Biology:Savoryellaceae

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Short description: Order of fungi

Savoryellaceae
Scientific classification e
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Sordariomycetes
Subclass: Savoryellomycetidae
Order: Savoryellales
Boonyuen, Suetrong, S. Sivichai, K.L. Pang & E.B.G. Jones[2]
Family: Savoryellaceae
Jaklitsch & Réblová[1]
Genera

See text.

Savoryellaceae is a family of aquatic based fungi. It is the only family in the monotypic order Savoryellales within the class Sordariomycetes, division Ascomycota.[3][4]

The family contains the genera: Ascotaiwania (11 species), Canalisporium (21 species), Dematiosporium (1 species; Dematiosporium aquaticum Z.L. Luo, K.D. Hyde & H.Y. Su (2019)),[5] Monotosporella (3 species), Neoascotaiwania (4 species) and lastly, Savoryella (13 species).[4]

History

The taxonomic placement of freshwater and marine Savoryella species has been widely debated, and the genus had been tentatively assigned to various orders within the Sordariomycetes class.[2]

The genus had been tentatively placed in order Sphaeriales incertae sedis (now Diaporthales) by Kohlmeyer & Kohlmeyer in 1979,[6] Ascomycetes incertae sedis by Eriksson & Hawksworth in 1986,[7] family Amphisphaeriaceae by Eriksson & Hawksworth in 1987,[8] order Sordariales by Jones & Hyde in 1992,[9] and order Halosphaeriales by Read et al. in 1993 based on morphological features.[10] Vijaykrishna et al. in 2006 showed Savoryella belongs to Hypocreales order based on phylogenetic analysis of partial small subunit rRNA (SSU).[11] The genus Savoryella, based on morphological features, was then placed in the Sordariales order genera incertae sedis by Jones et al. (2009),[12] and, later, Boonyuen et al. (2011),[2] showed that genera Savoryella, Ascotaiwania, Ascothailandia, and Canalisporium all cluster in the order Savoryellales within class Hypocreomycetidae,[13] Sordariomycetes.[14] According to the one fungus-one name concept by Hawksworth in 2011,[15] the genus Canalisporium was recommended for protection over Ascothailandia hence, it was then synonymized under Canalisporium based on sequence data (Sri-indrasutdhi et al., 2010).[16]

The family Savoryellaceae (Savoryellales) was then established by Jaklitsch and Réblová in 2015,[1] and was typified by the genus Savoryella. Boonyuen et al. (2011),[2] had earlier introduced the order Savoryellales, but without designating a family to it. According to phylogenetic and molecular clock analyses (Hongsanan et al., 2017;[17] Hyde et al., 2017),[18] the orders Conioscyphales, Fuscosporellales, Pleurotheciales, and Savoryellales cluster together as a distinct clade, with a stem age of 268 Mya.[17] Hence, the order Savoryellales was referred to a new subclass Savoryellomycetidae by Hongsanan et al. (2017),[17] which was then supported by other studies.[14][19]

In 2016, Savoryellales consisted of family Savoryellaceae with three genera, Ascotaiwania, Canalisporium and Savoryella.[20] Then Maharachchikumbura et al. (2016),[21] and Wijayawardene et al. (2018),[19] accepted the placement of Savoryellaceae in the order Savoryellales. With the inclusion of the genus Neoascotaiwania, Savoryellaceae comprised four genera: Ascotaiwania, Canalisporium, and Savoryella (Hernández-Restrepo et al.) in 2017.[22][23] Then freshwater genera and asexual fungus from Erhai lake in China, Dematiosporium was added to the Savoryellaceae family in 2019.[5] Lastly, anamorphic fungal genus Monotosporella was also added later to the order and family.[3]

Description

Generally, Savoryellaceae species 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;[24] Jones and Hyde, 1992;[9] Tsui and Hyde, 2003;[25] Jones et al., 2009;[12] Boonyuen et al., 2011,[2]).

Taxonomic studies of marine Ascomycotina with the ultrastructure of the asci, ascospores and appendages of Savoryella were studied in 1993.[10]

The sexual morphs of Savoryellaceae 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.[23][26][27]

Different types of asexual morphs (based on morphological observations only) have been experimentally linked to Ascotaiwania and Neoascotaiwania species such as, monotosporella-like in Ascotaiwania sawada (Sivichai et al., 1998),[28] and Ascotaiwania mitriformis (Ranghoo and Hyde, 1998),[29] monodictys-like in Ascotaiwania lignicola (Chang, 2001),[30] trichocladium-like in Ascotaiwania hsilio (Chang, 2001),[30] and bactrodesmium-like asexual morphs for Neoascotaiwania species (Hernández-Restrepo et al., 2017).[22] These various asexual morphs have previously led to confusion in the classification of various sexual genera in the family. Canalisporium species comprise muriform (spores arranged like bricks in a wall) conidia (with sexual morph – Ascothailandia; Sri-indrasutdhi et al., 2010),[16] but not all species have been sequenced. Therefore, Savoryellaceae asexual morphs are associated with dematiaceous hyphomycetes characterized by semi-macronematous conidiophores (conidiophores that are only slightly morphologically different from the vegetative hyphae) and monoblastic conidiogenous cells generating brown, thick-walled, dictyoseptate conidia that are transversely septate or cheiroid.[26] Cheiroid means "spores in which several to many columns of cells arise from a common origin and grow parallel or diverge moderately". The distant position of Helicoon farinosum (asexual morph of Ascotaiwania hughesii) (Fallah et al., 1999),[31] from the rest of members of Savoryellaceae was confirmed by phylogenetic analysis (Boonyuen et al., 2011;[2] Réblová et al., 2012[32])".[23]

Distribution and habitat

The various fungal species in the order are found in terrestrial, marine,[33] brackish and freshwater habitats,[5][34][35] this includes places such as water-cooling towers,[24][36] and also reservoirs.[14][29]

Some species of Savoryellaceae are found abundant on submerged wood in aquatic habitats and some species have been reported found on terrestrial woody plants.[37][38]

For example; Ascotaiwania species have been isolated from submerged and decaying wood in freshwater habitats,[39][40] and are widely distributed in countries such as Ecuador, France, Great Britain, Hong Kong, Malaysia, Mauritius, Taiwan, Thailand, and Australia. While Canalisporium species are saprobes (processing of decayed (dead or waste) organic matter), mostly on rotten wood and are distributed in tropical regions of both hemispheres of the world.[41] They are also found on woody plants,[42][43] and submerged, decaying wood in freshwater.[28][44] Canalisporium species have been recorded from Cuba,[45] India,[42] Kenya,[46] Malaysia,[47] Taiwan, Thailand,[16] and Uganda.[48] Though Savoryella species are cosmopolitan in distribution, although they are mostly common in tropical and subtropical ecosystems.[36] Neoascotaiwania species have been documented from forest soil in Spain,[22] and also on decaying wood collected in streams in Taiwan.[23][35] Lastly, genus Monotosporella is distributed worldwide,[28][43] and it is usually found growing on decaying wood submerged in freshwater.[28][44] but has also been reported as being found on woody plants outside of aquatic habitats.[42][43]

Species in the family Savoryellaceae generally have a scattered distribution worldwide.[49] Including places such as; North America,[31] (including Florida,[43]) Cuba,[45] parts of Southern Europe (including Spain,[22] and Hungary,[50]), Africa including Kenya,[46] India,[42][51] Thailand,[16] Malaysia,[47] Taiwan,[35][39] China,[52] Japan,[34] Hong Kong,[29][44] Australia,[53][54] and New Zealand.[55][56]

References

  1. ↑ 1.0 1.1 Jaklitsch, W. M.; Réblová, M. (2015). "Nomenclatural novelties, Savoryellaceae Jaklitsch & Réblová.". Index Fungorum 209: 1. doi:10.1007/978-3-319-23534-9_1. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 Boonyuen, Nattawut; Charuwan, Chuaseeharonnachai; Suetrong, Satinee; Sri-indrasutdhi, Veera; Sivichai, Somsak; Gareth Jones, E.B.; Pang, Ka-Lai (2011). "Savoryellales (Hypocreomycetidae, Sordariomycetes): a novel lineage of aquatic ascomycetes inferred from multiple-gene phylogenies of the genera Ascotaiwania, Ascothailandia, and Savoryella.". Mycologia 103 (6): 1351–1371. doi:10.3852/11-102. PMID 21642338. 
  3. ↑ 3.0 3.1 Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, Dolatabadi; Stadler, Marc; Haelewaters, Danny et al. (2020). "Outline of Fungi and fungus-like taxa". Mycosphere 11: 1060–1456. doi:10.5943/mycosphere/11/1/8. 
  4. ↑ 4.0 4.1 Wijayawardene, N.N.; Hyde, K.D.; Dai, D.Q.; Sánchez-García, M.; Goto, B.T.; Saxena, R.K. et al. (2022). "Outline of Fungi and fungus-like taxa – 2021". Mycosphere 13 (1): 53–453 [160]. doi:10.5943/mycosphere/13/1/2. https://www.researchgate.net/publication/358798332. 
  5. ↑ 5.0 5.1 5.2 Luo, Zong-Long; Hyde, Kevin D.; Liu, Jian-Kui (Jack); Maharachchikumbura, Sajeewa S. N.; Jeewon, Rajesh; Bao, Dan-Feng; Bhat, Darbhe Jayarama; Lin, Chuan-Gen et al. (2019). "Freshwater Sordariomycetes". Fungal Diversity 99: 451–660. doi:10.1007/s13225-019-00438-1. 
  6. ↑ Kohlmeyer, J.; Kohlmeyer, E. (1979). Marine Mycology: the higher fungi.. New York: Academic Press. pp. 690. 
  7. ↑ Eriksson, O.E.; Hawksworth, D.L. (1986). "An alphabetical list of the generic names of ascomycetes.". Systema Ascomycetum 5: 3–111. 
  8. ↑ Eriksson, O.E.; Hawksworth, D.L. (1987). "Notes on ascomycete systematics. Nos 225-463.". Systema Ascomycetum 6: 111–166. 
  9. ↑ 9.0 9.1 Jones, E. B. G.; Hyde, Kevin D. (1992). "Taxonomic studies on Savoryella Jones et Eaton (Ascomycotina).". Bot. Mar. 35 (2): 83–92. doi:10.1515/botm.1992.35.2.83. 
  10. ↑ 10.0 10.1 Read, S.J.; Jones, E.B.G.; Moss, S.T. (1993). "Taxonomic studies of marine Ascomycotina: ultrastructure of the asci, ascospores and appendages of Savoryella species.". Canadian Journal of Botany 71 (2): 273–283. doi:10.1139/b93-028. 
  11. ↑ Vijaykrishna, D.; Jeewon, R.; Hyde, Kevin D. (2006). "Molecular taxonomy, origins and evolution of freshwater ascomycetes.". Fungal Diversity 23: 351–390. 
  12. ↑ 12.0 12.1 Jones, E. B. G.; Sakayaroj, J.; Suetrong, S.; Somrithipol, S.; Pang, K. L. (2009). "Classification of marine Ascomycota, anamorphic taxa and Basidiomycota.". Fungal Divers. 35: 1–187. 
  13. ↑ E. B. Gareth Jones and Ka-Lai Pang (Editors) Marine Fungi: and Fungal-like Organisms (2012), p. 37, at Google Books
  14. ↑ 14.0 14.1 14.2 Gomdola, Deecksha (2 March 2021). "Savoryellales – Facesoffungi number: FoF 02096". https://www.facesoffungi.org/savoryellales/. 
  15. ↑ Hawksworth, D. L. (2011). "A new dawn for the naming of fungi: impacts of decisions made in Melbourne in July 2011 on the future publication and regulation of fungal names.". IMA Fungus 2 (2): 155–162. doi:10.5598/imafungus.2011.02.02.06. PMID 22679600. 
  16. ↑ 16.0 16.1 16.2 16.3 Sri-indrasutdhi, V.; Boonyuen, N.; Suetrong, S.; Chuaseeharonnachai, C.; Sivichai, S.; Jones, E. B. G. (2010). "Wood inhabiting freshwater fungi from Thailand: Ascothailandia gen. et sp. nov., Canalisporium sp. nov. with a key to Canalisporium species (Sordariomycetes Ascomycota).". Mycoscience 51: 411–420. doi:10.1007/s10267-010-0055-6. 
  17. ↑ 17.0 17.1 17.2 Hongsanan, S.; Maharachchikumbura, S.S.; Hyde, Kevin D.; Samarakoon, M. C.; Jeewon, R.; Zhao, Q. (2017). "An updated phylogeny of sordariomycetes based on phylogenetic and molecular clock evidence.". Fungal Divers. 84: 25–41. doi:10.1007/s13225-017-0384-2. 
  18. ↑ Hyde, Kevin D.; Maharachchikumbura, S. S.; Hongsanan, S.; Samarakoon, M. C.; Lücking, R.; Pem, D. (2017). "The ranking of fungi: a tribute to David L. Hawksworth on his 70th birthday.". Fungal Divers. 84: 1–23. doi:10.1007/s13225-017-0383-3. 
  19. ↑ 19.0 19.1 Wijayawardene, N. N.; Hyde, Kevin D.; Lumbsch, H.T.; Liu, J.K.; Maharachchikumbura, S.S.N.; Ekanayaka, A.H.; Tian, Q.; Phookamsak, R. (2018). "Outline of Ascomycota: 2017.". Fungal Diversity 88: 167−263. doi:10.1007/s13225-018-0394-8. 
  20. ↑ Réblová, M.; Seifert, K.A.; Fournier, J.; Štěpánek, V. (2016). "Newly recognized lineages of perithecial ascomycetes: the new orders Conioscyphales and Pleurotheciales.". Persoonia 37: 57−81. doi:10.3767/003158516X689819. PMID 28232761. 
  21. ↑ Maharachchikumbura, Sajeewa S. N.; Hyde, Kevin D.; Jones, E. B. Gareth; McKenzie, E. H. C.; Bhat, Jayarama D.; Dayarathne, Monika C.; Huang, Shi-Ke; Norphanphoun, Chada et al. (2016-06-03). "Families of Sordariomycetes" (in en). Fungal Diversity 79 (1): 1–317. doi:10.1007/s13225-016-0369-6. ISSN 1560-2745. 
  22. ↑ 22.0 22.1 22.2 22.3 Hernández-Restrepo, M.; Gené, J.; Castañeda-Ruiz, R. F.; Mena-Portales, J.; Crous, P. W.; Guarro, J. (2017). "Phylogeny of saprobic microfungi from Southern Europe.". Stud. Mycol. 86: 53–97. doi:10.1016/j.simyco.2017.05.002. PMID 28626275. 
  23. ↑ 23.0 23.1 23.2 23.3 Dayarathne, Monika C.; Maharachchikumbura, Sajeewa S. N.; Gareth Jones, E. B.; Dong, Wei; Devadatha, Bandarupalli; Yang, Jing; Ekanayaka, Anusha H.; De Silva, Wasana et al. (7 May 2019). "Phylogenetic Revision of Savoryellaceae and Evidence for Its Ranking as a Subclass". Front. Microbiol.. Section. Fungi and Their Interactions 10: 840. doi:10.3389/fmicb.2019.00840. PMID 31133992. 
  24. ↑ 24.0 24.1 Jones, E.B.G.; Eaton, R.A. (1969). "Savoryella lignicola gen. et sp. nov. from water cooling towers.". Transactions of the British Mycological Society 52: 161–174. doi:10.1016/S0007-1536(69)80169-5. 
  25. ↑ Tsui, C. K. M.; Hyde, K. D. (2003). "Freshwater mycology.". Fungal Divers. Res. Ser. 10: 1–350. doi:10.2307/1468086. 
  26. ↑ 26.0 26.1 Réblová, M.; Miller, A.N.; Rossman, A.Y.; Seifert, K.A.; Crous, P.W.; Hawksworth, D.L.; Abdel-Wahab, M.A.; Cannon, P.F. et al. (2016). "Recommendations for competing sexual-asexually typified generic names in Sordariomycetes (except Diaporthales, Hypocreales, and Magnaporthales).". IMA Fungus 7 (1): 131–153. doi:10.5598/imafungus.2016.07.01.08. PMID 27433444. 
  27. ↑ Zhang, Shengnan; Hyde, Kevin D.; Liu, Jian-Kui Jack; Jones, E. B. Gareth; Abdel-Wahab, Mohamed (July 2019). "Additions to the genus Savoryella (Savoryellaceae)". Phytotaxa 408 (3): 195–207. doi:10.11646/phytotaxa.408.3.4. 
  28. ↑ 28.0 28.1 28.2 28.3 Sivichai, S.; Hywel-Jones, N. L.; Jones, E. B. G. (1998). "Lignicolous freshwater Ascomycota from Thailand 1. Ascotaiwania sawada and its anamorph state Monotosporella.". Mycoscience 39 (3): 307–311. doi:10.1007/BF02464013. 
  29. ↑ 29.0 29.1 29.2 Ranghoo, V.M.; Hyde, Kevin D. (1998). "Ascomycetes from freshwater habitats: Ascolacicola aquatica gen. et sp. nov. and a new species of Ascotaiwania from wood submerged in a reservoir in Hong Kong.". Mycologia 90 (6): 1055−1062. doi:10.1080/00275514.1998.12027005. 
  30. ↑ 30.0 30.1 Chang, H. S. (2001). "Trichocladium anamorph of Ascotaiwania hsilio and monodictys-like anamorphic states of Ascotaiwania lignicola.". Fungal Sci. 16: 35–38. 
  31. ↑ 31.0 31.1 Fallah, P. M.; Crane, J. L.; Shearer, C. A. (1999). "Freshwater ascomycetes: two new species of Ascotaiwania from North America.". Can. J. Bot. 77 (87–92): 87–92. doi:10.1139/b98-202. 
  32. ↑ Réblová, M.; Seifert, K. A.; Fournier, J.; Štěpánek, V. (2012). "Phylogenetic classification of Pleurothecium and Pleurotheciella gen. nov. and its dactylaria-like anamorph (Sordariomycetes) based on nuclear ribosomal and protein-coding genes.". Mycologia 104 (6): 1299–1314. doi:10.3852/12-035. PMID 22684295. 
  33. ↑ Hyde, Kevin D.; Jones, E.B.G. (1998). "Marine mangrove fungi.". Marine Ecology 9 (1): 15−33. doi:10.1111/j.1439-0485.1988.tb00196.x. 
  34. ↑ 34.0 34.1 Minoura, K.; Muroi, T. (1978). "Some freshwater ascomycetes from Japan.". Transactions of the Mycological Society of Japan 19: 129−134. 
  35. ↑ 35.0 35.1 35.2 Chang, H.S.; Hsieh, S.Y.; Jones, E.B.G.; Read, S.J.; Moss, S.T. (1998). "New freshwater species of Ascotaiwania and Savoryella from Taiwan.". Mycological Research 102 (6): 709−718. doi:10.1017/S0953756297005637. 
  36. ↑ 36.0 36.1 Jones, E. B. G.; To-anun, C.; Suetrong, S.; Boonyuen, N. (2016). "Mycosphere Essays 12. Progress in the classification of the water cooling tower ascomycete Savoryella and a tribute to John Savory: a review.". Mycosphere 7: 570–581. doi:10.5943/mycosphere/7/5/4. 
  37. ↑ Linder, D. H. (1929). "A monograph of the helicosporous Fungi Imperfecti.". Ann. Missouri Bot. Gard. 16 (3): 227–388. doi:10.2307/2394038. https://www.biodiversitylibrary.org/part/35450. 
  38. ↑ Jones, E. B. G.; Suetrong, S.; Sakayaroj, J.; Bahkali, A. H. (2015). "Classification of marine ascomycota, basidiomycota, blastocladiomycota and chytridiomycota.". Fungal Divers. 73: 1–72. doi:10.1007/s13225-015-0339-4. 
  39. ↑ 39.0 39.1 Sivanesan, A.; Chang, H. S. (1992). "Ascotaiwania, a new amphisphaeriaceous ascomycete genus on wood from Taiwan.". Mycol. Res. 96 (6): 481–484. doi:10.1016/S0953-7562(09)81094-0. 
  40. ↑ Ranghoo, V. M.; Hyde, Kevin D.; Liew, E. C. Y.; Spatafora, J. W. (1999). "Family placement of Ascotaiwania and Ascolacicola based on DNA sequences from the large subunit rRNA gene.". Fungal Divers. 2: 159–168. 
  41. ↑ Goh, T. K.; Ho, W. H.; Hyde, Kevin D.; Whitton, S. R.; Umali, T. E. (1998). "New records and species of Canalisporium (Hyphomycetes), with a revision of the genus.". Can. J. Bot. 76: 142–152. doi:10.1139/b97-164. 
  42. ↑ 42.0 42.1 42.2 42.3 Rao, V.; de Hoog, G. S. (1986). "New or critical Hyphomycetes from India.". Stud. Mycol. 28: 1–84. 
  43. ↑ 43.0 43.1 43.2 43.3 Raja, H. A.; Shearer, C. A. (2007). "Freshwater ascomycetes: Aliquandostipite minuta (Jahnulales, Dothideomycetes), a new species from Florida.". Mycoscience 48 (6): 395–398. doi:10.1007/S10267-007-0375-3. 
  44. ↑ 44.0 44.1 44.2 Tsui, Clement K. M.; Hyde, Kevin D.; Hodgkiss, I. J. (2001). "Longitudinal and temporal distribution of freshwater ascomycetes and dematiaceous hyphomycetes on submerged wood in the Lam Tsuen River, Hong Kong.". J. N. Am. Benthol. Soc. 20 (4): 533–549. doi:10.2307/1468086. 
  45. ↑ 45.0 45.1 Holubová-Jechová, V.; Sierra, A. M. (1984). "Studies on hyphomycetes from Cuba II. Hyphomycetes from the Isla de la Juventudeská.". Česká Mykol. 38: 96–120. 
  46. ↑ 46.0 46.1 Kirk, P. M. (1985). "New or interesting microfungi. XIV. Dematiaceous hyphomycetes from Mt. Kenya.". Mycotaxon 23: 305–352. 
  47. ↑ 47.0 47.1 Nawawi, A.; Kuthubutheen, A. J. (1989). "Canalisporium, a new genus of lignicolous hyphomycetes from Malaysia.". Mycotaxon 34: 475–487. doi:10.1139/b97-164. 
  48. ↑ Matsushima, T. (1987). Matsushima Mycological Memoir, No. 5.. 
  49. ↑ "Savoryellales" (in en). https://www.gbif.org/species/7358616. 
  50. ↑ Gönczöl, J. (1976). "Monotosporella tuberculata, a new species of aquatic hyphomycetes from Hungary.". Nova Hedwigia 27: 493–500. 
  51. ↑ B.D. Borse, K.N. Borse, S.Y. Patil, C.M. Pawara, L.C. Nemade and V.R. Patil Freshwater Higher Fungi of India (2016), p. 6, at Google Books
  52. ↑ Du, Hong-Zhi; Yang, Jing; Liu, Ning-Guo; Cheewangkoon, Ratchadawa; Liu, Jian-Kui (2022). "Morpho-Phylogenetic Evidence Reveals New Species of Fuscosporellaceae and Savoryellaceae from Freshwater Habitats in Guizhou Province, China". J. Fungi 8 (11): 1138. doi:10.3390/jof8111138. PMID 36354905. 
  53. ↑ Hyde, Kevin D. (1993). "Tropical Australian freshwater fungi. V.* Bombardia sp., Jahnula australiensis sp. nov., Savoryella aquatica sp. nov. and S. lignicola sp. nov.". Australian Systematic Botany 6 (2): 161−167. doi:10.1071/SB9930161. 
  54. ↑ Abdel-Wahab, M.A.; Jones, E.B.G. (2000). "Three new marine Ascomycetes from driftwood in Australia sand dunes.". Mycoscience 41 (4): 379–388. doi:10.1007/BF02463951. 
  55. ↑ "NZOR Name Details - Savoryellales Boonyuen, Suetrong, S. Sivichai, K.L. Pang & E.B.G. Jones". https://www.nzor.org.nz/names/705d910e-0b4d-400a-85a3-9c58eca9f647. 
  56. ↑ "Savoryellales Boonyuen, Suetrong, S. Sivichai, K.L. Pang & E.B.G. Jones 2011 - Biota of NZ". https://biotanz.landcareresearch.co.nz/scientific-names/91180d0b-2f3e-440f-b9db-2c31e2508198. 

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