts within ecosystems through the ages [9] .

Life History

General depiction of ascospore production and lifecycle of fungus with fruitbodies in the form of perithecia. NOTE: This depicts a perithecia and not a psuedothecia as seen in Sporormiella. It also does not include the herbivorous digestion stage which is thought to trigger germination of ascospores. Ascomycota Lifecycle.webp
General depiction of ascospore production and lifecycle of fungus with fruitbodies in the form of perithecia. NOTE: This depicts a perithecia and not a psuedothecia as seen in Sporormiella. It also does not include the herbivorous digestion stage which is thought to trigger germination of ascospores.

Sporormiella generally follows the lifecycle of a coprophilous fungus in a majority of the species within this genus. In the ascus, meiosis occurs and produces four ascospores which is followed by mitosis allowing the ascus to develop eight total spores. Upon maturation, the spores will be expelled from the ascus, away from the dung and stick on vegetation such as a blade of grass [9] . The spore will stick to the vegetation until it succumbs to desiccation or is consumed by a herbivorous animal, where it will travel through the digestive tract; it is debated if this genus requires this stage for germination [10] , similarly to the phylum Neocallimastigomycota, or not [9] . After excretion from the animal, the fungi will grow on the digested plant matter substrate via septated hyphae and eventually enter its sexual stage as a telomorph where it will produce its fruitbody in the form of a pseudothesia [9] . In this pseudothecia, asci will develop and the process of meiosis and mitosis will occur again to produce spores.

No anamorph has been reported for this genus yet [16] .

Paleontological Importance

While this genus may seem inconspicuous, it has become a popular proxy used by paleoecologists worldwide in the pursuit of gathering data on ancient herbivores, with a major focus on megafauna in North America during the Quaternary period. In this area of study, the spores of Sporormiella are counted in lake core samples and converted into dung abundance and potential herbivore biomass [10] . These data points are then compared to the fossil record, pollen record, and other proxies to search for potential correlations and patterns. Spores enter the lake sediment records via slopewash of excrement from the surrounding landscape carrying spores into the lake ecosystem [10] .

The most well known application of this proxy is to track the decline and eventual extinction events of megafauna in North America with mammoths being of key interest [11] . Sporormiella spores have been found within the gut contents and on mammoth coprolites; however, a specific species has not been identified from these findings yet so scientists are unsure how specified this interaction is [10] . Regardless, Sporormiella remains one of the most utilized mycological proxies in the field. A larger herbivorous biomass present in an area could be associated with a bigger production of excrement, allowing the proliferation of this genus to potentially represent general distribution patterns.

There are two other genera that are also used as a proxy, Podospora and Sordaria, although Sporormiella remains the most popular as it is often the most abundant in samples and is relatively easy to identify [17] .

Limitations

This proxy contains some limitations and biases that are being rectified today. Spores can be easily confused with other coprophilous fungi but especially the genus Pruessia due to their similar morphologies [8] . There are also issues with spore concentrations nearest to the shore where herbivorous activity occurred and the center of lakes leaving an uneven distribution of spores throughout a basin and potentially impacting data collection [9] . The reason for lake sediment cores being taken in the middle of lakes is due to these areas often providing the best opportunities for pollen collection and stratification of sediment over the years with lower chances of sediment mixing as compared to the shoreline [8] . Another issue that is often brought up is the lack of understanding how spores preserve in different substrates and if different processes affect the spore abundance and thus data collected [9] .

It is also important to note that these coprophilous fungi are often generalists of herbivorous dung and have been found on smaller mammal dung and are being used as proxies for non-mammalian groups in some studies like one conducted in New Zealand that used this genus to trace the extinct Moa birds [15] . This versatility can be advantageous in some studies (as previously mentioned) but also can create "noise" in studies that are attempting to track abundances of specific groups, such as mammoths [9] .

Future Studies

While there are many papers that are scrutinizing the use of Sporormiella as a proxy, there are a few suggestions on future studies that could be conducted to mitigate potential biases.

Further Reading

Sporormia

Paleoecology

Related Research Articles

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of 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 defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<span class="mw-page-title-main">Megafauna</span> Large animals

In zoology, megafauna are large animals. The most common thresholds to be a megafauna are weighing over 45 kg (99 lb) or weighing over 1,000 kg (2,200 lb). The first occurrence of the term was in 1876. After the Cretaceous–Paleogene extinction event wiped out all non-avian dinosaurs, mammals and other vertebrates experienced an expansion in size. Millions of years of evolution led to gigantism on every major land mass. During the Quaternary extinction event, many species of megafauna went extinct as part of a slowly progressing extinction wave that affected ecosystems worldwide.

<i>Pilobolus</i> Genus of fungi

Pilobolus is a genus of fungi that commonly grows on herbivore dung.

<span class="mw-page-title-main">Australian megafauna</span> Large animals in Australia, past and present era

The term Australian megafauna refers to the megafauna in Australia during the Pleistocene Epoch. Most of these species became extinct during the latter half of the Pleistocene, and the roles of human and climatic factors in their extinction are contested.

<i>Sordaria fimicola</i> Species of fungus

Sordaria fimicola is a species of microscopic fungus. It is commonly found in the feces of herbivores. Sordaria fimicola is often used in introductory biology and mycology labs because it is easy to grow on nutrient agar in dish cultures. The genus Sordaria, closely related to Neurospora and Podospora, is a member of the large class Sordariomycetes, or flask-fungi. The natural habitat of the three species of Sordaria that have been the principal subjects in genetic studies is dung of herbivorous animals. The species S. fimicola is common and worldwide in distribution. The species of Sordaria are similar morphologically, producing black perithecia containing asci with eight dark ascospores in a linear arrangement. These species share a number of characteristics that are advantageous for genetic studies. They all have a short life cycle, usually 7–12 days, and are easily grown in culture. Most species are self-fertile and each strain is isogenic. All kinds of mutants are easily induced and readily obtainable with particular ascospore color mutants. These visual mutants aid in tetrad analysis, especially in analysis of intragenic recombination.

<span class="mw-page-title-main">Late Pleistocene extinctions</span> Extinctions of large mammals in the Late Pleistocene

The Late Pleistocene to the beginning of the Holocene saw numerous extinctions of predominantly megafaunal animal species, which resulted in a collapse in faunal density and diversity across the globe. The extinctions during the Late Pleistocene are differentiated from previous extinctions by the widespread absence of ecological succession to replace these extinct megafaunal species, and the regime shift of previously established faunal relationships and habitats as a consequence. The timing and severity of the extinctions varied by region and are thought to have been driven by varying combinations of human and climatic factors. Human impact on megafauna populations is thought to have been driven by hunting ("overkill"), as well as possibly environmental alteration. The relative importance of human vs climatic factors in the extinctions has been the subject of long-running controversy.

The Sporormiaceae are a family of fungi in the order Pleosporales. Taxa have a cosmopolitan distribution and are saprobic on dung (coprophilous) and rotting vegetation.

<span class="mw-page-title-main">Coprophilous fungi</span> Fungi that grow on animal dung

Coprophilous fungi are a type of saprobic fungi that grow on animal dung. The hardy spores of coprophilous species are unwittingly consumed by herbivores from vegetation, and are excreted along with the plant matter. The fungi then flourish in the feces, before releasing their spores to the surrounding area.

Sporormia is a genus of fungi in the family Sporormiaceae and part of the phylum Ascomycota. It is morphologically distinct from other genera in this family due to the orientation of cylindrical ascospores contained in a gelatinous sheath within the asci of the fruitbody. It is mostly known for being coprophilous but known to utilize wood as a substrate as well.

Polytolypa is a monotypic genus of fungus containing the single species Polytolypa hystricis. First classified in the Onygenaceae family, as of 2008 it is considered to be in the Ajellomycetaceae, although there is still uncertainty as to its phylogenetic relationships with other similar genera. This species is only known from a single specimen derived in the laboratory from a specimen of dung of the North American porcupine, Erethizon dorsatum, collected in Ontario, Canada. Polytolypa hystricis contains bioactive compounds that have antifungal activity.

Podonectria is a genus in the monotypic Podonectriaceae family of fungi. They are parasitic fungus on scale insects, other fungi, or on substrates that had previously colonized by other fungi.

<i>Coprinopsis radiata</i> Species of fungus

Coprinopsis radiata, formerly known as Coprinus radiatus, and commonly known as the miniature woolly inky cap, is a coprophilous fungus that grows on herbivore dung. It is heterothallic.

Collariella bostrychodes is a fungal decomposer of lignin and carbohydrate in the family Chaetomiaceae commonly found in soil and dung. The fungus is distinguished by a darkened collar-like ostiole around the ostiolar pore, giving the fungus its name. The fungus is highly variable in shape and form, giving raise to the belief that there are two subclades in the species. The ascospores range from lemon-shaped to nearly spherical with slightly pointed ends. It can grow to be pale green and later turn pale bluish grey or olivaceous with age. The fungus produces the toxic secondary metabolite, chaetochromin.

Zopfiella ebriosa is an unharmful fungus discovered covering the corks of wine bottles in 1991 in Tarragona, Spain. A member of the division Ascomycota, Zopfiella ebriosa is characterized by small and asymmetrical asci, presence of ostioles, and possession of germ slits.

Podospora appendiculata is a coprophilous fungus that is most commonly found in the dung of lagomorphs, such as hares and rabbits, in temperate to warm climates. A member of the division Ascomycota, P. appendiculata is characterized by ovoid, hair-studded perithecia which can bear a distinctive violaceous colouring and peridia which are coriaceous, or leathery, in texture. Podospora appendiculata has been shown to produce three compounds with antimicrobial properties.

Triangularia setosa is a member of the Ascomycota, and of the genus Triangularia. This genus is notable for its widespread appearance on the excrement of herbivores, and is therefore seen as a coprophilous fungus. The fungus itself is characteristically dark in colour and produces sac-like perithecium with a covering of hair. Its dispersion involves the ingestion, passage, and projectile ejection of spores. It has preference for colonizing the dung of lagomorphs, such as hares and rabbits.

Ascodesmis nigricans is a coprophilous fungus that could be isolated from the dung of various animals. It was firstly introduced by Philippe Édouard Léon Van Tieghem, a French botanist, and was the type species of the genus Ascodesmis. It is an uncommon species but its development of the fruit body has been the subject of much laboratory study due to the easy nature of its cultivation. Ascodesmis nigricans is not pathogenic to human, animals or plants. This species has world-wide distribution.

Anopodium ampullaceum is a species of fungus first discovered by Nils Lundqvist in Sweden, in the year 1964. A. ampullaceum became one of the first few fungi along with Anopodium epile and Podospora dagonerii, to be placed in the new genus Anopodium due to their unique spores that did not suit the description of the spores of the Podospora genus, which P. dagonerii had previously belonged to. The genus Anopodium deviates from other members of the Sordariomycetes class by two spore characteristics; firstly the pedicels of its spore in the apical position, and secondly due to its immature spores having spherical bodies with cylindrical apical regions. As of 1998 all three of these species are now considered to be one species, using the name A. ampullaceum.

Cercophora areolata is a member of the Ascomycota division, and is grouped into the Lasiosphaeriaceae family based on morphology. C. areolata is a coprophilous fungus that has been most recently isolated from porcupine dung. Defining features of C. areolata include: 1) ovoid-conical, glabrous ascomata, 2) black, carbonaceous, areolate peridium and 3) clavate-shaped, single-walled asci. From studies on C. areolata, this fungus produces multiple antifungal compounds, which inhibit other competitor fungi.

<span class="mw-page-title-main">Wood-pasture hypothesis</span> Ecological theory

The wood-pasture hypothesis is a scientific hypothesis positing that open and semi-open pastures and wood-pastures formed the predominant type of landscape in post-glacial temperate Europe, rather than the common belief of primeval forests. The hypothesis proposes that such a landscape would be formed and maintained by large wild herbivores. Although others, including landscape ecologist Oliver Rackham, had previously expressed similar ideas, it was the Dutch researcher Frans Vera, who, in his 2000 book Grazing Ecology and Forest History, first developed a comprehensive framework for such ideas and formulated them into a theorem. Vera's proposals, although highly controversial, came at a time when the role grazers played in woodlands was increasingly being reconsidered, and are credited for ushering in a period of increased reassessment and interdisciplinary research in European conservation theory and practice. Although Vera largely focused his research on the European situation, his findings could also be applied to other temperate ecological regions worldwide, especially the broadleaved ones.

References

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