| Savoryella | |
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| Scientific classification | |
| Kingdom: | |
| Division: | |
| Class: | Sordariomycetes, (subclass Savoryellomycetidae)
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| Order: | |
| Family: | |
| Genus: | Savoryella E.B.G. Jones & R.A. Eaton,[1]
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| Type species | |
| Savoryella lignicola E.B.G.Jones & R.A.Eaton
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Savoryella is a genus of freshwater and marine based fungi in the family Savoryellaceae and the order Savoryellales.[2][3]
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.[4]
The genus had been tentatively placed in order Sphaeriales incertae sedis by Kohlmeyer & Kohlmeyer in 1979,[5] Ascomycetes incertae sedis by Eriksson & Hawksworth in 1986,[6] family Amphisphaeriaceae by Eriksson & Hawksworth in 1987,[7] order Sordariales by Jones & Hyde in 1992,[8] and order Halosphaeriales by Read et al. in 1993 based on morphological features.[9] Vijaykrishna et al. in 2006 showed Savoryella belongs to Hypocreales order based on phylogenetic analysis of partial small subunit rRNA (SSU).[10] The genus Savoryella, based on morphological features, was then placed in the Sordariales order genera incertae sedis by Jones et al. (2009),[11] and, later, Boonyuen et al. (2011),[4] showed that genera Savoryella, Ascotaiwania, Ascothailandia, and Canalisporium all cluster in the order Savoryellales within class Hypocreomycetidae,[12] Sordariomycetes.[13]
The family Savoryellaceae was then established by Jaklitsch and Réblová in 2015,[14] and was typified by the genus Savoryella. Boonyuen et al. (2011),[4] had earlier introduced the order Savoryellales, but without designating a family to it. According to phylogenetic and molecular clock analyses (Hongsanan et al., 2017;[15] Hyde et al., 2017),[16] the orders Conioscyphales, Fuscosporellales, Pleurotheciales, and Savoryellales cluster together as a distinct clade, with a stem age of 268 Mya.[15] Hence, the order Savoryellales was referred to a new subclass Savoryellomycetidae by Hongsanan et al. in 2017,[15] which was then supported by other studies.[17][13]
Generally, Savoryellales taxa share a set of characters including immersed, semi-immersed to superficial, non-stromatic, heavily pigmented, coriaceous (leathery; stiff and tough, but flexible) [ascomata]], mostly lying horizontally to the host, partly deliquescing (liquefying or melting), paraphyses, unitunicate (single-walled) asci comprises non-amyloid apical annulus, and fusiform (spindle or rod-shaped) to ellipsoidal shaped, transversely septate (walled) ascospores with hyaline (translucent) end cells and brown median cells (Jones and Eaton, 1969;[18] Jones and Hyde, 1992;[8] Tsui and Hyde, 2003;[19] Jones et al., 2009;[11] Boonyuen et al., 2011,[4]).
Taxonomic studies of marine Ascomycotina with the ultrastructure of the asci, ascospores and appendages of Savoryella were studied in 1993.[9]
The sexual morphs of Savoryellales species have perithecial (spherical, cylindrical, or flask-shaped hollow) ascomata (fruiting body) with elongate necks, while the asexual morphs are dematiaceous (produce melanin in their cell walls, giving them a characteristic brown colour especially when grown on agar) hyphomycetes with semi-macronematous conidiophores (morphologically different conidiophore from the vegetative hyphae) and monoblastic (one primary germ layer) conidiogenous (producing conidia) cells.[20][21][22][13]
Ascomata of Savoryella are long-necked and pale in color [19]
the apical pore or apparatus of ascus was not described (Jones and Eaton, 1969).[18] Later, Jones and Hyde (1992) observed the asci and apical apparatus of Savoryella appendiculata, Savoryella longispora, and Savoryella paucispora.[8] Ultrastructural 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.[23]
Paraphyses of Savoryella are inconspicuous at maturity.[4]
Ultrastructural studies of Savoryella lignicola 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 (Read et al., 1993).[9][22]
Savoryella ascospores are ellipsoid to fusiform, with several septa, and versicolorous with brown median cells and hyaline polar cells. The ascospores of Savoryella appendiculata and Savoryella paucispora have mucilaginous sheath around the central cells (Boonyuen et al., 2011). 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;[8] Read et al., 1993).[9]
Members of Savoryella are characterized by immersed to superficial ascomata, papillate, periphysate, clavate to cylindrical,
Savoryella species occur on decaying wood or substrates submerged in freshwater, marine and brackish water habitats, and are currently represented by eleven species that include five marine and six freshwater species (Jones & Eaton 1969, Minoura & Muroi 1978,[24] Koch 1982, Hyde & Goh 1998, Jones & Hyde 1992, Hyde 1993, 1994, Ho et al. 1997, 2002, Hyde & Jones 1998, Abdel-Wahab & Jones 2000,[25] Jones et al. 2016, Dayarathne et al. 2019).
Though Savoryella species are cosmopolitan in distribution, although they are mostly common in tropical and subtropical ecosystems.[26]
It has a scattered distribution worldwide.[27]
Australia,[28]
As accepted by Species Fungorum;[29]
Former species;