Savoryella
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''Savoryella'' is a genus of freshwater and marine based
fungi 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 ...
in the family '' Savoryellaceae'' and the order
Savoryellales Savoryellaceae is a Family (biology), family of aquatic based Fungus, fungi. It is the only family in the monotypic order (biology), order Savoryellales within the class Sordariomycetes, phylum, division Ascomycota. The family contains the gener ...
.


History

The taxonomic placement of ''Savoryella'' species has been widely debated since it was published, and the genus had been tentatively assigned to various orders within the Sordariomycetes class. The genus had been tentatively placed in order Sphaeriales (now
Diaporthales Diaporthales is an order (biology), order of sac fungi. Wijayawardene et al. in 2020 added a number of name families to the order. Diaporthales includes a number of plant pathogenic fungi, the most notorious of which is ''Cryphonectria parasiti ...
) incertae sedis by Kohlmeyer & Kohlmeyer in 1979, ''Ascomycetes'' incertae sedis by Eriksson & Hawksworth in 1986, family ''
Amphisphaeriaceae The ''Amphisphaeriaceae'' are a family of fungi that is mainly found in parts of New Zealand, South America, Asia and parts of Europe. According to the 2007 Outline of Ascomycota, there were 41 genera placed within the family, although the positi ...
'' by Eriksson & Hawksworth in 1987, order
Sordariales The order Sordariales is one of the most diverse taxonomic groups within the Sordariomycetes (subdivision Pezizomycotina, division Ascomycota). Species in the order Sordariales have a broad range of ecological diversity, containing lignicolous, ...
by Jones & Hyde in 1992, and order Halosphaeriales by Read et al. in 1993 based on morphological features. Vijaykrishna et al. in 2006 showed ''Savoryella'' belongs to
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 ...
order based on phylogenetic analysis of partial small subunit
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosomal ...
(SSU). The genus ''Savoryella'', based on morphological features, was then placed in the
Sordariales The order Sordariales is one of the most diverse taxonomic groups within the Sordariomycetes (subdivision Pezizomycotina, division Ascomycota). Species in the order Sordariales have a broad range of ecological diversity, containing lignicolous, ...
order genera
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
by Jones et al. (2009), and, later, Boonyuen et al. (2011), showed that genera ''Savoryella'', ''Ascotaiwania'', ''Ascothailandia'', and ''Canalisporium'' all cluster in the order Savoryellales within class
Hypocreomycetidae Hypocreomycetidae is a subclass of sac fungi. It is a highly diverse group of fungi with species from various habitats. This subclass has been reported as pathogenic, endophytic, parasitic, saprobic, fungicolous, lichenicolous, algicolous, ...
, Sordariomycetes. The family ''Savoryellaceae'' was then established by Jaklitsch and Réblová in 2015, and was typified by the genus ''Savoryella''. Boonyuen et al. (2011), had earlier introduced the order Savoryellales, but without designating a family to it. According to phylogenetic and molecular clock analyses (Hongsanan et al., 2017; Hyde et al., 2017), the orders Conioscyphales, Fuscosporellales, Pleurotheciales, and Savoryellales cluster together as a distinct clade, with a stem age of 268 Mya. Hence, the order ''Savoryellales'' was referred to a new subclass ''Savoryellomycetidae'' by Hongsanan et al. in 2017, which was then supported by other studies. Fossilised fungal spores have been found in topsoil samples on
Tibetan Plateau The Tibetan Plateau, also known as the Qinghai–Tibet Plateau or Qingzang Plateau, is a vast elevated plateau located at the intersection of Central Asia, Central, South Asia, South, and East Asia. Geographically, it is located to the north of H ...
in 2021. They were collected from different vegetation zones and show lots of variation. ''Savoryella'' spp. was found to be the dominant taxon in cropland samples.


Taxonomy

The genus name of ''Savoryella'' is derived from John George Savory (1917-2003), who was a
mycologist Mycology is the branch of biology concerned with the study of fungi, including their taxonomy, genetics, biochemical properties, and use by humans. Fungi can be a source of tinder, food, traditional medicine, as well as entheogens, poison, and ...
, born in Sacriston,
Co Durham County Durham, officially simply Durham, is a ceremonial county in North East England.UK General Acts 1997 c. 23Lieutenancies Act 1997 Schedule 1(3). From legislation.gov.uk, retrieved 6 April 2022. The county borders Northumberland and Tyne an ...
, England. He was the UK's leading expert on the special group of fungi that attack wood and wood-based materials. He made substantial contributions to testing technologies and to the derivation of national and European standards, as well as inspiring new studies on the microbial ecology of decay in wooden windows in the late 1960s and early 1970s. Then in the mid-1950s, he recognised and described a new group of wood-destroying microbes causing what he called "soft rot" - on account of the slimy texture conferred to the wood they degraded, especially under aquatic conditions. Then in 1969 a completely new genus of fungi was discovered by researchers at
Portsmouth Polytechnic The University of Portsmouth (UoP) is a public university in Portsmouth, England. Comprising five Faculty (division), faculties, the university offers a wide range of academic disciplines. in 2022, with around 28,280 students enrolled in Unde ...
to cause soft rot in the marine environment, its identifier, Evan Benjamin Gareth Jones, gave it the name of ''Savoryella'' in tribute to Savory. with Latin diminutive suffix ''ellus''.


Description

Members of the ''Savoryella'' genus, are characterized by immersed (growing wholly under water) to superficial (being on or near the surface)
ascomata An ascocarp, or ascoma (: ascomata), is the fruiting body (sporocarp (fungi), sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded ascus, asci, each of which typically contains four to ...
, (fruiting body), papillate (covered in papillae/hairs), periphysate, clavate (club-shaped) to cylindrical, unitunicate (single-walled) asci with a non-
amyloid Amyloids are aggregates of proteins characterised by a fibrillar morphology of typically 7–13 nm in diameter, a β-sheet secondary structure (known as cross-β) and ability to be stained by particular dyes, such as Congo red. In the human ...
apical thickening containing a pore, and ellipsoidal, 3-septate (walled)
ascospores In fungi, an ascospore is the sexual spore formed inside an ascus—the sac-like cell that defines the division Ascomycota, the largest and most diverse division of fungi. After two parental nuclei fuse, the ascus undergoes meiosis (halving of ...
(spores contained in an ascus) are ellipsoid to fusiform (spindle or rod-shaped), with brown central cells and hyaline (translucent)
polar cells Atmospheric circulation is the large-scale movement of air and together with ocean circulation is the means by which thermal energy is redistributed on the surface of the Earth. The Earth's atmospheric circulation varies from year to year, but ...
.
Paraphyses Paraphyses are erect sterile filament-like support structures occurring among the reproductive apparatuses of fungi, ferns, bryophytes and some thallophytes. The singular form of the word is paraphysis. In certain fungi, they are part of the f ...
of ''Savoryella'' are inconspicuous at maturity. The apical pore or apparatus of
ascus An ascus (; : asci) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some gen ...
was not originally described by Jones and Eaton, 1969). Later, Jones and Hyde (1992) observed the asci and apical apparatus of species; ''Savoryella appendiculata'', ''Savoryella longispora'', and ''Savoryella paucispora''. Ultra-structural observations of asci and ascospores with transmission electron microscopy (TEM) by Read et al. (1992) described the ascal apical ring of ''Savoryella appendiculata'' and ''Savoryella longispora'' as extending subapically on to the side walls of the ascus. Taxonomic studies of marine Ascomycotina with the ultra-structure of the asci, ascospores and appendages of ''Savoryella'' were studied in 1993. They revealed that the unitunicate ascus wall comprised an outer, 30–40 nm electron-dense layer and an inner, 420–450 nm, thick, electron-transparent layer. The sexual morphs of Savoryellales species have perithecial (spherical, cylindrical, or flask-shaped hollow)
ascomata An ascocarp, or ascoma (: ascomata), is the fruiting body (sporocarp (fungi), sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded ascus, asci, each of which typically contains four to ...
(fruiting body) with elongate necks, while the asexual morphs are dematiaceous (produce
melanin Melanin (; ) is a family of biomolecules organized as oligomers or polymers, which among other functions provide the pigments of many organisms. Melanin pigments are produced in a specialized group of cells known as melanocytes. There are ...
in their cell walls, giving them a characteristic brown colour especially when grown on
agar Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from " ogonori" and " tengusa". As found in nature, agar is a mixture of two components, t ...
) hyphomycetes with semi-macronematous
conidiophores A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
(morphologically different conidiophore from the vegetative
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 ...
) and monoblastic (one primary germ layer) conidiogenous (producing
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
) cells. The ascospores of species ''Savoryella appendiculata'' and ''Savoryella paucispora'' have mucilaginous (sticky or viscous when wet) sheath around the central cells (Boonyuen et al., 2011). Also, ''Savoryella appendiculata'' is the solitary species in this genus with ascospores with polar tetradiate appendages, formed as an outgrowth of the hyaline apical cell of the ascospore on release from the ascus (Jones and Hyde, 1992; Read et al., 1993).


Distribution and habitats

''Savoryella'' species are found on decaying wood or substrates submerged in freshwater, marine, and
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
habitats (Jones & Eaton 1969, Minoura & Muroi 1978, Koch 1982, Hyde & Goh 1998, Jones & Hyde 1992, Hyde 1993, Hyde 1994, Ho et al. 1997, Ho 2002, Hyde & Jones 1998, Abdel-Wahab & Jones 2000, Jones et al. 2016, Dayarathne et al. 2019,). Including places such as water-cooling towers. Species ''Savoryella lignicola'' was initially described from test panels of '' Fagus sylvestris'' and ''
Pinus sylvestris ''Pinus sylvestris'', the Scots pine (UK), Scotch pine (US), Baltic pine, or European red pine is a species of tree in the pine family Pinaceae that is native plant, native to Eurasia. It can readily be identified by its combination of fairly s ...
'' which were exposed in a water cooling tower exposed to brackish water. Most ''Savoryella'' species have a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
, although they are mostly common in tropical and subtropical ecosystems. They are found in places such as India, Sri Lanka, Hong Kong, Japan, and Australia. ''Savoryella melanospora'' has been found with other fungi in intertidal
mangrove A mangrove is a shrub or tree that grows mainly in coastal saline water, saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen a ...
forests within Thailand.


Species

In 2019, the genus had eleven species that include five marine, and six freshwater species. As accepted by
Species Fungorum ''Index Fungorum'' is an international project to index all formal names (Binomial nomenclature, scientific names) in the fungus Kingdom (biology), kingdom. As of 2015, the project is based at the Royal Botanic Gardens, Kew, one of three partn ...
; *'' Savoryella appendiculata'' *'' Savoryella aquatica'' *'' Savoryella curvispora'' *'' Savoryella fusiformis'' *'' Savoryella grandispora'' *'' Savoryella lignicola'' *'' Savoryella longispora'' *'' Savoryella melanospora'' *'' Savoryella nypae'' *'' Savoryella paucispora'' *'' Savoryella sarushimana'' *'' Savoryella verrucosa'' *'' Savoryella yunnanensis'' Former species; * ''S. limnetica'' = '' Neoascotaiwania limnetica'', Savoryellaceae


References


External links

{{Taxonbar, from=Q10662170 Sordariomycetes genera