Phacopsis

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Phacopsis
Phacopsis vulpina - Flickr - pellaea (3).jpg
Closeup of Phacopsis vulpina gall on Letharia sp.
Scientific classification e
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Parmeliaceae
Genus: Phacopsis
Tul. (1852)
Type species

Tul. (1852)

Phacopsis is a genus of lichenicolous lichens. They grow on parasitically on members of the lichen family Parmeliaceae, of which they are also a member. Phacopsis species appear as partially immersed shiny brown to black apothecia that cause gall-like deformations on the thallus of the host lichen. Phacopsis are distinguished from each other by the shape of their spores, the colour and amyloid staining reaction of the hypothecium (a tissue layer under the spore-bearing hymenium), and the identity of their host lichen.

Systematics[]

Phacopsis was circumscribed by Edmond Tulasne in 1852 as a segregate of genus Abrothallus. Tulasne included three species: P. clemens, P. varia, and P. vulpina.[1] The last of these species was chosen as the type by Frederic Clements and Cornelius Shear (1931),[2] and by John Axel Nannfeldt (1932).[3] Of the other two species originally included in Phacopsis by Tulasne, P. clemens has since been referred to the genus Arthonia,[4] while P. varia is now known as .[5]

In 1988, Dagmar Triebel and Gerhard Rambold proposed that Phacopsis should be considered synonymous with Nesolechia, owing to similarities in their apothecial morphology and hymenial characters.[6] This proposed synonymy, however, was not accepted by several authors in the years following,[7][8][9] until 1995 when Triebel, Rambold, and John Elix showed that the supposed differences that separated the two genera were not consistent.[10] Following this, Ove Eriksson and David L. Hawksworth used an expanded generic concept of Phacopsis in the next update of their regular publication on ascomycete systematics.[11] In a 2018 review, the authors recommended not synonymizing Nesolechia and Phacopsis, suggesting that the separation between these two genera is still poorly known.[12]

The higher-level classification of Phacopsis has changed several times in its taxonomic history. It has been placed in the order Arthoniales,[13] in the families Graphidaceae,[2] and the Acarosporaceae.[14] Using electron microscopy, Josef Hafellner examined the asci (spore-bearing cells) of the type species P. vulpina, showing them to be similar to those commonly found in members of the Lecanoraceae, and so he considered the genus most appropriately classified in this family.[15] This familial placement, however, was not corroborated in Triebel and colleagues' 1995 review and revision of the genus.[10] Molecular analysis eventually revealed its true phylogenetic placement in the family Parmeliaceae. This relationship is an example of adelpho-parasitism, in which the host species is closely related to the parasite;[16] this type of relationship is fairly common in the Lecanorales. The genus Protousnea has a sister taxon relationship with Phacopsis, which has been estimated to have evolved from its ancestors during the Miocene (23.03 to 5.333 mya).[17]

Description[]

Microscopy of Phacopsis vulpina asci. Several spores are visible in some of the asci, and the non-amyloid zone above the axial body is visible in some instances.

All species are endokapylic, meaning they possess a thallus in which no morphologically distinct lichenized structure is formed. Their apothecia are circular to irregular in shape, and may be dispersed or aggregated. The excipulum (the outer margin of the apothecium) range from colourless to blackish-brown, and often comprises "a few rows of cells which may be difficult to recognize". The hypothecium (a layer of dense hyphal tissue just below the hymenium) is colourless to blackish-brown. In some species, the hyphae in the hypothecium have a violet staining reaction with iodine; this characteristic can be used to distinguish between some species. The hymenium is colourless, pale brownish, or pale olivaceous. Asci are more or less club-shaped, and contain eight spores. They are surrounded by a thin outer amyloid wall layer and a thicker, non-amyloid inner wall layer; a non-amyloid zone rests above the axial body of the ascus.[10]

The ascospores are colourless, lack septa, and have smooth walls with occasional thickening at either end. The spores have a range of shapes; depending on the species, the following shapes have been recorded: ellipsoid, ovoid (egg-shaped), fusiform (spindle-shaped), lemon-shaped, falciform (sickle-shaped), fabiform (bean-shaped), and partly curved. The pycnidia of Phacopsis are immersed in the thallus of the host. Pycnospores have a bacilliform shape.[10]

Distribution[]

Members of the host family, Parmeliaceae, are found worldwide and the family has centres of distribution in Asia and in the Southern Hemisphere; Phacopsis probably occurs wherever the host does. A 1995 survey of Phacopsis accepted 13 taxa (eight species and several varieties) found on 41 host species on 20 genera. Many of the Phacopsis species are poorly known, and have only been found at their type locality.[10] The most southernly distributed member of the genus is Phacopsis usneae, known only to occur in southern Chile and Antarctica.[18]

Species[]

Phacopsis species are distinguished from each other by the characteristics of their ascospores, the colour and amyloid reaction of the hypothecium, and the identity of their host.[10] The following list of Phacopsis species indicates its name, taxonomic authority and year of publication, type locality, and host species (or genus).

Several species formerly placed in Phacopsis have since been transferred to other genera. These include:

  • Phacopsis doerfeltii Alstrup & P.Scholz (1998) now (Alstrup & P.Scholz) Diederich (2018)[24]
  • Phacopsis huuskonenii Räsänen (1948) now Raesaenenia huuskonenii (Räsänen) D.Hawksw., C.Boluda & H.Lindgren (2015))[17]
  • Phacopsis lesdainii Vouaux (1914) now (Vouaux) Etayo & Diederich (2000)[25]
  • Phacopsis macrospora Uloth (1861) now (Uloth) Hafellner & Nav.-Ros. (2004)[26]
  • Phacopsis mulleri Willey (1892) now Skyttella mulleri (Willey) D.Hawksw. & R.Sant. (1988)[27]
  • Phacopsis oxyspora (Tul.) Triebel & Rambold (1988) now (Tul.) A.Massal. (1856))

References[]

  1. ^ Tulasne, L.-R. (1852). "Mémoire pour servir à l'histoire organographique et physiologique des Lichens". Annales des Sciences Naturelles Botanique. Série 3 (in Latin). 17: 124.
  2. ^ a b Clements, Frederic E.; Shear, Cornelius L. (1931). The Genera of Fungi (2nd ed.). New York: Hafner Publishing. p. 306.
  3. ^ Nannfeldt, J.A. "Studien über die Morphologie und Systematik der nichtlichenisierten inoperculaten Discomyceten". Nova Acta Regiae Societatis Scientiarum Upsaliensis (in German). 8 (2): 1–368.
  4. ^ "Record Details: Phacopsis clemens Tul., Annls Sci. Nat., Bot., sér. 3 17: 124 (1852)". Index Fungorum. Retrieved 14 May 2021.
  5. ^ "Record Details: Phacopsis varia Tul., Annls Sci. Nat., Bot., sér. 3 17: 125 (1852)". Index Fungorum. Retrieved 14 May 2021.
  6. ^ a b c Triebel, D.; Rambold, G. (1988). "Cecidonia und Phacopsis (Lecanorales): zwei lichenicole Pilzgattungen mit cecidogenen Arten" [Cecidonia and Phacopsis (Lecanorales): two lichenicolous fungus genera with cecidiogen species]. Nova Hedwigia (in German). 47 (3–4): 279–309.
  7. ^ Eriksson, O.E.; Hawksworth, David L. (1989). "Notes on ascomycete systematics. Nos. 804–888". Systema Ascomycetum. 8 (1): 59–86.
  8. ^ Alstrup, V.; Hawksworth, D.L. (1990). "The lichenicolous fungi of Greenland". Meddelelser om Grønland Biosciences. 31: 1–90.
  9. ^ Hafellner, J.; Sancho, L.G. (1990). "Über einige Pilze und Flechten aus den Gebirgen Zentralspaniens und den Ostalpen (mit 8 Abbildungen)". Herzogia (in German). 8 (3–4): 363–382.
  10. ^ a b c d e f g h i Triebel, Dagmar; Rambold, Gerhard; Elix, John A. (1995). "A conspectus of the genus Phacopsis (Lecanorales)". The Bryologist. 98 (1): 71–83. doi:10.2307/3243643. JSTOR 3243643.
  11. ^ Eriksson, O.E.; Hawksworth, David L. (1995). "Notes on ascomycete systematics. Nos. 1885–2023". Systema Ascomycetum. 14: 41–77.
  12. ^ Thell, Arne; Kärnefelt, Ingvar; Seaward, Mark D. (2018). "Splitting or synonymizing – genus concept and taxonomy exemplified by the Parmeliaceae in the Nordic region" (PDF). Graphis Scripta. 30 (6): 130–137. ISSN 2002-4495.
  13. ^ Eriksson, O.E.; Hawksworth, David L. (1987). "An alphabetical list of the generic names of the ascomycetes - 1987". Systema Ascomycetum. 6 (1): 1–109.
  14. ^ Kirk PM, Cannon PF, David JC, Stalpers JA. (2001). Ainsworth & Bisby's Dictionary of the Fungi (9th ed.). Oxon, UK: CABI Bioscience. ISBN 978-0-85199-377-5.CS1 maint: uses authors parameter (link)
  15. ^ Hafellner, Josef (1987). "Studien über lichenicole Pilze und Flechten V. Über die Gattung Phacopsis Tul. emend. Körber (Lecanorales)". Herzogia (in German). 7: 343–352.
  16. ^ Peršoh, Derek; Rambold, Gerhard (2002). "Phacopsis — A lichenicolous genus of the family Parmeliaceae" (PDF). Mycological Progress. 1 (1): 43–55. doi:10.1007/s11557-006-0004-0. S2CID 9935563.
  17. ^ a b Divakar, Pradeep K.; Crespo, Ana; Wedin, Mats; Leavitt, Steven D.; Hawksworth, David L.; Myllys, Leena; et al. (2015). "Evolution of complex symbiotic relationships in a morphologically derived family of lichen-forming fungi". New Phytologist. 208 (4): 1217–1226. doi:10.1111/nph.13553. PMID 26299211.
  18. ^ Alstrup, Vagn; Olech, Maria; Wietrzyk-Pełka, Paulina; Węgrzyn, Michał Hubert (2018). "The lichenicolous fungi of the South Shetland Islands, Antarctica: species diversity and identification guide". Acta Societatis Botanicorum Poloniae. Polish Botanical Society. 87 (4): 22. doi:10.5586/asbp.3607.
  19. ^ Aptroot, André; Triebel, Dagmar (2002). "A new Phacopsis species on Paraparmelia and Xanthoparmelia in southern Africa". Nova Hedwigia. 74 (3–4): 405–409. doi:10.1127/0029-5035/2002/0074-0405.
  20. ^ Zhurbenko, Mikhail P. (2010). "New and interesting lichenicolous fungi from Eurasia. II" (PDF). Mycosphere. 1 (3): 213–222. open access
  21. ^ a b Rambold, G.; Triebel, D. (1992). The inter-lecanoralean associations. Bibliotheca Lichenologica. 48. p. 169.
  22. ^ Dodge, Carroll William (1948). Lichens and Lichen Parasites. Reports (B.A.N.Z. Antarctic Research Expedition, (1929–1931)). Series B, Zoology and botany. 7. Adelaide: Hassell Press. p. 264. OCLC 19094928.
  23. ^ Zhurbenko, Mikhail P.; Enkhtuya, Ochirbat; Javkhlan, Samiya (2019). "A first synopsis of lichenicolous fungi of Mongolia, with the description of five new species". Plant and Fungal Systematics. 64 (2): 345–366. doi:10.2478/pfs-2019-0023.
  24. ^ Diederich, Paul; Lawrey, James D.; Ertz, Damien (2018). "The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa". The Bryologist. 121 (3): 340–425. doi:10.1639/0007-2745-121.3.340. S2CID 92396850.
  25. ^ Etayo, Javier; Diederich, Paul (2000). "Echinodiscus lesdainii gen. et comb. nov., a new name for Phacopsis lesdainii Vouaux (lichenicolous Ascomycetes, Leotiales)". Bulletin de la Société des naturalistes luxembourgeois: 63–66.
  26. ^ "Record Details: Phacopsis macrospora Uloth, Flora, Regensburg 44: 653 (1861)". Index Fungorum. Retrieved 6 November 2021.
  27. ^ Hawksworth, David L.; Santesson, Rolf (1988). "Skyttella, a new genus for Phacopsis mulleri Willey (syn. Agyrium flavescens Rehm)" (PDF). Graphis Scripta. 2 (2): 33–37.
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