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Parmeliaceae

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Parmeliaceae
Parmelia saxatilis
Parmelia saxatilis
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Parmeliaceae
Zenker(1827)
Type genus
Parmelia
Ach.(1803)
Genera[1]

See text

TheParmeliaceaeis a large anddiversefamily ofLecanoromycetes.With over 2700species[2]in 71genera,it is the largest family oflichen-formingfungi.The most speciose genera in the family are the well-known groups:Xanthoparmelia(822 species),Usnea(355 species),Parmotrema(255 species), andHypotrachyna(262 species).[3]

Nearly all members of the family have asymbioticassociation with agreen alga(most oftenTrebouxiaspp., butAsterochlorisspp. are known to associate with some species).[4]The majority of Parmeliaceae species have afoliose,fruticose,or subfruticose growth form. Themorphologicaldiversity and complexity exhibited by this group is enormous, and many specimens are exceedingly difficult to identify down to the species level.

The family has acosmopolitan distribution,and is present in a wide range ofhabitatsandclimatic regions.[5]This includes everywhere from roadside pavement to alpine rocks, from tropical rainforest trees to subshrubs in theArctic tundra.Members of the Parmeliaceae are found in mostterrestrialenvironments. Several Parmeliaceae species have been assessed for the globalIUCN Red List.

Taxonomy

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Based on several molecularphylogeneticstudies, the Parmeliaceae as currently circumscribed has been shown to be amonophyleticgroup.[6]This circumscription is inclusive of the previously described families Alectoriaceae, Anziaceae, Hypogymniaceae, and Usneaceae, which are all no longer recognised by most lichensystematists.However, despite the family being one of the most thoroughly studied groups of lichens, several relationships within the family still remain unclear.Phylogeneticanalysis supports the existence of seven distinctcladesin the family. The Parmelioid clade is the largest, containing 27 genera and about 1850 species – about two-thirds of the species in the family.[7][8]

  • Alectorioid clade (5 genera)
  • Cetrarioid clade (17 genera)
  • Hypogymnioid clade (4 genera)
  • Letharioid clade (2 genera)
  • Parmelioid clade (27 genera)
  • Psiloparmelioid clade (2 genera)
  • Usneoid clade (1 genus)

Many Parmeliaceae genera do not group phylogenetically into any of these clades, and these, along with genera that have not yet had their DNA studied, are classed as "genera with uncertain affinities".[7]

The Parmeliaceae has been divided into two subfamilies, Protoparmelioideae and Parmelioideae.[9][10]Thediversificationof various Parmelioideaelineagesmay have been a result of gaining innovations that providedadaptiveadvantages, such asmelaninproduction in the genusMelanohalea.[11]Diversification of the Protoparmelioideae occurred during theMiocene.[12]The Parmelioid clade is the largest in the Parmeliaceae, with more than 1800 species and a centre of distribution in the Southern Hemisphere.[13]

Evolutionary history

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Although fossil records of extant lichen species are scarce, the existence of someamberinclusions has allowed for a rough estimate of thedivergenceof the Parmeliaceae from itsmost recent common ancestor.AnAnziainclusion from 35–40Myr-oldBaltic amberandParmeliafrom 15–45 Myr-oldDominican ambersuggest a minimum age estimate for the Parmeliaceae of about 40 Myr.[14][15]A fossil-calibrated phylogeny has estimated the Parmeliaceae to have diversified much earlier, around theCretaceous–Paleogene boundary,58–74 Myr ago.[16]

Characteristics

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Thallus

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Parmeliaceae thalli are most often foliose, fruticose or subfruticose, but can be umblicate, peltate, caespitose, crustose, or subcrustose. Two genera,NesolechiaandRaesaenenia,containlichenicolous fungi.They can be a variety of colours, from whitish to grey, green to yellow, or brown to blackish (or any combination therein). Many genera are lobe forming, and nearly all are heteromerous (which arecorticateon both sides). Species are usually rhizinate on the lower surface, occasionally with holdfasts, rhizohyphae, or a hypothallus. Only a few genera have a naked lower surface (for exampleUsnea,HypogymniaandMenegazzia). The upper surface has a pored or non-pored epicortex.Medullais solid, but often loosely woven.[17]

Apothecia

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Apothecia are lecanorine, produced along the lamina or margin, and sessile to pedicellate (or less often sunken). Thalline exciple is concolorous with the thallus. Asci are amyloid, and the vast majority of species have eight spores per ascus, though a few species are many-spored, and severalMenegazziaspecies have two spores per ascus.[17]

Spores

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Ascosporesare simple,hyaline,and often small.Conidiagenerally arise laterally from the joints of conidiogenous hyphae (Parmelia-type), but arise terminally from these joints in a small number of species (Psora-type). The conidia can have a broad range of shapes: cylindrical to bacilliform, bifusiform, fusiform, sublageniform, unciform, filiform, or curved.Pycnidiaare immersed or rarely emergent from the upper cortex, are produced along the lamina or margins, pyriform in shape, and dark-brown to black in colour.[17]

Chemistry

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Members of the Parmeliaceae exhibit a diverse chemistry, with several types oflichenan(Xanthoparmelia-type,Cetraria-type, intermediate-type),isolichenanand/or otherpolysaccharidesbeing known from thecell wallsof many species.[17]The wide diversity in the types of chemical compounds includesdepsides,depsidones,aliphaticacids,triterpenes,anthraquinones,secalonic acids,pulvinic acidderivatives, andxanthones.The compoundsusnic acidandatranorin,which are found exclusively in the Parmeliaceae, are of great importance in thesystematicsof the family, and the presence or absence of these chemicals have been used in several instances to help define genera.ParmeliaandUsneaare the best chemically characterized genera, while the speciesCetraria islandicaandEvernia prunastrihave attracted considerable research attention for theirbioactive compounds.[18]

A study of three parmelioid lichens (Bulbothrix setschwanensis,Hypotrachyna cirrhata,andParmotrema reticulatum) collected from high-altitude areas ofGarhwal Himalaya,showed considerable variation in the chemical content with the rising altitude. This suggests that there is a prominent role for secondary metabolites in the wider ecological distribution of Parmelioid lichens at higher altitudes.[19]

Photobiont

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The mainphotobiontgenus that associates with Parmeliaceae species is thechlorophyteTrebouxia.In particular, the speciesTrebouxia jamesiiappears to be especially prominent. Some Parmeliaceae genera are also known to associate withAsterochloris,[4]but the frequency of this association is not yet known. In general, photobiont diversity within the Parmeliaceae is a little studied subject, and much is left to discover here.

Genera

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These are the genera that are in the Parmeliaceae (including estimated number of species in each genus). Following the genus name is thetaxonomic authority(those who firstcircumscribedthe genus; standardised author abbreviations are used), year of publication, and the estimated number of species.

Conservation

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Parmeliaceae species that have been assessed for the globalIUCN Red Listinclude the following:Anzia centrifuga(vulnerable,2014);[41]Sulcaria badia(endangered,2019);[42]Lethariella togashii(vulnerable, 2017);[43]Hypotrachyna virginica(critically endangered,2020);[44]Sulcaria isidiifera(critically endangered, 2017);[45]Sulcaria spiralifera(endangered, 2020);[46]andXanthoparmelia beccae(vulnerable, 2017).[47]

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References

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  2. ^Lücking, Robert; Hodkinson, Brendan P.; Leavitt, Steven D. (2017). "The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota–Approaching one thousand genera".The Bryologist.119(4): 361–416.doi:10.1639/0007-2745-119.4.361.S2CID90258634.
  3. ^Wijayawardene, Nalin; Hyde, Kevin; L.K.T., Al-Ani; S., 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.hdl:11336/151990.
  4. ^abMiadlikowska, J.et al.(2006). New insights into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota) from phylogenetic analyses of three ribosomal RNA- and two protein-coding genes.Mycologia98:1088-1103.http://www.mycologia.org/cgi/reprint/98/6/1088.pdf
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  8. ^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.hdl:10652/3753.PMID26299211.
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  12. ^Singh, Garima; Dal Grande, Francesco; Schnitzler, Jan; Pfenninger, Markus; Schmitt, Imke (2018)."Different diversification histories in tropical and temperate lineages in the ascomycete subfamily Protoparmelioideae (Parmeliaceae)".MycoKeys(36): 1–19.doi:10.3897/mycokeys.36.22548.PMC6037653.PMID29997448.
  13. ^abCrespo, Ana; Kauff, Frank; Divakar, Pradeep K.; del Prado, Ruth; Pérez-Ortega, Sergio; de Paz, Guillermo Amo; et al. (2010). "Phylogenetic generic classification of parmelioid lichens (Parmeliaceae, Ascomycota) based on molecular, morphological and chemical evidence".Taxon.59(6): 1735–1753.doi:10.1002/tax.596008.
  14. ^Poinar, G.O.; Peterson, E.B.; Platt, J.L. (2000). "FossilParmeliain new World Amber ".The Lichenologist.32(3): 263–269.Bibcode:2000ThLic..32..263P.doi:10.1006/lich.1999.0258.S2CID86227172.
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  16. ^DeSalle, Robert; Amo de Paz, Guillermo; Cubas, Paloma; Divakar, Pradeep K.; Lumbsch, H. Thorsten; Crespo, Ana (2011)."Origin and diversification of major clades in parmelioid lichens (Parmeliaceae, Ascomycota) during the Paleogene inferred by bayesian analysis".PLOS ONE.6(12): e28161.Bibcode:2011PLoSO...628161A.doi:10.1371/journal.pone.0028161.PMC3234259.PMID22174775.
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  40. ^Bystrek, J. (1971). "Taxonomic studies on the genusAlectoria".Annales Universitatis Mariae Curie-Sklodowska.26:265–279.
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  43. ^Ohmura, Y.; Randlane, T.; Spribille, T. (2017)."Lethariella togashii".IUCN Red List of Threatened Species.2017.Retrieved19 April2021.
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