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Lobaria
	Lobaria pulmonaria
Scientific classification
Domain:	Eukaryota
Kingdom:	Fungi
Division:	Ascomycota
Class:	Lecanoromycetes
Order:	Peltigerales
Family:	Peltigeraceae
Genus:	Lobaria; (Schreb.) Hoffm. (1796)
Type species
	Lobaria pulmonaria; (L.) Hoffm. (1796)
Synonyms
	Lichen sect. Lobaria Schreb. (1791); Pulmonaria Hoffm. (1789) [1790]; Reticularia Baumg. (1790); Lobariomyces E.A.Thomas (1939); Ricasoliomyces E.A.Thomas ex Cif. & Tomas. (1953); Stictomyces E.A.Thomas ex Cif. & Tomas. (1953); Anomalobaria Moncada & Lücking (2013);

Lobaria is a genus of foliose lichens, formerly classified in the family Lobariaceae, but now placed in the Peltigeraceae. They are commonly known as "lung wort" or "lungmoss" as their physical shape somewhat resembles a lung, and their ecological niche is similar to that of moss.

Lobaria are unusual in that they have a three-part symbiosis, containing a fungus, and an alga (as other lichens do), but also a cyanobacterium that fixes nitrogen.

Taxonomy

Lobaria was originally described as a section of the eponymous genus Lichen by German naturalist Johann Christian Daniel von Schreber in 1786. It was proposed as a genus by Georg Franz Hoffmann in 1796.^[2] The establishment of Lobaria remained uncertain until Edvard Vainio also described it. He divided the genus into two sections based on different morphologies of the mature spore: Lobaria and Ricasolia.^[3] In 2013, the concept of family Lobariaceae was revised with the help of molecular phylogenetics, and, in addition to the creation of several new genera, Ricasolia was promoted to generic status.^[4] The family Lobariaceae was synonymized with the Peltigeraceae in 2018.^[5]

Ecology

Lichenicolous fungi that have been found growing on Lobaria species include Stigmidium lobariae,^[6] Calycina alstrupii,^[7] and Abrothallus halei.^[8]

A Lobaria-associated actinobacterium, Subtercola lobariae, was isolated from L. retigera collected from the Jiaozi Snow Mountain in Yunnan Province, China.^[9] About a third of the bacteria found colonizing the thallus surface of Lobaria pulmonaria were found to belong to the Rhizobiales. This order of bacteria is well-known in their role as beneficial partners in plant-microbe interactions. Advantages conferred by the presence of the bacteria include auxin and vitamin production, nitrogen fixation, and stress protection. Although the bacteria were most prevalent on the thallus surface, they were shown to be able to penetrate into the interhyphal gelatinous matrix of the upper lichen cortical layer. Occasionally, some bacteria colonize the interior of the fungal hyphae.^[10]

Hydration traits determine much of a lichen's distribution pattern along a climatic gradient. A study demonstrated that Lobaria amplissima thalli with external cephalodia need more rain than thalli without, consistent with reports of decreasing frequency of external cephalodia from wet to drier climates.^[11]

A study using ecological niche modelling of occurrence data of three Lobaria species found in Italy predicts that climate change will impact their distribution range across the country and that there is a high extinction risk resulting from reduction of their range.^[12]

Evolutionary history

A fossil impression found in a 12–24 Myr-old Miocene deposit from Trinity County in northern California^[13] has strong similarities to extant species of Lobaria, particularly L. pulmonaria and other species with reticulated edges, including L. anomala and L. retigera.^[14] Using a molecular clock-calibrated phylogeny to obtain a time estimate for Lobaria yielded a stem age (the time that that clade descended from a common ancestor with its sister clade) of nearly 30 Mya. The evidence suggests that the paleoclimate and the closing or opening of the Bering Strait played a significant role in determining the distribution of most Lobaria species.^[15]

Species

Lobaria anomala (Brodo & Ahti) T.Sprib. & McCune (2014)
Lobaria anthraspis (Ach.) T.Sprib. & McCune (2014)
Lobaria discolor (Bory) Hue (1901)
Lobaria endochroma Sipman (2004)^[16] – New Guinea
Lobaria hartmannii (Müll.Arg.) Zahlbr. (1925)
Lobaria hengduanensis C.C.Miao & Li S.Wang (2018)^[17] – China
Lobaria hertelii Sipman (2004)^[16] – New Guinea
Lobaria himalayensis Upreti & Divakar (2008)^[18] – India
Lobaria irrugulosa C.C.Miao & Li S.Wang (2018)^[17] – China
Lobaria isidiophora Yoshim. (1971)^[19] – Asia
Lobaria latilobulata C.C.Miao & Li S.Wang (2018)^[17] – China
Lobaria macaronesica C.Cornejo & Scheid. (2010)^[20] – Macaronesia
Lobaria oregana (Tuck.) Müll.Arg. (1889)
Lobaria orientalis (Asahina) Yoshim. (1969) – Asia
Lobaria plurimiseptata (C.Knight) Zahlbr. (1925)
Lobaria pseudoretigera Sipman (2004)^[16] – New Guinea
Lobaria pulmonaria (L.) Hoffm. (1796)
Lobaria retigera (Bory) Trevis. (1869)
Lobaria rhaphispora (C.Knight) Zahlbr. (1925)
Lobaria spathulata (Inumaru) Yoshim. (1971)

Several species formerly classified in Lobaria have been transferred to other genera in view of modern molecular phylogenetic studies. Examples include Lobaria quercizans and Lobaria amplissima (now in Ricasolia), and Lobaria scrobiculata (now in Lobarina).

Chemistry

Retigeranic acid is a sesterterpene compound isolated from Lobaria retigera.^[21] The ethyl acetate extract of Lobaria orientalis collected in Central Vietnam led to the isolation of new β-orcinol depsidones, lobarientalones A and B, and several diphenyl ethers, lobariethers A–E.^[22]

Uses

Three species of Lobaria are used as food by ethnic peoples in Yunnan Province (China): L. isidiophora, L. kurokawae, and L. yoshimurae.^[23]

References

^ "Synonymy: Lobaria (Schreb.) Hoffm". Species Fungorum. Retrieved 23 March 2022.
^ Hoffmann GF. Deutschlands Flora: oder, botanisches Taschenbuch für das Iahr. Erlangen: Bey Iohann lacob Palm;1796. p. 1–14
^ Wainio EA. Etude sur la classification naturelle et lamorphologie des lichens du Brésil. Acta Soc Fauna Fla Fenn1890;7:1-247
^ Moncada, Bibiana; Lücking, Robert; Betancourt-Macuase, Luisa (2013). "Phylogeny of the Lobariaceae (lichenized Ascomycota: Peltigerales), with a reappraisal of the genus Lobariella". The Lichenologist. 45 (2): 203–263. doi:10.1017/S0024282912000825. S2CID 86082520.
^ Kraichak, Ekaphan; Huang, Jen-Pan; Nelsen, Matthew; Leavitt, Steven D.; Lumbsch, H. Thorsten (2018). "A revised classification of orders and families in the two major subclasses of Lecanoromycetes (Ascomycota) based on a temporal approach". Botanical Journal of the Linnean Society. 188 (3): 233–249. doi:10.1093/botlinnean/boy060.
^ Zhurbenko, M.P.; Etayo, J. (2012). "Stigmidium lobariae, a new lichenicolous fungus from the Holarctic". Mycosphere. 3 (1): 62–64. doi:10.5943/mycosphere/3/1/7.
^ Suija, Ave; Motiejūnaitė, Jurga (2017). "Calycina alstrupii sp. nov. (Pezizellaceae, Helotiales), a new lichenicolous fungus from Norway". Phytotaxa. 307 (2): 113–122. doi:10.11646/phytotaxa.307.2.2.
^ Suija, Ave; Pérez-Ortega, Sergio; Hawksworth, David L. (2010). "Abrothallus halei (Ascomycota, incertae sedis), a new lichenicolous fungus on Lobaria species in Europe and North America". The Lichenologist. 43 (1): 51–55. doi:10.1017/S002428291000054X. S2CID 86335273.
^ Si, Hong-Li; Shi, Fei-Xiang; Zhang, Lu-Lu; Yue, Hong-Shan; Wang, Hai-Ying; Zhao, Zun-Tian (2017). "Subtercola lobariae sp. nov., an actinobacterium of the family Microbacteriaceae isolated from the lichen Lobaria retigera". International Journal of Systematic and Evolutionary Microbiology. 67 (5): 1516–1521. doi:10.1099/ijsem.0.001753. PMID 28005519.
^ Erlacher, Armin; Cernava, Tomislav; Cardinale, Massimiliano; Soh, Jung; Sensen, Christoph W.; Grube, Martin; Berg, Gabriele (2015). "Rhizobiales as functional and endosymbiontic members in the lichen symbiosis of Lobaria pulmonaria L." Frontiers in Microbiology. 6: 53. doi:10.3389/fmicb.2015.00053. PMC 4322706. PMID 25713563.
^ Gauslaa, Yngvar; Johlander, Stina; Nordén, Björn (2019). "Lobaria amplissima thalli with external cephalodia need more rain than thalli without". The Lichenologist. 51 (3): 281–286. doi:10.1017/S0024282919000197. S2CID 196685184.
^ Nascimbene, Juri; Casazza, Gabriele; Benesperi, Renato; Catalano, Immacolata; Cataldo, Daniela; Grillo, Maria; Isocrono, Deborah; Matteucci, Enrica; Ongaro, Silvia; Potenza, Giovanna; Puntillo, Domenico; Ravera, Sonia; Zedda, Luciana; Giordani, Paolo (2016). "Climate change fosters the decline of epiphytic Lobaria species in Italy". Biological Conservation. 201: 377–384. doi:10.1016/j.biocon.2016.08.003. hdl:11585/587210.
^ MacGinitie, H. D. (1937) The flora of the Weaverville beds of Trinity County, California. Carnegie Institution of Washington Publication 465: 83–151.
^ Peterson, E.B. (2000). "An overlooked fossil lichen (Lobariaceae)". The Lichenologist. 32 (3): 298–300. doi:10.1006/lich.1999.0257. S2CID 83629248.
^ Cornejo, Carolina; Scheidegger, Christoph (2018). "Estimating the timescale of Lobaria diversification". The Lichenologist. 50 (1): 113–121. doi:10.1017/S0024282917000676. S2CID 89794079.
^ ^a ^b ^c Sipman, H.J.M. (2004). "The species of Lobaria (lichenized Ascomycetes) in New Guinea". Bibliotheca Lichenologica. 88: 573–606.
^ ^a ^b ^c Miao, C.C.; Wang, X.Y.; Scheidegger, C.; Wang, L.S.; Zhao, Z.T. (2018). "Three new cyanobacterial species of Lobaria (Lobariaceae, Peltigerales) from the Hengduan Mountains, China". Mycosystema. 37 (7): 838–848. doi:10.13346/j.mycosystema.180018.
^ Upreti, D.K.; Divakar, P.K. (2008). "Notes on some interesting macrolichens from India". Nova Hedwigia. 86 (3): 525–528. doi:10.1127/0029-5035/2008/0086-0525. S2CID 84359249.
^ Yoshimura, I. (1971). "The genus Lobaria of Eastern Asia". Journal of the Hattori Botanical Laboratory. 34: 231–364.
^ Cornejo, Carolina; Scheidegger, Christoph (2010). "Lobaria macaronesica sp. nov., and the phylogeny of Lobaria sect. Lobaria (Lobariaceae) in Macaronesia". The Bryologist. 113 (3): 590–604. doi:10.1639/0007-2745-113.3.590. S2CID 84925065.
^ Kaneda, Miyuki; Takahashi, Rumiko; Iitaka, Yoichi; Shibata, Shoji (1972). "Retigeranic acid, a novel sesterterpene isolated from the lichens of group". Tetrahedron Letters. 13 (45): 4609–4611. doi:10.1016/S0040-4039(01)94378-3.
^ Nguyen, Dung M.T.; Do, Lien M.T.; Nguyen, Vy T.; Chavasiri, Warinthorn; Mortier, Jacques; Nguyen, Phung P.K. (2017). "Phenolic compounds from the lichen Lobaria orientalis". Journal of Natural Products. 80 (2): 261–268. doi:10.1021/acs.jnatprod.6b00465. PMID 28181807.
^ Wang, Li-song; Narui, Takao; Harada, Hiroshi; Culberson, Chicita F.; Culberson, William Louis (2001). "Ethnic uses of lichens in Yunnan, China". The Bryologist. 104 (3): 345–349. doi:10.1639/0007-2745(2001)104[0345:EUOLIY]2.0.CO;2. S2CID 86316534.