Punctelia

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Punctelia
Punctelia borreri (26167464298).jpg
Punctelia borreri
Scientific classification e
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Parmeliaceae
Genus: Punctelia
Krog (1982)
Type species

(Sm.) Krog (1982)

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 (asexual spores), simple rhizines (root-like structures that attach the lichen thallus to its substrate), and point-like pseudocyphellae (tiny pores on the thallus surface that facilitate gas exchange). It is this last feature that is alluded to in the vernacular names speckled shield lichens or speckleback lichens.

Punctelia lichens grow on bark, wood, and rocks. The genus is cosmopolitan, occurring on all continents but Antarctica. Species are found in temperate to subtropical locations. Punctelia has centres of distribution in the Neotropics and Africa; about half of the known species occur in South America. The photobiont partners of Punctelia are green algae in the genus Trebouxia. Some Punctelia species have been proposed for use as bioindicators of air pollution.

Systematics[]

The genus was circumscribed by Norwegian lichenologist Hildur Krog in 1982. The genus originally contained 22 species segregated from Parmelia based on differences in the development of the pseudocyphellae, secondary chemistry, and phytogeography.[1] Before Krog's publication, the Parmelia species with point-like pseudocyphellae were referred to as the Parmelia borreri group, and categorized in Parmelia subgenus Parmelia, section Parmelia, subsection Simplices.[2] Krog divided Punctelia into two subgenera: Punctelia subgenus Punctelia, characterized by hook-shaped (unciform) spermatia and atranorin as a major cortical substance, and Punctelia subgenus Flavopunctelia characterized by bifusiform spermatia and usnic acid as a major cortical substance.[1] Based on differences in spermatia shape as well as additional chemical characters, Flavopunctelia was later recognized by Mason Hale as a separate genus consisting of four species.[3] A preliminary molecular phylogenetic analysis of these two genera, published in 2005, corroborated their segregation from Parmelia, and confirmed the genus delimitations.[4]

In North America, member of the genus are commonly known as "speckled shield lichens" or "speckleback lichens".[5] The genus name also refers to the pseudocyphellae: it is derived from the Latin punctum, meaning "small spot" or "dot".[6]

Phylogenetics[]

 
 
 
 
 
 
A
 

P. guanchica

 
 

P. rudecta

 
 
 

P. toxodes

 
 
B
 
 
 

P. missouriensis

 
 

P. aff rudecta

 
 
 

P. ruderata

 
 
 
 

P. perreticulata

 
 

P. subrudecta

 
 
 
 
C
 
 

 
 

P. caseana

 
 
 

P. pseudocoralloidea

 
 
 
D
 

P. bolliana

 
 

P. appalachensis

 
 
 
 

P. hypoleucites

 
 
E
 
 
 
 

 
 

P. subpraesignis

 
 
 

P. reddenda

 
 
 
 

 
 
 
Phylogeny of Punctelia, using sequences from about a third of its species. Letter labels on the lines correspond to the five monophyletic clades recognized in Punctelia.[7]

Punctelia is a member of the large lichen family Parmeliaceae.[8] In 2017, Divakar and colleagues used a then-recently developed "temporal phylogenetic" approach to identify temporal bands for specific taxonomic ranks in the family Parmeliaceae, suggesting that groups of species that diverged within the time window of 29.45–32.55 million years ago represent genera. They proposed to synonymize the lichenicolous genus Nesolechia with Punctelia (its lichen-forming sister group), because the former group of species originated relatively recently and fell under the timeframe threshold for genus level.[9] This synonymy was not accepted in a later analysis.[10]

Molecular phylogenetic analysis has been used to more accurate delimit Punctelia species and identify previously undetected cryptic species, a research trend that has become common in the family Parmeliaceae, where dozens of cryptic species have been identified.[11] In the case of Punctelia, P. rudecta was thought until recently to have an expansive global distribution, with a range including North and South America, Africa, and Asia. Phylogenetic analysis revealed a species complex that has subsequently been split into four distinct cryptic lineages with more restricted distributional ranges; the range of P. rudecta has been reduced to North America. This research also revealed that there are five major clades in Punctelia, and each clade has characteristic patterns in medullary chemistry. Clades A, B, and C have species with lecanoric acid, clade D has species with gyrophoric acid as the main compound, while clade E has two species that have fatty acids as the main secondary chemical.[7]

Description[]

Herbarium specimen of Punctelia rudecta showing rounded pseudocyphellae on a lobe.

Punctelia lichens are medium-sized, foliose (leafy), and grey to greyish-green,[5] although collected specimens gradually lose their colour tone.[7] The size range for most typical specimens is 4 to 20 cm (1.6 to 7.9 in) in diameter.[6] The lobes that comprise the thallus are typically 3–10 mm across. The medulla is white, while the lower surface ranges from pale to black. Simple, unbranched rhizines are present that extend to the edge of the lobe;[5] they are usually more or less the same colour as the thallus underside, although individuals with light rhizines on a dark background are not unusual.[1]

A major characteristic of Punctelia is the presence of point-like (punctate) pseudocyphellae on the surface of the thallus. These are tiny pores that facilitate gas exchange. In the genus Parmelia, pseudocyphellae are straight and without a distinct form, and typically situated on the thallus surface (laminal) and/or on the margins (marginal). In comparison, Punctelia pseudocyphellae are rounded (orbicular) and laminal, although in some species the cortex gets pushed around the edges of the thallus, giving them a marginal appearance. Electron microscopy of Parmelia pseudocyphellae reveals a perforated polysaccharide layer; this layer is absent in Punctelia.[1] Pseudocyphellae are termed conspicuous when then can be viewed with the naked eye, inconspicuous when a hand lens or microscope is needed to see them, and subtle for intermediate states where they can be seen only with concerted effort.[12]

The apothecia (sexual reproductive structures) are lecanorine, with brown discs. Ascospores are colourless, ellipsoid, and number eight per ascus;[5] they range in size from 10–27 to 6–18 μm.[6] The unciform (hook-like) shape of the conidia is another major characteristic of genus Punctelia. They are short rods measuring 4–7 μm long with one end curved. Although not all Punctelia species have unciform conidia, this conidial shape only occurs in Punctelia.[1] Some species have filiform (threadlike) conidia that are in the size range 7–12 μm long by 0.8–1 μm wide.[6] The size and shape of the conidia is an important character in some instances; for example, P. graminicola and P.  hypoleucites are morphologically indistinguishable from each other, and they can only be reliably identified by differences in their conidia.[13] Cell walls of Punctelia lichens contain the alpha glucan polysaccharide isolichenan.[6]

Other Parmeliaceae genera that are superficially similar to Punctelia and have pseudocyphellae are Flavopunctelia and Cetrelia. Flavopunctelia species tend to be yellower than Punctelia due to the presence of usnic acid in the cortex. Cetrelia is usually larger with lobes measuring 1–2 cm (0.4–0.8 in), a dark lower thallus surface, and few rhizines.[5]

Secondary chemicals found in the genus include atranorin in the cortex, and gyrophoric acid in the medulla.[1] Lecanoric acid has been detected as a minor component in Punctelia jujensis and P. subrudecta.[14]

Photobiont[]

Most lichen genera associate with a photobiont partner from one algal genus.[15] Punctelia is no exception to this general rule; it associates with species from the most common photobiont genus, Trebouxia. In a study of photobiont partner selectivity, Punctelia subrudecta specimens collected from central Europe were shown to have a moderate selectivity, associating with three species of Trebouxia: , , and (the latter most frequently).[15] The photobiont partner for P. rudecta is .[16]

Habitat and distribution[]

Punctelia lichens are found on bark, wood, and rocks.[5] In the biodiverse cerrado forests of Brazil, they are more or less limited to well-lit microhabitats without direct sunlight.[17] They have a temperate to subtropical distribution with centers of distribution in the Neotropics and Africa.[7] The genus has a cosmopolitan distribution,[6] occur on all continents, with the exception of Antarctica.[7] Only a few species are known to occur in Asia and Australia; in contrast, most Parmelia species occur in these regions.[1] Five species are known from Australia, including two cosmopolitan species and three Australasian species.[6] Sixteen Punctelia species occur in the continental United States and Canada.[18] About half of the known Punctelia species are found in Brazil.[17] Several European countries have revised the genus in recent decades, including Norway (2000),[19] Switzerland (2003),[20] Denmark (2007),[21] and Lithuania (2010).[22] Six species occur in Europe.[23]

Species[]

Punctelia appalachensis
Punctelia guanchica
Punctelia jeckeri
Punctelia rudecta

A recent (2020) estimate places 48 species in the genus Punctelia.[8] As of April 2021, Species Fungorum accepts 30 species of Punctelia.[24]

The species Punctelia pallescens, described by Syo Kurokawa in 1999 as a new species from western Australia,[30] is considered synonymous with P. subalbicans.[46] Parmelia helenae, described by in 1937[47] and transferred to Punctelia in 1998,[48] was considered by some lichenologists to be a questionable taxon because, according to Teuvo Ahti, "the type material is insufficient to resolve its taxonomic relationship with Punctelia perrituculata ... and P. subrudecta ... on the basis of conidial characters".[48] It is now placed in synonymy with P. subrudecta.[1][23] Punctelia semansiana (W.L.Culb. & C.F.Culb.) Krog is the same species as Punctelia graminicola.[33]

Human uses[]

Biomonitoring[]

Some members of Punctelia have been shown to be somewhat sensitive to air pollution. A study conducted in Spain observed Punctelia borreri and P. subrudecta reappearing in areas with a decline in SO2 pollution.[49] Two Punctelia species have been recommended for use as element bioindicators in air pollution monitoring studies in the eastern United States. Punctelia rudecta is suggested for use in cooler forested uplands, and P. missouriensis for use in isolated woodlands or urban areas.[50] Because of the widespread occurrence of P. hypoleucites in both urban and industrial sites in and around Tandil, Argentina, it has been proposed as a potential biomonitor of air pollution in that city.[51]

Dyeing[]

Punctelia rudecta can be used to create a dye by a color-extraction with ammonia as a solvent. A pink color is obtained using this method.[52]

References[]

  1. ^ Jump up to: a b c d e f g h i j k l m n o p q r s t u v w x y 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.
  2. ^ Hale, Mason E.; Kurokawa, Syo (1964). "Studies on Parmelia subgenus Parmelia" (PDF). Contributions from the United States National Herbarium: 121–192. open access
  3. ^ Hale Jr, Mason E. (1983). "Flavopunctelia, a new genus in the Parmeliaceae (Ascomycotina)". Mycotaxon. 20 (2): 681–682.
  4. ^ Thell, Arne; Herber, B.; Aptroot, A.; M.T., Adler; T., Feuerer; Kärnefelt, E.I. (2005). "A preliminary phylogeographic study of Flavopunctelia and Punctelia inferred from rDNA ITS-sequences" (PDF). Folia Cryptogamica Estonica. 41: 115–122.
  5. ^ Jump up to: a b c d e f Brodo, Irwin M.; Sharnoff, Sylvia Duran; Sharnoff, Stephen (2001). Lichens of North America. Yale University Press. pp. 605–606. ISBN 978-0300082494.
  6. ^ Jump up to: a b c d e f g Elix, John A. (1994). "Punctelia". Lichens—Lecanorales 2, Parmeliaceae (PDF). Flora of Australia. 55. Canberra: Australian Biological Resources Study/CSIRO Publishing. ISBN 978-0-643-05676-3.
  7. ^ Jump up to: a b c d e f Alors, David; Lumbsch, H. Thorsten; Divakar, Pradeep K; Leavitt, Steven D.; Crespo, Ana (2016). "An integrative approach for understanding diversity in the Punctelia rudecta species complex (Parmeliaceae, Ascomycota)". PLOS ONE. 11 (2): 1–17. Bibcode:2016PLoSO..1146537A. doi:10.1371/journal.pone.0146537. PMC 4749632. PMID 26863231. open access
  8. ^ Jump up to: a b Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, 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.
  9. ^ Divakar, Pradeep K.; Crespo, Ana; Kraichak, Ekaphan; Leavitt, Steven D.; Singh, Garima; Schmitt, Imke; Lumbsch, H. Thorsten (2017). "Using a temporal phylogenetic method to harmonize family- and genus-level classification in the largest clade of lichen-forming fungi". Fungal Diversity. 84: 101–117. doi:10.1007/s13225-017-0379-z. S2CID 40674310.
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