Cladoniaceae

The Cladoniaceae are a family of lichenized fungi in the order Lecanorales.[2] It is one of the largest families of lichen-forming fungi, with about 560 species distributed amongst 17 genera. The reindeer moss and cup lichens (Cladonia) belong to this family. The latter genus, which comprises about 500 species, forms a major part of the diet of large mammals in taiga and tundra ecosystems.[5]

Cladoniaceae
Cladonia subuluata
Scientific classification
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Cladoniaceae
Zenker (1827)[1]
Type genus
Cladonia
P.Browne (1756)
Synonyms[2]
  • Cetradoniaceae J.C.Wei & Ahti (2002)[3]
  • Squamarinaceae Hafellner (1984)[4]

Taxonomy

Cladoniaceae was formally introduced to science in 1827 by German naturalist Jonathan Carl Zenker in a publication of Karl Goebel.[1] It is one of the largest families of lichen-forming fungi,[6] with about 560 species distributed amongst 18 genera. The type genus, after which the family was named, is Cladonia, circumscribed by Irish physician and botanist Patrick Browne in 1756. He included 8 species in his new genus. Of their occurrence, he wrote: "All these species are found in great abundance in the mountains of Liguanea: they grow mostly on the ground, among other sorts of moss, but a few ... species chiefly are found upon the decaying trunks of trees."[7]

Synonymy

Several phylogenetic studies have shown that Cladoniaceae is a member of the order Lecanorales, and is closely related to the family Stereocaulaceae.[8][9][10][11][12] The family Cetradoniaceae, which was created in 2002 to contain the endangered species Cetradonia linearis, was folded into the Cladoniaceae in 2006.[13]

In 2018, Kraichak and colleagues used a technique called temporal banding to reorganize the Lecanoromycetes, proposing a revised system of classification based on correlating taxonomic rank with geological (evolutionary) age. They synonymized the families Squamarinaceae and Stereocaulaceae with the Cladoniaceae, resulting in a large increase in the number of genera and species.[14] The Squamarinaceae had already been included in the Cladoniaceae by previous authors.[6] Although this reorganization has been used in some later publications,[2] the folding of the Stereocaulaceae into the Cladoniaceae was not accepted in a recent analysis. As Robert Lücking explained, "merging of the two families under the name Cladoniaceae is not possible without a conservation proposal because Cladoniaceae (Zenker, 1827) is antedated by Stereocaulaceae (Chevallier, 1826) by one year."[15]

Description

The thallus of Cladoniaceae lichens are fruticose or foliose, and are often dimorphic–consisting of two distinct forms. The primary thallus is ephemeral to persistent, crustose, foliose or squamulose, while the secondary thallus is typically vertical and holds the ascomata. The photobiont is chlorococcoid (green algae with a coccoid shape). The ascomata are in the form of an apothecium, and are biatorine, meaning they are of the lecideine type – light in colour and soft in consistency. They often have a reduced margin. The hamathecium (referring to all hyphae between the asci in the hymenium) consists of branched paraphyses, and is amyloid. The asci are somewhat fissitunicate, meaning they have two layers that separate during ascus dehiscence. The ascus structure consists of an apical dome and a tube (both of which are amyloid), which is cylindrical to clavate. Ascospores number eight per ascus, and they are usually non-septate, ellipsoid to more or less spherical in shape, hyaline, and non-amyloid. The conidiomata are pycnidia; the conidia are non-septate, usually filiform (thread-like), and hyaline.[16]

Genera
Pilophorus acicularis

When Edvard August Vainio published his three-volume monograph on the Cladoniaceae (Monographia Cladoniarum universalis, 1887, 1894, 1897), he included 134 species and subspecies. Now, after more than a century of discovery and research, including recent advances in understanding revealed by molecular phylogenetic studies, the Cladoniaceae encompass 17 genera and more than 550 species.

This is a list of the genera contained within the Cladoniaceae, based on the Catalogue of Life; this includes taxa formerly classified in the Squamarinaceae, but does not include the Stereocaulaceae. Following the genus name is the taxonomic authority, year of publication, and the number of species:

Myelorrhiza was transferred from the Cladoniaceae to the Ramalinaceae by Kistenich and colleagues in 2018.[18]

References
  1. Goebel, Karl Christian Traugott Friedemann; Kunze, G. (1827). Pharmaceutische Waarenkunde (in German). Eisenach: J.F. Bärecke. p. 124.
  2. Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, LKT; Dolatabadi, S; 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.
  3. Wei, Jiang-Chun; Ahti Teuvo (2002). "Cetradonia, a new genus in the new family Cetradoniaceae (Lecanorales, Ascomycota)". The Lichenologist. 34 (1): 19–31. doi:10.1006/lich.2001.0354.
  4. Hafellner, J. (1984). "Studien in Richtung einer natürlichen Gliederung der Sammelfamilien Lecanoracae und Lecideaceae". Beihefte zur Nova Hedwigia. 79: 241–371.
  5. Cannon, Paul F.; Kirk, Paul M. (2007). Fungal Families of the World. CAB International. p. 73. ISBN 978-0-85199-827-5.
  6. 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.
  7. Browne, Patrick (1756). Civil and natural history of Jamaica. London. p. 81.
  8. Wedin, Mats; Döring, Heidi; Ekman, Stefan (2000). "Molecular phylogeny of the lichen families Cladoniaceae, Sphaerophoraceae, and Stereocaulaceae (Lecanorales, Ascomycotina)". The Lichenologist. 32 (2): 171–187. doi:10.1006/lich.1999.0236.
  9. Stenroos, Soili; Myllys, Leena; Thell, Arne; Hyvönen, Jaakko (2002). "Phylogenetic hypotheses: Cladoniaceae, Stereocaulaceae, Baeomycetaceae, and Icmadophilaceae revisited". Mycological Progress. 1 (3): 267–282. doi:10.1007/s11557-006-0024-9.
  10. Arup, U.; Ekman, S.; Grube, M.; Mattsson, J.-E.; Wedin, M. (2007). "The sister group relation of Parmeliaceae (Lecanorales, Ascomycota)". Mycologia. 99 (1): 42–49. doi:10.1080/15572536.2007.11832599.
  11. Ekman, Stefan; Andersen, Heidi L.; Wedin, Mats; Buckley, Thomas (2008). "The limitations of ancestral state reconstruction and the evolution of the ascus in the Lecanorales (lichenized Ascomycota)". Systematic Biology. 57 (1): 141–156. doi:10.1080/10635150801910451.
  12. Miadlikowska, Jolanta; Kauff, Frank; Högnabba, Filip; Oliver, Jeffrey C.; Molnár, Katalin; Fraker, Emily; et al. (2014). "A multigene phylogenetic synthesis for the class Lecanoromycetes (Ascomycota): 1307 fungi representing 1139 infrageneric taxa, 317 genera and 66 families". Molecular Phylogenetics and Evolution. 79: 132–168. doi:10.1016/j.ympev.2014.04.003. PMC 4185256. PMID 24747130.
  13. Zhou, Qi-Ming; Wei, Jiang-Chun; Ahti, Teuvo; Stenroos, Soili; Högnabba, Filip (2006). "The systematic position of Gymnoderma and Cetradonia based on SSU rDNA sequences". Journal of the Hattori Botanical Laboratory. 100: 871–880.
  14. 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/boy060/5091569.
  15. Lücking, Robert (2019). "Stop the abuse of time! Strict temporal banding is not the future of rank-based classifications in fungi (including lichens) and other organisms". Critical Reviews in Plant Sciences. 38 (3): 199–253. doi:10.1080/07352689.2019.1650517. S2CID 202859785.
  16. Jaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert; Lumbsch, H. Thorsten (2016). Frey, Wolfgang (ed.). Syllabus of Plant Families: Adolf Engler's Syllabus der Pflanzenfamilien. 1/2 (13 ed.). Berlin Stuttgart: Gebr. Borntraeger Verlagsbuchhandlung, Borntraeger Science Publishers. p. 121. ISBN 978-3-443-01089-8. OCLC 429208213.
  17. Stenroos, Soili; Pino‐Bodas, Raquel; Ahti, Teuvo (2019). "Rexiella, a new name for Rexia S. Stenroos, Pino‐Bodas & Ahti (2018), non Rexia D. A. Casamatta, S. R. Gomez & J. R. Johansen (2006)". Cladistics. 35 (5): 603. doi:10.1111/cla.12401.
  18. Kistenich, Sonja; Timdal, Einar; Bendiksby, Mika; Ekman, Stefan (2018). "Molecular systematics and character evolution in the lichen family Ramalinaceae (Ascomycota: Lecanorales)". Taxon. 67 (5): 871–904. doi:10.12705/675.1. hdl:10852/67955.
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