Biology:Candelariaceae

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Candelariaceae is a family of lichen-forming fungi in the order Candelariales. [1] It contains seven genera and about 73 species of lichens that are typically characterised by their bright yellow colour. These lichens have diverse growth forms, including crusty, scaly, and small leaf-like structures, and are found worldwide, particularly in arid and mountainous regions of both hemispheres. The family includes well-known genera such as Candelaria, Candelariella, and Placomaronea, with their classification continuing to evolve based on molecular studies. Recent phylogenies find Candelina and Placomaronea to be well supported, but many relationships among Candelaria and Candelariella lineages remain unresolved.

Taxonomy

The Finnish lichenologist Rainar Hakulinen proposed Candelariaceae in 1954 when he removed the bright-yellow foliose genus Candelaria from Parmeliaceae and united it with the crustose Candelariella and the umbilicate Placomaronea.⁠[2] Candelaria had formerly been placed in the Parmeliaceae by Alexander Zahlbruckner in 1926,[3] while the crustose lichen genus Candelariella was classified among the crustose Lecanoraceae. The family remained morphologically delimited until Josef Poelt (1974) showed that cortex anatomy separated the placodioid Candelina—which has a densely pigmented outer cortex—from Placomaronea, where the cortex is [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] and capped by "peppered" pigment hoods.⁠[4] Early molecular work (Westberg et al. 2007) confirmed the monophyly of the family within Candelariales and recovered well-supported clades corresponding to Candelina and Placomaronea, but also revealed that Candelariella in its traditional sense is paraphyletic.⁠[5]

Subsequent phylogenies that sampled a wider array of internal transcribed spacer (ITS) sequences reiterated these patterns and showed that the ability to produce more than eight ascospores per ascus (polyspory) has evolved repeatedly inside the family.⁠[6] In an effort to impose monophyly, in 2020 Sergey Kondratyuk and colleagues erected three additional genera—Candelinella, Opeltiella and Protocandelariella—to house small but distinct ITS lineages, chiefly the grey-thalline Candelariella subdeflexa group.⁠[7] Because their multilocus backbone had weak support and relied on sparsely sampled loci, those proposals have been treated as provisional in some recent catalogues.[8]

The most comprehensive analysis to date, an ITS-only dataset of 303 accessions that included two newly described Placomaronea species, again retrieved robust clades for Candelina and Placomaronea and confirmed the basal position of the Protocandelariella lineage, but failed to give meaningful support to Candelinella or Opeltiella, and left relationships among the various Candelariella and Candelaria lineages in an unresolved polytomy.⁠[8] The authors therefore recommended a conservative taxonomy that recognises five well-supported genera—Candelaria, Candelariella sensu lato (in the broad sense), Candelina, Placomaronea and the anatomically distinctive Protocandelariella—while awaiting multilocus data that might justify further segregation. Under this pragmatic circumscription the family comprises roughly 80 accepted species distributed worldwide, especially in arid and montane regions of the Northern and Southern hemispheres.[8]

Description

Candelariella xanthostigma

The Candelariaceae exhibits a diverse range of growth forms, including crustose, [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]-[[Glossary of lichen terms#{{biology:{1}}}|subumbilicate]], and [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] thalli, typically characterized by their bright yellow colour. The [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] in these lichens is [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], contributing to their photosynthetic abilities. The reproductive structures, or ascomata, are predominantly [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] apothecia, although [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] apothecia are also found in some instances.[9]

The [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], which is part of the ascomata, is made up of unbranched or slightly branched, amyloid [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]. The asci, or spore-bearing structures, have an apical [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] with an amyloid outer wall. The tholus is weakly amyloid, except for the darker, strongly amyloid lower portion, and is typically [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] in shape. Each ascus contains 8 to 64 [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], which are either non-septate or indistinctly 1-septate. These spores are hyaline, non-amyloid, and vary in shape from ellipsoid to [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (lemon-shaped).[9]

Candelariaceae lichens also produce [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], which are [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]]-containing structures. The [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]], or asexual spores, are non-septate, hyaline, and range in shape from ellipsoid to [[Glossary of lichen terms#{{biology:{1}}}|{{Biology:{1}}}]] (rod shaped), sometimes exhibiting a curved form. The secondary chemistry of Candelariaceae lichens includes derivatives of pulmonic acid, a lichen product that contributes to their unique characteristics.[9]

Field and microscope identification

A rock-dwelling Placomarea lichen in Peru

Placodioid Placomaronea and Candelina are easily confused in the field; sections of the upper cortex resolve them: Placomaronea shows a thick paraplectenchymatous cortex with hooded apical cells "peppered" in pigment and a thin hyaline coating that flakes, whereas Candelina has a thinner cortex capped by a dense, cemented pigment layer and lacks any hyaline coating. Candelariella typically has a thinner, looser pigment layer and a mostly prosoplectenchymatous cortex; C. kansuensis is an outlier with a Placomaronea-like hooded cortex plus a very thick hyaline coating.[8]

Genera

Many authors provisionally recognise five core genera (Candelaria, Candelariella (in the loose sense), Candelina, Placomaronea, Protocandelariella) based on well-supported clades;[5] Candelinella and Opeltiella have been proposed[7] but remain weakly supported in ITS-only backbones and are treated conservatively by some workers pending more evidence with multiple genetic loci.[8]

References

  1. ↑ Wijayawardene, N.N.; Hyde, K.D.; Dai, D.Q.; Sánchez-García, M.; Goto, B.T.; Saxena, R.K. et al. (2022). "Outline of Fungi and fungus-like taxa – 2021". Mycosphere 13 (1): 53–453. doi:10.5943/mycosphere/13/1/2. https://www.researchgate.net/publication/358798332. 
  2. ↑ Hakulinen, R. (1954). "Die Flechtengattung Candelariella Müller Argoviensis" (in de). Annales Botanici Societatis Zoologicae Botanicae Fennicae "Vanamo" 27 (3): 11. 
  3. ↑ Zahlbruckner, A. (1926). "Lichenes. Spezieller Teil." (in de). Die natürlichen Pflanzenfamilien. 2 (8 ed.). pp. 61–270. 
  4. ↑ 4.0 4.1 Poelt, J. (1974). "Zur Kenntnis der Flechtenfamilie Candelariaceae" (in de). Phyton 16: 189–210. https://www.zobodat.at/pdf/PHY_16_1_4_0189-0210.pdf. 
  5. ↑ 5.0 5.1 Westberg, Martin; Arup, Ulf; Kärnefelt, Ingvar (2007). "Phylogenetic studies in the Candelariaceae (lichenized Ascomycota) based on nuclear ITS DNA sequence data". Mycological Research 111 (11): 1277–1284. doi:10.1016/j.mycres.2007.08.007. PMID 18006290. 
  6. ↑ Westberg, Martin; Morse, Caleb A.; Wedin, Mats (2011). "Two new species of Candelariella and a key to the Candelariales (lichenized Ascomycetes) in North America". The Bryologist 114 (2): 325–334. doi:10.1639/0007-2745-114.2.325. 
  7. ↑ 7.0 7.1 7.2 7.3 7.4 Kondratyuk, S.Y.; Lőkös, L.; Jeong, M.-H.; Oh, S.-O.; Kondratiuk, A.S.; Hur, J.-S. (2020). "Contributions to molecular phylogeny of lichen-forming fungi, 1. The family Candelariaceae". Acta Botanica Hungarica 62 (3–4): 293–307. doi:10.1556/034.62.2020.3-4.4. http://real.mtak.hu/143241/1/article-p293.pdf. 
  8. ↑ 8.0 8.1 8.2 8.3 8.4 Ramos, Daniel; Hollinger, Jason; Bungartz, Frank (2025). "Two new species of Placomaronea (Candelariaceae: lichenized Ascomycota) in Peru, with a revision of secondary chemistry and cortical anatomy of Placomaronea, Candelina and Candelariella". The Lichenologist 57 (2): 61–81. doi:10.1017/S0024282925000040. Bibcode: 2025ThLic..57...61R. 
  9. ↑ 9.0 9.1 9.2 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. 118. ISBN 978-3-443-01089-8. OCLC 429208213. 
  10. ↑ Müller, J. (1894). "Conspectus systematicus lichenum Novae Zelandiae" (in la). Bulletin de l'Herbier Boissier 2 (App. 1): 11, 47. 
  11. ↑ Räsänen, V. (1944). "Lichenes novi I". Annales Botanici Societatis Zoologicae Botanicae Fennicae "Vanamo" 20 (3): 1–34. 

Wikidata ☰ Q10441420 entry



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