Pannaria
Pannaria | |
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Pannaria conoplea | |
Scientific classification ![]() | |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Lecanoromycetes |
Order: | Peltigerales |
Family: | Pannariaceae |
Genus: | Delise ex Bory (1828) |
Type species | |
Pannaria rubiginosa (Thunb.) Delise (1828)
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Synonyms[1] | |
Pannaria is a genus of lichen-forming fungi in the family Pannariaceae.[2] These lichens form leaf-like, scaly, or crusty patches that typically arrange themselves in loose rosettes on tree bark in humid forests. Most species partner with nitrogen-fixing cyanobacteria, though some contain green algae instead. The genus was established in 1828 and now includes about 80 recognized species found worldwide, particularly in tropical and subtropical regions.
Description
Pannaria forms a thallus that can be leaf-like (foliose), a mosaic of tiny scales (squamulose), or a thin crust (crustose). Many species arrange their lobes in loose rosettes and, unlike several related genera, they rarely develop the blue-black felt (hypothallus) seen beneath other cyanobacterial lichens. The upper surface ranges from grey-blue to deep brown-black and may carry a light dusting of frost-like crystals (pruina). Most species partner with the filamentous cyanobacterium Nostoc, but members of the Pannaria sphinctrina group instead contain green algal cells.[3]
The lichen's sexual fruiting bodies are sessile apothecia whose red-brown to black discs are framed by a persistent rim of thallus tissue. That rim has two zones: an outer layer of tightly packed, brick-like cells and an inner layer that is looser and packed with photobiont cells. A thin, pale exciple of pseudoparenchymatous cells surrounds the hymenium. The asci hold eight ascospores, show no blue reaction to potassium–iodide stain (K/I–) and lack the amyloid plug found in many related taxa. Their spores are colourless, single-celled, ellipsoidal and often end in one or two small points; the outer wall is finely warted or ridged.[3]
Asexual reproduction is limited to scattered pycnidia that release straight, rod-shaped conidia. Thin-layer chromatography usually detects no secondary metabolites, though some species contain pannarin, which produces an orange-red colour with the para-phenylenediamine (Pd) spot test.[3]
Photobiont
Like most members of the Pannariaceae, species of Pannaria partner with filamentous, nitrogen-fixing cyanobacteria belonging to the genus Nostoc. A multilocus survey of 37 thalli representing 21 species showed that these cyanobionts are drawn from two broad Nostoc lineages previously known as the "Nephroma guild" and the "Peltigera guild"; the Pannaria sequences are scattered across both, with a gradual transition between them.[4]
Host choice is flexible. Bipartite (cyanobacteria-only) and tripartite (cyanobacteria plus green-algal) species do not segregate cleanly by photobiont: some tripartite southern-hemisphere taxa share identical Nostoc strains with corticolous (bark-dwelling) bipartite species from both hemispheres, whereas other Pannaria species show marked selectivity, keeping to a narrow subset of strains. This breadth of associations contrasts with the tighter photobiont fidelity reported for genera such as Nephroma and Peltigera.[4]
Ecology
Species of Pannaria grow mainly as epiphytes on the bark of trees in sheltered, humid forests. They are encountered far less often on shaded rocks or on thin, moss-covered soil, where the micro-climate is similarly cool and moist.[3]
Species
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As of July 2025, Species Fungorum (in the Catalogue of Life) accept 78 species of Pannaria:[5]
- Pannaria adpressa Zahlbr. (1930)
- Pannaria aenea Müll.Arg. (1896)[6]
- Pannaria andina P.M.Jørg. & Sipman (2004)[7]
- Pannaria allorhiza (Nyl.) Elvebakk & D.J.Galloway (2003)
- Pannaria aotearoana Elvebakk & Elix (2016)[8]
- Pannaria applanata Eckfeldt (1894)[9]
- Pannaria areolata Delise (1828)
- Pannaria asahinae P.M.Jørg. (2001)[10]
- Pannaria athroophylla (Stirt.) Elvebakk & D.J.Galloway (2003)
- Pannaria atrofumosa C.Knight (1880)
- Pannaria auctorum Bory (1828)
- Pannaria brasiliensis Zahlbr. (1909)
- Pannaria byssoidea Passo & Calvelo (2011)[11]
- Pannaria caesiocinerea (Vain.) P.M.Jørg. (2004)
- Pannaria caespitosa P.M.Jørg. (2000)[12]
- Pannaria calophylla (Müll.Arg.) Passo & Calvelo (2006)[13]
- Pannaria cameroonensis C.W.Dodge (1964)
- Pannaria carneopallens Vain. (1921)
- Pannaria cassa Elvebakk (2022)
- Pannaria centrifuga P.M.Jørg. (2001)[14]
- Pannaria cheirolepis F.Wilson (1889)
- Pannaria chilensis Fée (1852)
- Pannaria cinerascens (Nyl.) P.M.Jørg. (2004)
- Pannaria complanata P.M.Jørg. (2001)[15]
- Pannaria conoplea (Ach.) Bory (1828)
- Pannaria contorta (Müll.Arg.) Passo & Calvelo (2006)[13]
- Pannaria craspedia Körb. (1859)
- Pannaria crenulata P.M.Jørg. (1983)[16]
- Pannaria crispella (Nyl.) Elvebakk (2022)
- Pannaria crustata Stirt. (1873)[17]
- Pannaria decipiens P.M.Jørg. & D.J.Galloway (1992)
- Pannaria delicata P.M.Jørg. & D.J.Galloway (1999)[18]
- Pannaria delisei Bory (1838)
- Pannaria dissecta P.M.Jørg. (2001)[14]
- Pannaria durietzii (P.James & Henssen) Elvebakk & D.J.Galloway (2003)
- Pannaria ekistophylla Colmeiro (1867)
- Pannaria elatior Stirt. (1899)
- Pannaria elegantior P.M.Jørg. (2003)[19]
- Pannaria elixii P.M.Jørg. & D.J.Galloway (1992)
- Pannaria emodii P.M.Jørg. (2001)[15]
- Pannaria euphylla (Nyl.) Elvebakk & D.J.Galloway (2003)
- Pannaria exasperata H.Magn. (1944)
- Pannaria farinosa Elvebakk & Fritt-Rasm. (2007)[20]
- Pannaria fimbriata P.M.Jørg. (2001)[14]
- Pannaria flabellata P.M.Jørg. (2012)[21]
- Pannaria formosana P.M.Jørg. (2001)[15]
- Pannaria fulvescens (Mont.) Nyl. (1857)
- Pannaria fumbris Kremp. (1874)
- Pannaria funebris Kremp. (1874)
- Pannaria gallowayi Elvebakk & Elix (2016)[8]
- Pannaria glacialis Anzi (1860)
- Pannaria granulifera Müll.Arg. (1896)
- Pannaria globigera Hue (1909)
- Pannaria holospoda Nyl. (1888)
- Pannaria hookeri (Borrer) Nyl. (1857)
- Pannaria howeana Elvebakk (2012)[22]
- Pannaria hypnorum (Vahl) Körb. (1855)
- Pannaria immixta Nyl. (1867)[23]
- Pannaria implexa (Stirt.) Passo, Calvelo & S.Stenroos (2008)
- Pannaria insularis P.M.Jørg. & Kashiw. (2001)[10]
- Pannaria isidiosa Elvebakk & Elix (2006)[24]
- Pannaria italica Gyeln. (1940)
- Pannaria japonica Räsänen (1940)
- Pannaria kantvilasii Elvebakk (2022)
- Pannaria kerguelensis C.W.Dodge (1966)
- Pannaria laciniata Zahlbr. (1941)
- Pannaria laciniosa Hue (1902)
- Pannaria lanuginosa (Hoffm.) Szatala (1930)
- Pannaria lasiella Stirt. (1878)
- Pannaria lepidophora Vain. (1921)
- Pannaria leproloma (Nyl.) P.M.Jørg. (2001)[14]
- Pannaria limbata Vain. (1921)
- Pannaria lobulifera Elvebakk (2007)[25]
- Pannaria lurida (Mont.) Nyl. (1857)
- Pannaria luridula Nyl. (1876)
- Pannaria macrocarpa Müll.Arg. (1892)
- Pannaria malmei C.W.Dodge (1933)
- Pannaria mangroviana P.M.Jørg. (2001)[14]
- Pannaria melanesica Elvebakk (2020)
- Pannaria melanotricha Müll.Arg. (1885)
- Pannaria microphyllizans (Nyl.) P.M.Jørg. (2001)[14]
- Pannaria minutiphylla Elvebakk (2013)[26]
- Pannaria molkenboeri (Mont. & Bosch) Hue (1902)
- Pannaria molybdodes F.Wilson (1889)
- Pannaria mosenii C.W.Dodge (1933)
- Pannaria multifida P.M.Jørg. (2004)
- Pannaria myrioloba Müll.Arg. (1896)
- Pannaria neocaledonica B.de Lesd. (1910)
- Pannaria nilgherriensis P.M.Jørg. & Upreti (2003)
- Pannaria obscura Müll.Arg. (1895)
- Pannaria oregonensis McCune & M.Schultz (2022)[27]
- Pannaria pannosa (Sw.) Nyl. (1855)
- Pannaria papuana (Aptroot & Diederich) P.M.Jørg. & Sipman (2006)
- Pannaria parmeliae F.Wilson (1889)
- Pannaria patagonica (Malme) Elvebakk & D.J.Galloway (2003)
- Pannaria perfossa Stirt. (1875)
- Pannaria phloeodes Stirt. (1877)
- Pannaria phyllidiata Elvebakk (2011)
- Pannaria placodioides Nyl. ex Cromb. (1875)
- Pannaria placodiopsis Nyl. (1875)
- Pannaria planiuscula P.M.Jørg. (2003)[28]
- Pannaria prolifera Müll.Arg. (1882)
- Pannaria prolificans Vain. (1896)
- Pannaria pruinosa P.M.Jørg. & Timdal (2004)[29]
- Pannaria pulverulacea Elvebakk (2013)[26]
- Pannaria pulvinula P.M.Jørg. (2000)[12]
- Pannaria pyxinoides (Nyl.) Elvebakk (2018)
- Pannaria ramosii Vain. (1921)[30]
- Pannaria ramulosa P.M.Jørg. (2001)[15]
- Pannaria reflectens (Nyl.) P.M.Jørg. (2010)
- Pannaria rolfii Elvebakk (2012)[31]
- Pannaria romanoana Hue (1915)
- Pannaria rubiginea Nyl. (1859)
- Pannaria rubiginella P.M.Jørg. & Sipman (2004)[7]
- Pannaria rubiginosa (Thunb. ex Ach.) Delise (1828)
- Pannaria squamulosa P.M.Jørg. (2003)[28]
- Pannaria streimannii Elvebakk (2012)[22]
- Pannaria subcrustacea (Räsänen) P.M.Jørg. (2001)[14]
- Pannaria subfusca P.M.Jørg. (2000)[32]
- Pannaria subsimilis C.Knight (1880)
- Pannaria superior Nyl. (1868)
- Pannaria taylorii Tuck. (1875)
- Pannaria tavaresii P.M.Jørg. (1978)[33]
- Pannaria tenuis P.M.Jørg. & Sipman (2006)
- Pannaria tetraspora Maheu & A.Gillet (1926)
- Pannaria thoroldii C.W.Dodge (1964)
- Pannaria thraustolepis F.Wilson (1889)
- Pannaria tjibodensis Zahlbr. (1928)[34]
- Pannaria vischii C.W.Dodge (1971)
- Pannaria wrightiorum Elvebakk (2022)
References
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- ^ Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Dolatabadi, Somayeh; 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. hdl:10481/61998.
- ^ a b c d Cannon, P.; Aptroot, A.; Coppins, B.; Sanderson, N.; Simkin, J. (2021). Peltigerales: Pannariaceae, including the genera Fuscopannaria, Leptogidium, Nevesia, Pannaria, Parmeliella, Pectenia, Protopannaria and Psoroma (PDF). Revisions of British and Irish Lichens. Vol. 9. p. 7.
- ^ a b Elvebakk, Arve; Papaefthimiou, Dimitra; Robertsen, Eli Helene; Liaimer, Anton (2008). "Phylogenetic patterns among Nostoc cyanobionts within bi- and tripartite lichens of the genus Pannaria". Journal of Phycology. 44 (4): 1049–1059. doi:10.1111/j.1529-8817.2008.00556.x. PMID 27041623.
- ^ "Pannaria". Catalogue of Life. Species 2000: Leiden, the Netherlands. Retrieved 14 July 2025.
- ^ Müller, J. (1896). "Analecta Australiensia". Bulletin de l'Herbier Boissier. 4: 87–96.
- ^ a b Jørgensen, P.M.; Sipman, H.J.M. (2004). "A revision of the Pannaria rubiginosa complex in South America". Nova Hedwigia. 78 (3–4): 311–327. Bibcode:2004NovaH..78..311J. doi:10.1127/0029-5035/2004/0078-0311.
- ^ a b Elvebakk, A.; Elix, J.A. (2016). "A trio of endemic New Zealand lichens: Pannaria aotearoana and P. gallowayi, new species with a new chemo-syndrome, and their relationship with P. xanthomelana". Nova Hedwigia. 105 (1–2): 167–184. doi:10.1127/nova_hedwigia/2016/0385.
- ^ Eckfeldt, J.W. (1894). "Lichens new to North America". Bulletin of the Torrey Botanical Club. 21 (9): 393–396. doi:10.2307/2478240. JSTOR 2478240.
- ^ a b Jørgensen, P.M.; Kashiwadani, H. (2001). "New and misunderstood species of Japanese Pannaria (lichenes)". Journal of Japanese Botany. 76 (1): 1–10.
- ^ Passo, Alfredo; Calvelo, Susana (2011). "Pannaria byssoidea (Pannariaceae), a new squamulose species from southern South America". The Bryologist. 114 (4): 756–763. doi:10.1639/0007-2745-114.4.756.
- ^ a b Jørgensen, Per M. (2000). "Studies in the lichen family. Pannariaceae IX. A revision of Pannaria subg. Chryopannaria". Nova Hedwigia. 71 (3–4): 405–411. Bibcode:2000NovaH..71..405J. doi:10.1127/nova/71/2000/405.
- ^ a b Passo, Alfredo; Calvelo, Susana (2006). "New reports and combinations in the family Pannariaceae (Lecanorales, lichenized Ascomycota)". The Lichenologist. 38 (6): 549–555. Bibcode:2006ThLic..38..549P. doi:10.1017/S0024282906005688.
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- ^ a b c d Jørgensen, Per M. (2001). "Four new Asian species in the lichen genus Pannaria". The Lichenologist. 33 (4): 297–302. Bibcode:2001ThLic..33..297J. doi:10.1006/lich.2001.0333.
- ^ Galloway, J.D.; James, P.W.; Jørgensen, P.M. (1983). "Pannaria crenulata, a new lichen from New Zealand". New Zealand Journal of Botany. 21: 101–104. doi:10.1080/0028825X.1983.10428544.
- ^ Stirton, J. (1873). "Additions to the lichen flora of New Zealand". Transactions of the Glasgow Society of Field Naturalists. 1: 15–23.
- ^ Jørgensen, P.M. (1999). "Studies in the lichen family Pannariaceae VIII. Seven new parmelielloid lichens from New Zealand". New Zealand Journal of Botany. 37 (2): 257–268. Bibcode:1999NZJB...37..257J. doi:10.1080/0028825X.1999.9512632.
- ^ Jørgensen, Per M. (2003). "Studies in the lichen family Pannariaceae XI. The isidiate species of the genus Pannaria Del". Nova Hedwigia. 76 (1–2): 245–255. Bibcode:2003NovaH..76..245J. doi:10.1127/0029-5035/2003/0076-0245.
- ^ Elvebakk, Arve; Fritt-Rasmussen, Janne; Elix, John A. (2007). "The New Zealand lichen Pannaria leproloma (Nyl.) P. M. Jørg. and its panaustral relative P. farinosa nom. nov". The Lichenologist. 39 (4): 349–359. Bibcode:2007ThLic..39..349E. doi:10.1017/s0024282907006913.
- ^ Jørgensen, P.M.; Gjerde, I. (2012). "Notes on some pannariaceous lichens from New Caledonia". Cryptogamie, Mycologie. 33 (1): 3–9. doi:10.7872/crym.v33.iss1.2012.003.
- ^ a b Elvebakk, Arve (2012). "Pannaria howeana and Pannaria streimannii, two related new lichen species endemic to Lord Howe Island, Australia". The Lichenologist. 44 (4): 457–463. Bibcode:2012ThLic..44..457E. doi:10.1017/s0024282912000047.
- ^ Nylander, W. (1865). "Lichenes Novae Zelandiae, quos ibi legit anno 1861 Dr. Lauder Lindsay". Journal of the Linnean Society. Botany (in Latin). 9 (36): 244–259. doi:10.1111/j.1095-8339.1866.tb01283.x.
- ^ Elvebakk, Arve; Elix, John A. (2006). "Pannaria isidiosa, a new Australian lichen with a new chemosyndrome". The Lichenologist. 38 (6): 557–563. Bibcode:2006ThLic..38..557E. doi:10.1017/s0024282906006141.
- ^ Elvebakk, A. (2007). "The panaustral lichen Pannaria sphinctrina (Mont.) Tuck. and the related new species P. lobulifera from New Caledonia". Cryptogamie, Mycologie. 28 (3): 225–235.
- ^ a b Elvebakk, Arve (2013). "Pannaria minutiphylla and P. pulverulacea, two new and common, austral species, previously interpreted as Pannaria microphyllizans (Nyl.) P. M. Jørg". The Lichenologist. 45 (1): 9–20. Bibcode:2013ThLic..45....9E. doi:10.1017/s0024282912000679.
- ^ McCune, Bruce; Schultz, Matthias; Fennell, Terry; Passo, Alfredo; Rodriguez, Juan Manuel (2022). "A new endemic, Pannaria oregonensis, replaces two misapplied names in the Pacific Northwest of North America". The Bryologist. 125 (1). doi:10.1639/0007-2745-125.1.170.
- ^ a b Jørgensen, Per M. (2003). "Notes on African Pannariaceae (lichenized ascomycetes)". The Lichenologist. 35 (1): 11–20. Bibcode:2003ThLic..35...11J. doi:10.1006/lich.2002.0424.
- ^ Jørgensen, P.M. (2004). "Further contributions to the Pannariaceae (lichenized Ascomycetes) of the Southern Hemisphere". Bibliotheca Lichenologica. 88: 229–253.
- ^ Vainio, E.A. (1921). "Lichenes insularum Philippinarum, III". Annales Academiae Scientiarum Fennicae (in Latin). 15 (6): 16.
- ^ Elvebakk, Arve (2012). "Pannaria rolfii, a new name for a recently described lichen species". Nova Hedwigia. 94 (3–4): 505–506. Bibcode:2012NovaH..94..505E. doi:10.1127/0029-5035/2012/0011.
- ^ Jørgensen, Per M. (2000). "Survey of the lichen family Pannariaceae on the American continent, north of Mexico". The Bryologist. 103 (4): 670–704. doi:10.1639/0007-2745(2000)103[0670:sotlfp]2.0.co;2.
- ^ Jørgensen, P.M. (1978). "The lichen family Pannariaceae in Europe". Opera Botanica. 45: 1–124.
- ^ Zahlbruckner, A. (1928). "Neue und ungenügend beschriebene javanische Flechten" [New and insufficiently described Javanese lichens]. Annales de Cryptogamie Exotique (in German). 1: 109–212 [165].