Parmeliaceae

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Parmeliaceae
Parmelia.saxatilis 29,Plogoff,Kougon.Vizen 2020-01-29 03.jpg
Parmelia saxatilis
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Parmeliaceae
Zenker (1827)
Type genus
Parmelia
Ach. (1803)
Genera [1]

See text

The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species [2] in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: Xanthoparmelia (822 species), Usnea (355 species), Parmotrema (255 species), and Hypotrachyna (262 species). [3]

Contents

Nearly all members of the family have a symbiotic association with a green alga (most often Trebouxia spp., but Asterochloris spp. are known to associate with some species). [4] The majority of Parmeliaceae species have a foliose, fruticose, or subfruticose growth form. The morphological diversity and complexity exhibited by this group is enormous, and many specimens are exceedingly difficult to identify down to the species level.

The family has a cosmopolitan distribution, and is present in a wide range of habitats and climatic regions. [5] This includes everywhere from roadside pavement to alpine rocks, from tropical rainforest trees to subshrubs in the Arctic tundra. Members of the Parmeliaceae are found in most terrestrial environments. Several Parmeliaceae species have been assessed for the global IUCN Red List.

Taxonomy

Based on several molecular phylogenetic studies, the Parmeliaceae as currently circumscribed has been shown to be a monophyletic group. [6] This circumscription is inclusive of the previously described families Alectoriaceae, Anziaceae, Hypogymniaceae, and Usneaceae, which are all no longer recognised by most lichen systematists. However, despite the family being one of the most thoroughly studied groups of lichens, several relationships within the family still remain unclear. Phylogenetic analysis supports the existence of seven distinct clades in 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]

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] The diversification of various Parmelioideae lineages may have been a result of gaining innovations that provided adaptive advantages, such as melanin production in the genus Melanohalea . [11] Diversification of the Protoparmelioideae occurred during the Miocene. [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]

Generic classification

The classification history of Parmeliaceae reflects evolving approaches to fungal taxonomy over two centuries. When Erik Acharius first described Parmelia in 1803, it encompassed a broad range of foliose lichens with rounded apothecia. By the mid-1800s, researchers began segregating genera based on ascospore characteristics, leading to the recognition of distinct groups like Physcia and Xanthoria. The most dramatic period of generic splitting occurred in the 1970s and 1980s, when Mason Hale and others proposed numerous new genera based primarily on morphological features such as lobe shapes, rhizine types, and cortical chemistry. [14]

The advent of molecular phylogenetics techniques in the late 1990s provided new tools for evaluating which morphological and chemical characters were most reliable for defining genera. These studies led to significant refinements in generic concepts, supporting some previously proposed splits while showing others to be artificial. For example, molecular data revealed that the brown-fruited genus Neofuscelia needed to be merged into Xanthoparmelia , while confirming that groups like Parmotrema and Cetrelia represented distinct evolutionary lineages. [14]

Current understanding of generic relationships in Parmeliaceae emphasises the importance of reproductive characters over vegetative features. Characters of the ascomata (especially exciple anatomy and ascospore characteristics), conidial types, and cell wall polysaccharides have proven particularly valuable for defining natural groups. In contrast, some previously emphasised features such as thallus growth form and the presence of specific cortical substances have been shown to be more variable within lineages than previously thought. [14]

Modern molecular studies have established that approximately 75% of Parmeliaceae species belong to well-defined major clades, including groups like Xanthoparmelia, Parmotrema , and their close relatives. The relationships among the remaining genera continue to be refined through ongoing research. Rather than being defined by single diagnostic features, most genera are now recognised as monophyletic groups characterised by unique combinations of multiple morphological, chemical, and anatomical traits. [14]

Evolutionary history

Although fossil records of extant lichen species are scarce, the existence of some amber inclusions has allowed for a rough estimate of the divergence of the Parmeliaceae from its most recent common ancestor. An Anzia inclusion from 35–40 Myr-old Baltic amber and Parmelia from 15–45 Myr-old Dominican amber suggest a minimum age estimate for the Parmeliaceae of about 40 Myr. [15] [16] A fossil-calibrated phylogeny has estimated the Parmeliaceae to have diversified much earlier, around the Cretaceous–Paleogene boundary, 58–74 Myr ago. [17]

Characteristics

Thallus

Parmeliaceae thalli are most often foliose, fruticose or subfruticose, but can be umblicate, peltate, caespitose, crustose, or subcrustose. Two genera, Nesolechia and Raesaenenia , contain lichenicolous 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 are corticate on 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 example Usnea , Hypogymnia and Menegazzia ). The upper surface has a pored or non-pored epicortex. Medulla is solid, but often loosely woven. [18]

Apothecia

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 several Menegazzia species have two spores per ascus. [18]

Spores

Ascospores are simple, hyaline, and often small. Conidia generally 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. Pycnidia are 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. [18]

Chemistry

Members of the Parmeliaceae exhibit a diverse chemistry, with several types of lichenan ( Xanthoparmelia -type, Cetraria -type, intermediate-type), isolichenan and/or other polysaccharides being known from the cell walls of many species. [18] The wide diversity in the types of chemical compounds includes depsides, depsidones, aliphatic acids, triterpenes, anthraquinones, secalonic acids, pulvinic acid derivatives, and xanthones. The compounds usnic acid and atranorin, which are found exclusively in the Parmeliaceae, are of great importance in the systematics of the family, and the presence or absence of these chemicals have been used in several instances to help define genera. Parmelia and Usnea are the best chemically characterized genera, while the species Cetraria islandica and Evernia prunastri have attracted considerable research attention for their bioactive compounds. [19]

A study of three parmelioid lichens ( Bulbothrix setschwanensis , Hypotrachyna cirrhata , and Parmotrema reticulatum ) collected from high-altitude areas of Garhwal 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. [20]

Photobiont

The main photobiont genus that associates with Parmeliaceae species is the chlorophyte Trebouxia . In particular, the species Trebouxia jamesii appears to be especially prominent. Some Parmeliaceae genera are also known to associate with Asterochloris , [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

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

A genus FoveolariaChesnokov, Prokopiev, Konoreva & Davydov was proposed in 2023 to contain the species historically known as Cetraria nivalis and transferred to several genera (including Allocetraria, Flavocetraria, and Nephromopsis), [30] but this naming proposal was not valid, as the name has already been used for a plant genus; its current taxonomic status is unclear. [43]

Conservation

Parmeliaceae species that have been assessed for the global IUCN Red List include the following: Anzia centrifuga (vulnerable, 2014); [44] Sulcaria badia (endangered, 2019); [45] Lethariella togashii (vulnerable, 2017); [46] Hypotrachyna virginica (critically endangered, 2020); [47] Sulcaria isidiifera (critically endangered, 2017); [48] Sulcaria spiralifera (endangered, 2020); [49] and Xanthoparmelia beccae (vulnerable, 2017). [50]

Related Research Articles

<i>Parmelia</i> (fungus) Genus of lichens

Parmelia is a genus of medium to large foliose (leafy) lichens. It has a global distribution, extending from the Arctic to the Antarctic continent but concentrated in temperate regions. There are about 40 species in Parmelia. In recent decades, the once large genus Parmelia has been divided into a number of smaller genera according to thallus morphology and phylogenetic relatedness.

<i>Cetraria</i> Genus of lichens

Cetraria is a genus of fruticose lichens that associate with green algae as photobionts. Most species are found at high latitudes, occurring on sand or heath, and are characterised by their "strap-like" form with spiny lobe edges. The lobes can range from narrow and linear to broader and flattened, often forming loose or densely packed cushions. Their distinctive spiny margins serve both a defensive role and aid in vegetative reproduction through fragmentation. The genus was created by Erik Acharius in 1803 and belongs to the large family Parmeliaceae. While originally a species-rich genus, taxonomic revisions since the 1960s have split many species into new genera, though the exact circumscription remains debated among lichenologists.

<i>Ahtiana</i> Single-species genus of lichen

Ahtiana is a fungal genus in the family Parmeliaceae. A monotypic genus, it contains the single species Ahtiana sphaerosporella, the mountain candlewax lichen, found in western North America. The species was originally classified as Parmelia sphaerosporella by Johannes Müller Argoviensis in 1891, before Trevor Goward established the new genus Ahtiana in 1985, naming it after Finnish lichenologist Teuvo Ahti. This foliose lichen is characterised by its pale yellowish-green thallus, spherical spores, laminal apothecia, and the presence of usnic and caperatic acids. It primarily grows on the bark of whitebark pine in subalpine and montane regions, though it occasionally colonises other conifers outside its preferred host's range.

<i>Allocetraria</i> Genus of lichens

Allocetraria is a genus of lichenized fungi in the family Parmeliaceae. It consists of 12 species, with a center of distribution in China.

<i>Arctocetraria</i> Genus of lichens

Arctocetraria is a genus of foliose lichens in the family Parmeliaceae. It has three species.

<i>Cetrariella</i> Genus of lichen

Cetrariella is a genus of foliose lichens in the family Parmeliaceae. It contains three species.

<i>Kaernefeltia</i> Genus of fungi

Kaernefeltia is a genus of lichenized fungi in the family Parmeliaceae.

<i>Masonhalea</i> Genus of fungi

Masonhalea is a genus of two species of lichenized fungi in the family Parmeliaceae.

<i>Tuckermannopsis</i> Genus of lichens

Tuckermannopsis is a genus of foliose lichens in the family Parmeliaceae.

<i>Melanohalea</i> Genus of lichen

Melanohalea is a genus of foliose lichens in the family Parmeliaceae. It contains 30 mostly Northern Hemisphere species that grow on bark or on wood. The genus is characterised by the presence of pseudocyphellae, usually on warts or on the tips of isidia, a non-pored epicortex and a medulla containing depsidones or lacking secondary metabolites. Melanohalea was circumscribed in 2004 as a segregate of the morphologically similar genus Melanelia, which was created in 1978 for certain brown Parmelia species. The methods used to estimate the evolutionary history of Melanohalea suggest that its diversification primarily occurred during the Miocene and Pliocene epochs.

<i>Cetrelia</i> Genus of lichens in the family Parmeliaceae

Cetrelia is a genus of leafy lichens in the large family Parmeliaceae. They are commonly known as sea-storm lichens, alluding to the wavy appearance of their lobes. The name of the genus, circumscribed in 1968 by the husband and wife lichenologists William and Chicita Culberson, alludes to the former placement of these species in the genera Cetraria and Parmelia.

<i>Parmotrema</i> Genus of fungi

Parmotrema is a genus of lichen belonging to the family Parmeliaceae. It is a large genus, containing an estimated 300 species, with a centre of diversity in subtropical regions of South America and the Pacific Islands.

<i>Punctelia</i> Genus of foliose lichens

Punctelia is a genus of foliose lichens belonging to the large family Parmeliaceae. The genus, which contains about 50 species, was segregated from genus Parmelia in 1982. Characteristics that define Punctelia include the presence of hook-like to thread-like conidia, simple rhizines, and point-like pseudocyphellae. It is this last feature that is alluded to in the vernacular names speckled shield lichens or speckleback lichens.

<i>Xanthoparmelia</i> Genus of fungi

Xanthoparmelia is a genus of foliose lichens in the family Parmeliaceae. This genus of lichen is commonly found in the United States, South America, southern Africa, Europe, Australia, and New Zealand.

Coelopogon is a genus of lichen-forming fungi in the family Parmeliaceae. The genus contains two species found in southern South America and South Africa.

<i>Crespoa</i> Genus of fungi

Crespoa is a genus of five species of lichen in the family Parmeliaceae. Species in this genus are characterized by having an upper thallus surface that is wrinkled and reticulately ridged to coarsely foveolate.

Remototrachyna is a genus of foliose lichens in the large family Parmeliaceae. It was separated from the genus Hypotrachyna based on the structure of the excipulum and genetic differences.

Emodomelanelia is a lichen genus in the family Parmeliaceae. It is monotypic, containing the single foliose Himalayan species Emodomelanelia masonii.

<i>Notoparmelia</i> Genus of lichens

Notoparmelia is a genus of foliose lichens in the family Parmeliaceae. It includes 18 species that grow on bark and rocks, and are mostly distributed in the Southern Hemisphere. The genus was created in 2014 as a segregate of Parmelia.

References

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  15. ↑ Poinar, G.O.; Peterson, E.B.; Platt, J.L. (2000). "Fossil Parmelia in new World Amber". The Lichenologist. 32 (3): 263–269. Bibcode:2000ThLic..32..263P. doi:10.1006/lich.1999.0258.
  16. ↑ Rikkinen, Jouko; Poinar, George O. (2002). "Fossilised Anzia (Lecanorales, lichen-forming Ascomycota) from European Tertiary amber". Mycological Research. 106 (8): 984–990. doi:10.1017/S0953756202005907.
  17. ↑ 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 . PMC   3234259 . PMID   22174775.
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  26. 1 2 3 4 Hale, Mason E. (1974). "Bulbothrix, Parmelina, Relicina and Xanthoparmelia, four new genera in the Parmeliaceae". Phytologia. 28 (5): 479–490.
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  29. ↑ Brusse, F.A.; Kärnefelt, I. "The new southern hemisphere lichen genus Coelopogon (Lecanorales, Ascomycotina), with a new species from Southern Africa". Mycotaxon. 42: 35–41.
  30. 1 2 Chesnokov, Sergey V.; Davydov, Evgeny A.; Konoreva, Liudmila A.; Prokopiev, Ilya A.; Poryadina, Lena N.; Zheludeva, Elena V.; Shavarda, Alexey L. (2023). "The monotypic genus Flavocetraria and two new genera: Cladocetraria and Foveolaria, in the cetrarioid core". Plant Systematics and Evolution. 309 (4): e24. doi:10.1007/s00606-023-01862-2.
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  37. ↑ Nylander, W. (1896). Les Lichens des Environs de Paris [The Lichens of the Paris Region] (in French). Paris: Typographye Paul. Schmidt. pp. 39, 139.
  38. ↑ Tulasne, L.-R. (1852). "Mémoire pour servir à l'histoire organographique et physiologique des Lichens" [Memoir on the organographic and physiological history of Lichens]. Annales des Sciences Naturelles Botanique. Série 3 (in Latin). 17: 124.
  39. ↑ Petrak, F. (1931). "Fungi Adeani. Ein Beitrag zur Pilzflora Bayerns und der angrenzenden Länder" [Fungi Adeani: A contribution to the mycological flora of Bavaria and adjacent regions]. Kryptogamische Forschungen Kryptogamenkommission der Bayerischen Botanischen Gesellschaft zur Erforschung der heimischen Flora. II (in German) (2): 155–194.
  40. ↑ Zopf, Wilhelm (1903). "Vergleichende Untersuchungen über Flechten in Bezug auf ihre Stoffwechselprodukte" [Comparative studies on lichens regarding their metabolic products]. Beihefte zum Botanischen Centralblatt (in German). 14: 95–126.
  41. ↑ Krog, Hildur (1982). "Punctelia, a new lichen genus in the Parmeliaceae". Nordic Journal of Botany. 2 (3): 287–292. doi:10.1111/j.1756-1051.1982.tb01191.x.
  42. ↑ Bystrek, J. (1971). "Taxonomic studies on the genus Alectoria". Annales Universitatis Mariae Curie-Sklodowska. 26: 265–279.
  43. ↑ "Record Details: Foveolaria Chesnokov, Prokopiev, Konoreva & Davydov, in Chesnokov, Davydov, Konoreva, Prokopiev, Poryadina, Zheludeva & Shavarda, Pl. Syst. Evol. 309(no. 24): 14 (2023)". Index Fungorum . Retrieved 1 January 2025.
  44. ↑ Aptroot, A.; Perez-Ortega, S.; Scheidegger, C. (2014). "Anzia centrifuga". IUCN Red List of Threatened Species . 2014. Retrieved 19 April 2021.
  45. ↑ McMullin, T.; Allen, J.; Lendemer, J. (2019). "Sulcaria badia". IUCN Red List of Threatened Species . 2019. Retrieved 19 April 2021.
  46. ↑ Ohmura, Y.; Randlane, T.; Spribille, T. (2017). "Lethariella togashii". IUCN Red List of Threatened Species . 2017. Retrieved 19 April 2021.
  47. ↑ Allen, J.; Lendemer, J.; McMullin, T. (2020). "Hypotrachyna virginica". IUCN Red List of Threatened Species . 2020. Retrieved 19 April 2021.
  48. ↑ McMullin, T.; Allen, J.; Lendemer, J. (2019). "Sulcaria isidiifera". IUCN Red List of Threatened Species . 2019. Retrieved 19 April 2021.
  49. ↑ McMullin, T.; Stone, D.; Lendemer, J.; Allen, J. (2021). "Sulcaria spiralifera". IUCN Red List of Threatened Species . 2021. Retrieved 19 April 2021.
  50. ↑ Aptroot, A.; Perez-Ortega, S. (2018). "Xanthoparmelia beccae". IUCN Red List of Threatened Species . 2018. Retrieved 21 June 2021.
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