Biology:Cladoniaceae

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Short description: Family of lichens

Cladoniaceae
Cladonia subulata - Flickr - pellaea.jpg
Cladonia subuluata is the type species of the type genus of the family Cladoniaceae
Scientific classification e
Domain: Eukaryota
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]

The Cladoniaceae are a family of lichen-forming 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] Many Cladoniaceae lichens grow on soil, but others can use decaying wood, tree trunks, and, in a few instances, rocks as their substrate. They grow in places with high humidity, and cannot tolerate aridity.

Many Cladoniaceae species are characterised by a thallus that has two distinct forms: a scaly or crust-like primary thallus that, depending on the species, can be permanent or temporary, and a secondary fruticose thallus called a podetium or pseudopodetium. Cladoniaceae members form symbiotic associations with green algae from the class Trebouxiophyceae, usually the genus Asterochloris. Six Cladoniaceae species are included in the International Union for Conservation of Nature's Red List of Threatened Species.

Systematics

Historical taxonomy

Cladoniaceae was formally introduced to science in 1827 by German naturalist Jonathan Carl Zenker in a publication of Karl Goebel.[1] Zenker's initial concept of the family included genera that are now recognised as separate families, including Baeomycetaceae, Icmadophilaceae, and Stereocaulaceae.[6] William Nylander included 53 Cladonia species worldwide in his 1860 work Synopsis lichenum.[7] When Edvard August Vainio published his three-volume monograph on the Cladoniaceae (Monographia Cladoniarum universalis, 1887,[8] 1894,[9] and 1897[10]), he included 134 species and subspecies. In his circumscription of the family, the genera Pycnothelia, Cladia, and Cladina were included in the genus Cladonia.[11] In more recent history, significant progress in the knowledge of the taxonomy and biogeography of the family can be attributed to several decades of devoted research by Teuvo Ahti.[12]

Cladoniaceae is now one of the largest families of lichen-forming fungi,[13] with about 560 species distributed amongst 18 genera. The type genus is Cladonia, circumscribed by Irish physician and botanist Patrick Browne in 1756. He included eight 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."[14]

Etymology

As is standard practice in botanical nomenclature,[15] the name Cladoniaceae is based on the name of the type genus, Cladonia, with the ending -aceae indicating the rank of family. The genus name comprises the Greek word: κλάδος (klādos), meaning "branch", "bud", or "shoot"; and the Latin -ia, a suffix meaning "quality or state of a thing".[16]

Synonymy

Several phylogenetic studies have shown that Cladoniaceae is a member of the order Lecanorales, and is closely related to the family Stereocaulaceae.[17][18][19][20][21] The family Cetradoniaceae, which was created in 2002 to contain the endangered species Cetradonia linearis,[3] was folded into the Cladoniaceae in 2006.[22]

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 synonymised the families Squamarinaceae and Stereocaulaceae with the Cladoniaceae, resulting in a large increase in the number of genera and species.[23] The Squamarinaceae had already been included in the Cladoniaceae by previous authors.[13] Although this reorganisation 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."[24] In a 2021 treatment of the British and Irish Cladoniaceae, the authors also keep these families separate, noting "both families are monophyletic and easily distinguishable on both morphological and molecular terms".[25]

Description

In Cladonia coccifera, the conidia are embedded in a scarlet-coloured slime on the margins of the scyphi (cups).

The thallus of Cladoniaceae lichens are fruticose (bushy) or foliose (leafy), and are often dimorphic–consisting of two distinct forms; this particular type of growth is also known as cladoniiform.[26] The primary thallus is ephemeral to persistent, crustose, foliose or squamulose, while the secondary thallus is typically vertical and holds the ascomata.[27] The secondary thallus ranges in height from a few millimetres to more than 25 centimetres (10 in). Some species, however, form neither a primary thallus nor any fruticose structures.[28] As for vegetative propagules, isidia occur very rarely in this family, whereas soredia are common.[25]

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.[27] Their colour is typically dark brown (sometimes pale brown), red, ochraceous, or black.[28] The hamathecium (referring to all hyphae between the asci in the hymenium) consists of sparsely branched paraphyses, and is amyloid. The asci (spore-bearing cells) 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 (club-shaped). Ascospores number eight per ascus, and they are usually non-septate, ellipsoid to more or less spherical in shape, hyaline (translucent), and non-amyloid.[27] Except for a few genera that produce septate ascospores (Calathaspis, Pycnothelia and Pilophorus), the hymenium does not generally have characters that are useful in taxonomy.[28] The conidiomata are pycnidia; the conidia are non-septate, usually filiform (thread-like), and hyaline.[27]

Chemistry

In the Cladoniaceae, over 70 different secondary metabolites (lichen products), primarily polyphenols, have been identified. To observe the fluorescence of certain depsides and depsidones, ultraviolet light is directly applied to the specimens. Traditional colour spot tests with reagents like KOH can be ineffective at low concentrations. Consequently, thin-layer chromatography analysis becomes crucial for identifying the major lichen products in the specimens and verifying species identification.[29]

Photobionts

The symbiotic algal partner (photobiont) of most Cladoniaceae taxa are unicellular green algae, usually in the genus Asterochloris, but occasionally in the genus Chlorella;[29] both of these genera are in the class Trebouxiophyceae. Eleven species of Asterochloris have found to be associated with genus Cladonia; the algal genus – one of the most common lichen symbionts – occurs in the thalli of more than 20 lichen genera.[30] The most common photobionts in this genus that associate with Cladonia are A. glomerata, A. italiana, and A. mediterranea, with some lineages showing dominance in one or several climatic regions.[31] In contrast, Myrmecia was shown to be the main photobiont for the Mediterranean species Cladonia subturgida.[32] The algal genus Trebouxia, a very common lichen photobiont, has not been recorded associating with the Cladoniacae.[28] Some Pilophorus species associate with cyanobacterial symbionts (in addition to their association with green algae) in structures called cephalodia.[29] The cyanobacterial genera Nostoc and Stigonema are involved in these tripartite associations.[28] In a study of several Cladonia lichens collected from Southern Finland, the associated microbial community, which was found to be consistent amongst the different species, consisted largely of Alphaproteobacteria and Acidobacteriota.[33]

Development

Cladonia coniocraea (left) has subulate podetia, while the podetia of Cladonia fimbriata (middle) feature cup-shaped scyphi, commonly called "pixie cups", Cladonia cervicornis (right) has a more complex branching pattern.

The development of several Cladoniaceae genera have been studied in detail,[34][35][36][37][38][39] although the interpretation of results has sometimes been controversial.[28] Cladoniaceae species begin development with the formation of a prothallus – a fungal layer upon which an algae-containing thallus will develop. It comprises the hyphae from the germination of an ascospore. After the protothallus contacts the alga, lichenisation begins with the development of small squamules (scale-like thallus segments) that make up the primary thallus. Most Cladoniaceae have a mixed thallus, consisting of two parts: a base, parallel to the substrate, called the primary thallus and the other erect, the secondary thallus. The primary thallus is squamulose (scaly) or crustose (crustose-like). The secondary thallus consists of vertical structures that are shrubby and hollow, although they can be solid in rare cases. If these structures are made of generative tissue, they are called podetia; when they are made of vegetative tissue, they are called pseudopodetia. The morphology of these structures determines to a large part the taxonomy of the Cladoniaceae, which can range from simple to very complex branching patterns.[29] Cladonia minisaxicola, found in the mountains of Bahia (Brazil) is the only species in that large genus that is completely crustose and does not develop podetia.[40]

The tips of the podetia have a wide range of morphology in the Cladoniaceae. They can be straight, tapering from a wide base to a point (called subulate), or flaring on cup-shaped scyphi. The scyphi are sometimes closed, or have a central perforation, forming structures called funnels.[29] The podetia are slow-growing, with an annual growth rate generally ranging from 1 to 15 mm.[41]

Branching in the Cladoniaceae occurs on the podetium due to the growth dynamics of fungal meristem tissue at its apex. Two primary branching patterns exist: one where branches emerge from late divisions of a large meristem that alters its shape, and another where branches come from small meristems that split early but maintain their shape. These meristem growth dynamics are crucial for interpreting phylogeny in Cladoniaceae mycobionts, with the trend towards smaller, early-splitting meristems seen as an evolutionary advancement.[42] In cladoniiform lichens, especially within the Cladoniaceae, a shift in meristem growth from isotropous (having uniform properties in all directions) to anisotropous (having properties that differ depending on the direction in which they are measured) leads to pronounced lateral elongation of the apical meristem. This change offers developmental flexibility, transitioning from a symmetrical growth to a more varied, asymmetrical growth, hinting at evolutionary processes within the Cladoniaceae. Despite this variability, such morphogenetic activities appear to be highly conserved even among species that are presumably distantly related.[43]

Habitat and distribution

Cladoniaceae species have been recorded growing in many habitats and on a diversity of substrates, including soil, tree trunks, and rotten wood.[28] In a few cases, Cladoniaceae can grow on rocks, such as Cladonia salmonea which grows on the rock faces of vertical cliffs, or Cladonia pyxidata, which can grow on thin soil on rocks.[44] They are absent from very dry regions. The range of their habitats includes boreal forests, bogs, temperate forests, the tundra of the Arctic and Antarctic, man-made habitats (e.g. roadsides), tropical highlands, and the sandy tropical lowlands of the Amazon rainforest.[28]

In his 2000 monograph on the Cladoniaceae of the Neotropical realm, Ahti included 184 species in 4 genera, and showed that that South America is a hotspot of biodiversity for genus Clanodia.[12] Bioclimatic variables significantly influence the distribution of Cladoniaceae species richness in the Neotropics, particularly under conditions of low precipitation and temperature, and high climatic variability. Areas with stable climates and higher temperatures and precipitation tend to support greater species richness.[45] Twenty-six Cladoniaceae species (25 Cladonia and 1 Cladia) are known to occur in the Galápagos Islands. There, some species form mats on lava flows that have developed little soil.[46] A 2013 monograph of Northern European Cladoniaceae treated 100 species (95 Cladonia, 4 Pilophorus, and the monotypic genus Pycnothelia).[47] In the 2021 key to lichen species in Italy, 86 Cladoniaceae are included.[48] In Bulgaria, 55 species in two genera were reported in 2022.[49] In a study of the lichen biodiversity in Kazakhstan's Burabay National Park, the Cladoniaceae made up about 30 percent of the species diversity.[50]

A study on the distributional ecology of Cladina and Cladonia in western North America found that the Coast Mountains of British Columbia act as a key phytogeographic barrier. This results in distinct oceanic and continental taxa groupings on either side. The research also suggests that the southern boundaries of certain species may be determined more by historical rather than purely ecological factors, indicating possible range expansions.[51]

Conservation

Cladonia perforata has been an endangered species on the IUCN redlist since 2003.

Each of the six Cladoniaceae species that have been assessed for the global IUCN Red List face a variety of threats impacting their survival. Cetradonia linearis (vulnerable, 2015) is endangered by ecosystem changes in spruce–fir forests, specifically the balsam woolly adelgid's impact on Fraser fir, and changes in humidity regimes and cloud immersion. The species is also vulnerable to threats from logging, mining, and road building if its legal protection status is removed.[52] Cladonia appalachiensis (endangered, 2020) growing on high-elevation Anakeesta Knob rock, faces threats from visitor disruption and changes in cloud cover and humidity.[53] The main threats to Cladonia perforata (endangered, 2003) include habitat loss, hurricanes, and improper fire management, with a single natural event potentially causing substantial subpopulation reduction.[54] Cladonia submitis (endangered, 2020) is primarily threatened by habitat loss and degradation due to land development, particularly around metropolitan areas. Climate change also poses significant risks through altered fire regimes and sea level rise, affecting its pine barren and sand dune habitats.[55] Pilophorus fibula (endangered, 2020) is threatened by habitat loss, alteration of hydrological regimes, recreational damage, and declining water quality.[56] Lastly, Gymnoderme insulare (endangered, 2014), primarily found in old-growth forests in Japan and Taiwan, faces threats from natural hazards like typhoons and is affected by the decline of its tree hosts, Cryptomeria japonica and Chamaecyparis obtusa.[57]

On the red list of China's macrofungi, Cladonia delavayi (vulnerable), Cladonia pseudoevansii (critically endangered), Gymnoderma coccocarpum (endangered), and Gymnoderma insulare (endangered) are the representatives of the Cladoniaceae.[58]

Genera

After more than a century of discovery and research, including recent advances in understanding revealed by molecular phylogenetics 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:

Pilophorus acicularis
Pycnothelia papillaria

Myelorrhiza was transferred from the Cladoniaceae to the Ramalinaceae by Kistenich and colleagues in 2018.[72] Neophyllis, originally classified in the Cladoniaceae, was transferred to Sphaerophoraceae in 1999.[73]

References

  1. ↑ 1.0 1.1 Goebel, Karl Christian Traugott Friedemann; Kunze, G. (1827) (in de). Pharmaceutische Waarenkunde. Eisenach: J.F. Bärecke. p. 124. 
  2. ↑ 2.0 2.1 2.2 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. ↑ 3.0 3.1 3.2 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. https://books.google.com/books?id=Lw1VLSH1xnAC&pg=PA73. 
  6. ↑ Ahti 2000, p. 3.
  7. ↑ Nylander, William (1860) (in la). Synopsis methodica lichenum. Paris: Imprimerie de L. Martinet. https://bibdigital.rjb.csic.es/records/item/12889-synopsis-methodica-lichenum-tomus-primus. 
  8. ↑ Wainio, Edvard August (1887). Monographia Cladoniarum universalis: I. Universal Monograph of Cladonias: I. Acta Societatis pro Fauna et Flora Fennica. 4. pp. 1–509. https://www.biodiversitylibrary.org/page/5587642. 
  9. ↑ Wainio, Edvard August (1894). Monographia Cladoniarum universalis: II. Acta Societatis pro Fauna et Flora Fennica. 10. pp. 1–499. https://www.biodiversitylibrary.org/page/5583213. 
  10. ↑ Wainio, Edvard August (1897). Monographia Cladoniarum universalis: III. Acta Societatis pro Fauna et Flora Fennica. 14. pp. 1–268. https://www.biodiversitylibrary.org/page/5634825. 
  11. ↑ Ahti 2000, p. 4.
  12. ↑ 12.0 12.1 Coppins, B. (2002). "Cladoniaceae. Teuvo Ahti. Flora Neotropica Monograph No. 78. New York: New York Botanical Garden Press. 2000. 362pp., 215 figures, 6 tables. ISBN 0 89327 431 3.". Edinburgh Journal of Botany 59 (3): 459–466. doi:10.1017/S0960428602230292. 
  13. ↑ 13.0 13.1 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. 
  14. ↑ 14.0 14.1 Browne, Patrick (1756). Civil and natural history of Jamaica. London: T. Osborne and J. Shipton. p. 81. https://www.biodiversitylibrary.org/page/11267602. 
  15. ↑ Hawksworth, David L. (1974). Mycologist's Handbook. Kew: Commonwealth Mycological Institute. p. 39. ISBN 978-0-85198-300-4. 
  16. ↑ Ulloa, Miguel; Aguirre-Acosta, Elvira (2020). Illustrated Generic Names of Fungi. St. Paul, Minnesota: APS press. p. 79. ISBN 978-0-89054-618-5. 
  17. ↑ 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. 
  18. ↑ 18.0 18.1 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. Bibcode: 2002MycPr...1..267S. 
  19. ↑ 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. PMID 17663122. 
  20. ↑ 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. PMID 18300027. 
  21. ↑ 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. PMID 24747130. 
  22. ↑ 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. 
  23. ↑ 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. 
  24. ↑ 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. Bibcode: 2019CRvPS..38..199L. 
  25. ↑ 25.0 25.1 Pino-Bodas, Rachel; Sanderson, Neil; Cannon, Paul; Aptroot, André; Coppins, Brian; Orange, Alan; Simkin, Janet (2021). "Lecanorales: Cladoniaceae [revision 1 including the genera Cladonia, Pilophorus and Pycnothelia"]. Revisions of British and Irish Lichens. 26 (3rd ed.). UK: The British Lichen Society. pp. 16. https://britishlichensociety.org.uk/sites/default/files/Cladoniaceae%20rev%201.pdf. 
  26. ↑ Ahti, T. (1982). "The morphological interpretation of cladoniiform thalli in lichens". The Lichenologist 14 (2): 105–113. doi:10.1017/s0024282982000255. 
  27. ↑ 27.0 27.1 27.2 27.3 Jaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert; Lumbsch, H. Thorsten (2016). 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. 
  28. ↑ 28.0 28.1 28.2 28.3 28.4 28.5 28.6 28.7 28.8 Stenroos, Soili; Pino-Bodas, Raquel; Hyvönen, Jaakko; Lumbsch, Helge Thorsten; Ahti, Teuvo (2018). "Phylogeny of the family Cladoniaceae (Lecanoromycetes, Ascomycota) based on sequences of multiple loci". Cladistics 35 (4): 351–384. doi:10.1111/cla.12363. PMID 34633698. 
  29. ↑ 29.0 29.1 29.2 29.3 29.4 Burgaz, Ana Rosa; Ahti, Teuvo; Pino-Bodas, Raquel (2020). Mediterranean Cladoniaceae. Madrid: Spanish Lichen Society (SEL). ISBN 978-84-09-21610-9. 
  30. ↑ Škaloud, Pavel; Steinová, Jana; Řídká, Tereza; Vančurová, Lucie; Peksa, Ondřej; De Clerck, O. (2015). "Assembling the challenging puzzle of algal biodiversity: species delimitation within the genus Asterochloris (Trebouxiophyceae, Chlorophyta)". Journal of Phycology 51 (3): 507–527. doi:10.1111/jpy.12295. PMID 26986666. Bibcode: 2015JPcgy..51..507S. 
  31. ↑ Pino-Bodas, Raquel; Stenroos, Soili (2020). "Global biodiversity patterns of the photobionts associated with the genus Cladonia (Lecanorales, Ascomycota)". Microbial Ecology 82 (1): 173–187. doi:10.1007/s00248-020-01633-3. PMID 33150498. 
  32. ↑ Pino-Bodas, Raquel; Blázquez, Miguel; de los Ríos, Asunción; Pérez-Ortega, Sergio (2023). "Myrmecia, not Asterochloris, is the main photobiont of Cladonia subturgida (Cladoniaceae, Lecanoromycetes)". Journal of Fungi 9 (12): e1160. doi:10.3390/jof9121160. PMID 38132761. 
  33. ↑ Shishido, Tânia Keiko; Wahlsten, Matti; Laine, Pia; Rikkinen, Jouko; Lundell, Taina; Auvinen, Petri (2021). "Microbial communities of Cladonia lichens and their biosynthetic gene clusters potentially encoding natural products". Microorganisms 9 (7): e1347. doi:10.3390/microorganisms9071347. PMID 34206222. 
  34. ↑ Jahns, H.M. (1970) (in de). Untersuchungen zur Entwicklungsgeschichte der Cladoniaceen unter besonderer Berücksichtigung des Podetien- Problems. Nova Hedwigia. 20. pp. 1–177. 
  35. ↑ Jahns, H.M. (1970). "Induktion der Apothecienbildung bei Cladia aggregata (Sw.) Nyl." (in de). Berichte der Deutschen Botanischen Gesellschaft 83: 33–40. doi:10.1111/j.1438-8677.1970.tb02301.x. 
  36. ↑ Jahns, H.M.; Beltman, H.A. (2007). "Variations in the ontogeny of fruiting bodies in the genus Cladonia and their taxonomic and phylogenetic significance". The Lichenologist 5 (5–6): 349–367. doi:10.1017/S0024282973000447. 
  37. ↑ Hammer, Samuel (1992). "Development in Cladonia ochrochlora". Mycologia 85 (1): 84–92. doi:10.1080/00275514.1993.12026250. 
  38. ↑ Hammer, Samuel (2018). "Primary tissue and the structure of the podetium in Cladonia". Mycologia 87 (1): 46–53. doi:10.1080/00275514.1995.12026501. 
  39. ↑ Grube, Martin; Hawksworth, David L. (2007). "Trouble with lichen: the re-evaluation and re-interpretation of thallus form and fruit body types in the molecular era". Mycological Research 111 (9): 1116–1132. doi:10.1016/j.mycres.2007.04.008. PMID 17698333. 
  40. ↑ Aptroot, André; da Silva Cáceres, Marcela Eugenia (2018). "New lichen species from Chapada Diamantina, Bahia, Brazil". The Bryologist 121 (1): 67–79. doi:10.1639/0007-2745-121.1.067. 
  41. ↑ Ahti 2000, p. 12.
  42. ↑ Hammer, Samuel (2000). "Meristem growth dynamics and branching patterns in the Cladoniaceae". American Journal of Botany 87 (1): 33–47. doi:10.2307/2656683. PMID 10636828. 
  43. ↑ Hammer, Samuel (2001). "Lateral growth patterns in the Cladoniaceae". American Journal of Botany 88 (5): 788–796. doi:10.2307/2657031. PMID 11353704. 
  44. ↑ Ahti 2000, p. 23.
  45. ↑ Soto-Medina, Edier Alberto (2013). "Patrones de Riqueza de Especies de la Familia Cladoniaceae en el Neotrópico Patterns of Species Richness of Family Cladoniaceae in the Neotropics" (in es). Cryptogamie, Mycologie 34 (2): 137–148. doi:10.7872/crym.v34.iss2.2013.137. 
  46. ↑ Yánez, Alba; Ahti, Teuvo; Bungartz, Frank (2013). "The Family Cladoniaceae (Lecanorales) in the Galapagos Islands". Phytotaxa 129 (1): 1–33. doi:10.11646/phytotaxa.129.1.1. 
  47. ↑ Ahti, Teuvo; Stenroos, Soili; Moberg, Roland (2013). Nordic Lichen Flora. Cladoniaceae. 5. Uppsala: Museum of Evolution, Uppsala University. ISBN 978-91-85221-29-5. 
  48. ↑ Gheza, Gabriele; Nimis, Per Luigi (22 July 2021). "Keys to the Lichens of Italy. 61) Cladoniaceae (Cladonia, Pilophorus, and Pycnothelia)". http://italic.units.it/flora/index.php?procedure=ext_key_home&key_id=3975. 
  49. ↑ Burgaz, Ana Rosa; Márquez, Rodrigo; PinoBodas, Raquel (2022). "The Cladoniaceae (Lecanorales, Ascomycota) from Bulgaria". Herzogia 35 (2): 510–540. doi:10.13158/heia.35.2.2022.510. 
  50. ↑ Bukabayeva, Zhanilxan; Abiev, Sardarbek; Korolev, Alexander (2022). Species diversity of lichens on the territory of the Burabay National Park (Republic of Kazakhstan) (Report). pp. 1–13. doi:10.21203/rs.3.rs-2195522/v1. 
  51. ↑ Goward, Trevor; Ahti, Teuvo (1997). "Notes on the distributional ecology of the Cladoniaceae (lichenized ascomycetes) in temperate and boreal western North America". Journal of the Hattori Botanical Laboratory 82: 143–155. https://www.jstage.jst.go.jp/article/jhbl/82/0/82_143/_pdf. 
  52. ↑ Allen, J.; Lendemer, J.; McMullin, T. (11 August 2015). "Rock Gnome Lichen. Cetradonia linearis". The IUCN Red List of Threatened Species 2020. https://www.iucnredlist.org/species/70386009/70386019. 
  53. ↑ Lendemer, J.; Allen, J.; McMullin, T. (6 October 2019). "Cladonia appalachiensis". The IUCN Red List of Threatened Species 2020. https://www.iucnredlist.org/species/80702853/80702858. 
  54. ↑ Yahr, R. (30 April 2003). "Cladonia perforata". The IUCN Red List of Threatened Species 2003. https://www.iucnredlist.org/species/43994/10838980. 
  55. ↑ Hoffman, J.; Ohmura, Y.; Lendemer, J. (30 August 2020). "Mid-Atlantic Comb-over. Cladonia submitis". The IUCN Red List of Threatened Species 2021. https://www.iucnredlist.org/species/43994/10838980. 
  56. ↑ Lendemer, J.; Allen, J.; McMullin, T. (8 October 2019). "Appalachian Matchsticks. Pilophorus fibula". The IUCN Red List of Threatened Species 2020. https://www.iucnredlist.org/species/80703047/80703054. 
  57. ↑ Ohmura, Y.; Nadyeina, O.; Scheidegger, C. (14 August 2014). "Gymnoderme insulare". The IUCN Red List of Threatened Species 2014. https://www.iucnredlist.org/species/58520980/58520984. 
  58. ↑ Yijian, Yao; Jiangchun, Wei; Wenying, Zhuang; Tiezheng, Wei; Yi, Li; Xinli, Wei et al. (2020). "Threatened species list of China's macrofungi". Biodiversity Science 28 (1): 20–25. doi:10.17520/biods.2019174. 
  59. ↑ Lamb, I.M.; Weber, W.A. (1974). "Calathaspis, a new genus of lichens (Cladoniaceae) from New Guinea". Occasional Papers of the Farlow Herbarium of Cryptogamic Botany 4: 1–12. 
  60. ↑ Nylander, William (1870) (in fr). Recognitio Monographica Ramalinarum. Caen: Impr. de P. Le Blanc-Hardel. p. 69. 
  61. ↑ Müller, J. (1889). "Lichenologische Beiträge XXXII" (in de). Flora (Regensburg) 72 (5): 505–508. 
  62. ↑ Galloway, D.J.; James, P.W. (1987). "Metus, a new austral lichen genus and notes on an Australian species of Pycnothelia". Notes from the Royal Botanical Garden Edinburgh 44 (3): 561–579. 
  63. ↑ Jørgensen, P.M.; Jahns, H.M. (1987). "Muhria, a remarkable new lichen genus from Scandinavia". Notes from the Royal Botanical Garden Edinburgh 44: 581–599. 
  64. ↑ Hammer, Samuel (2003). "Notocladonia, a new genus in the Cladoniaceae". The Bryologist 106 (1): 162–167. doi:10.1639/0007-2745(2003)106[0162:NANGIT2.0.CO;2]. 
  65. ↑ Liu, Jian Kui; Hyde, Kevin D.; Jones, E. B. Gareth; Ariyawansa, Hiran A.; Bhat, Darbhe J.; Boonmee, Saranyaphat et al. (2015). "Fungal diversity notes 1–110: taxonomic and phylogenetic contributions to fungal species". Fungal Diversity 72 (1): 1–197. doi:10.1007/s13225-015-0324-y. 
  66. ↑ Fries, Th. M. (1857). "Ad amimadversiones Cl. W. Nylanderi" (in la). Botaniska Notiser 1857: 167–174. https://www.biodiversitylibrary.org/page/62267180. 
  67. ↑ Dufour, J.L.M. (1821). "Révision des genres Cladonia, Scyphophorus, Helopodium et Baeomyces de la flore française" (in fr). Annales Générales des Sciences Physiques 8: 41–73. https://books.google.com/books?id=eekrAQAAMAAJ&pg=PA41. 
  68. ↑ 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. 
  69. ↑ Vainio, E.A. (1890). "Étude sur la classification naturelle et la morphologie des Lichens du Brésil. Pars prima" (in la). Acta Societatis Pro Fauna et Flora Fennica 7 (1): 7. 
  70. ↑ Hammer, Samuel (2001). "Additions to the lichen family Cladoniaceae in Australia". The Bryologist 104 (4): 560–575. doi:10.1639/0007-2745(2001)104[0560:ATTLFC2.0.CO;2]. 
  71. ↑ Montagne, C.; Berkeley, M.J. (1846). "On Thysanothecium, a new genus of lichens". Hooker's Journal of Botany 5: 257–258. https://www.biodiversitylibrary.org/page/775626. 
  72. ↑ 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. 
  73. ↑ Wedin, Mats; Döring, Heidi (1999). "The phylogenetic relationship of the Sphaerophoraceae, Austropeltum and Neophyllis (lichenized Ascomycota) inferred by SSU rDNA sequences". Mycological Research 103 (9): 1131–1137. doi:10.1017/S0953756298008223. 

Cited literature

  • Ahti, Teuvo (2000). Cladoniaceae. Flora Neotropic Monograph. 78. Bronx, New York: New York Botanical Garden Press. ISBN 978-0-89327-431-3. 

Wikidata ☰ Q1363839 entry



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