Normandina pulchella

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Normandina pulchella
Normandina pulchella - Flickr - pellaea (1).jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Eurotiomycetes
Order: Verrucariales
Family: Verrucariaceae
Genus: Normandina
Species:
N. pulchella
Binomial name
Normandina pulchella
(Borrer) Nyl. (1861)
Synonyms [1]
  • Endocarpon pulchellumBorrer (1831)
  • Verrucaria pulchellaBorrer (1831)
  • Coccocarpia pulchella(Borrer) C.Bab. (1855)
  • Lenormandia jungermanniaeNyl. (1855) [2]
  • Normandina jungermanniae(Nyl.) Nyl. (1855)
  • Lenormandia pulchella(Borrer) A.Massal. (1856)

Normandina pulchella, commonly known as the elf-ear lichen or blue heart, is a species of squamulose lichen in the family Verrucariaceae. This cosmopolitan species is widely distributed across both hemispheres, where it thrives in moist microhabitats. It favours moss-covered deciduous trees and rocks, often colonising over mosses and bryophytes. It occasionally grows on bare bark and on other lichens. Distinctive features of N. pulchella include its bluish-green squamules (scales) with sharply raised margins, non-reactivity to standard chemical spot tests, and growth in humid habitats. Initially, Nannochloris normandinae, a green alga, was thought to be its photobiont . More recent studies, however, have revised this understanding, with Diplosphaera chodatii now recognised as the algal partner.

Contents

First named and scientifically described by the English botanist William Borrer in 1831, the clarification of Normandina pulchella's place within the Verrucariaceae, facilitated by molecular phylogenetics analysis in 2010, resolved long-standing taxonomic uncertainties. Prior classifications had varied widely, placing N. pulchella within groups such as the Basidiomycota (i.e., as a basidiolichen) and Fungi incertae sedis, largely due to differing interpretations of the perithecia (fruiting bodies) found within the lichen. These discrepancies stemmed from confusion over whether the perithecia belonged to the lichen itself or were instead associated with a parasitic lichenicolous fungus.

Systematics

Taxonomic history

Lobes with bluish colouration Normandina pulchella 257885145.jpg
Lobes with bluish colouration

The lichen was originally formally described in 1831 by the English botanist William Borrer, who classified it within the genus Verrucaria . He coined the vernacular name "little filmy-leaved Verrucaria" due to its distinctive morphology. [3] The specific epithet pulchella is the Latin diminutive of pulchra, 'beautiful' or 'fair'. [4] He observed the lichen to be characterised by thin, membranous, greenish-grey, leaf-like scales, with these features transitioning from smooth, rounded forms to crowded, waved, and lobed configurations, whilst adorned with powdery granules. The underside of these scales is distinguished by a pale brown colour and woolly fibres. Borrer described the lichen's tubercles as nearly globular and black. These tubercles reveal only their apex through the thallus surface, exposing a gelatinous, brownish nucleus with a central pore. He noted the lichen's frequent occurrence on mossy trees in Sussex, and that it thrived on the leafy liverwort species Jungermannia dilatata , forming wide but often interrupted patches. Ellen Hutchins was credited by Borrer for initially discovering the species on a mountain near Bantry (Ireland), growing on Lichen plumbeus on heath stems. He remarked on the lichen's uniqueness: [5]

This curious little production is so unlike to every other Lichen, that its very genus must have remained doubtful but for Miss Hutchins's fortunate discovery of the tubercles. Acharius, to whom Sussex specimens were communicated, thought it a Thelephora, thus excluding it even from the natural order to which we hold it to belong.

Despite its prevalence in Sussex, Borrer mentioned that it appeared to have been overlooked elsewhere. [5] In another publication that year, Borrer proposed to transfer it to the genus Endocarpon . However, this was nomenclaturally invalid because the name had already been for a different species, violating the International Code of Botanical Nomenclature guidelines. [3] The Finnish taxonomist William Nylander transferred the taxon to the genus Normandina in 1861. [6]

Classification

The taxonomic placement of N. pulchella was subject to significant debate among lichenologists for many decades. [7] Although the lichen produces ascospores, the origin of these spores and their associated structures was historically contentious. Prior to molecular studies, researchers proposed various classifications, including placement within the Basidiomycota due to morphological similarities with certain basidiolichens, particularly Coriscium viride, associated with the basidiomycete Lichenomphalia hudsoniana . The central controversy stemmed from perithecia found within the thallus, which were later attributed to the parasitic fungus Sphaerulina chlorococca rather than to N. pulchella itself. Some researchers considered these structures to be the lichen's own reproductive organs, while others interpreted them as belonging to a parasitic fungus. [7] The discovery that Normandina lacks dolipore septa — a key feature of basidiomycetes — cast doubt on its classification as a basidiolichen, but its exact systematic position remained unclear until 2010, when molecular phylogenetics analysis definitively established its position within the Verrucariaceae. This classification was further supported by the presence of features typical of ascomycetes, such as Woronin bodies and simple perforated septa. [8]

Microscopic cross section of a Normandina pulchella lobe illustrating its layered (heteromerous) structure Normandina pulchella microscopy.jpg
Microscopic cross section of a Normandina pulchella lobe illustrating its layered (heteromerous) structure

A cytological study revealed that N. pulchella has a heteromerous thallus, where the fungal and algal components are organised into distinct layers. This structure plays a key role in its life processes, distinguishing it from basidiolichens. Specifically, the medulla, or inner layer, contains hyaline (transparent) fungal filaments entwined around small clusters of algal cells. These cells have thick walls and a singular, lobate chloroplast containing a distinct metameric pyrenoid – a structure assisting in starch formation, along with small starch grains and fat-containing plastoglobuli. [8]

The fungal partner, or mycobiont , was found to have hyphae featuring simple perforated septa, a type of internal cell division within the hyphae, accompanied by Woronin bodies – organelles unique to ascomycetes, a large fungal group. Additionally, the close yet non-invasive relationship between the fungal and algal cells suggested a mutualistic association typical of ascolichens. [8] This cytological evidence, along with the 2010 molecular phylogenetics analysis by Lucia Muggia and colleagues, conclusively established N. pulchella's position within the family Verrucariaceae. [9]

Common names

In addition to Borrer's original suggestion ("little filmy-leaved Verrucaria"), other vernacular names that have been used to refer to this lichen are "blue heart", [10] and "elf ear lichen". [11]

Description

Close-up of sorediate lobes Normandina pulchella 176415411.jpg
Close-up of sorediate lobes

Normandina pulchella has a squamulose (scaly) thallus, composed of small, scale-like formations termed squamules . The squamules display colours ranging from glaucous—a bluish-green or grey—to pale grey or greenish-grey, intensifying to a richer green when moistened. Each squamule can span up to 5 mm across and may include one or more broadly rounded lobes , each up to 1.7 mm in diameter, reminiscent of shell or ear shapes. The upper surface of these lobes features concentric ridges, while the edges are sharply defined and raised, typically spanning 50–100  micrometres (μm) in width. The squamules may be dispersed or densely packed across the lichen's surface. [12] The thallus is closely appressed to the substrate . [11] Internally, the thallus shows a heteromerous structure, with distinct layering of fungal and algal components. [7] The medulla contains transparent fungal filaments that form a network around clusters of algal cells. These fungal hyphae feature simple perforated septa with distinctive Woronin bodies, while the algal cells are characterised by thick cell walls and single lobate chloroplasts containing pyrenoids. [8]

For vegetative reproduction, N. pulchella develops soralia—structures on the lobe surfaces and edges—that discharge granular particles known as soredia . These soralia are green or match the colour of the lobes and contain granular soredia measuring 40–80 μm in diameter. The underside of the lichen presents a whitish, slightly felted (tomentose) appearance and adheres to its substrate through numerous fungal strands, or hyphae. [12] Rhizines do not occur in this species. [11]

Zeorin is the only lichen product that occurs in Normandina pulchella. Zeorin.svg
Zeorin is the only lichen product that occurs in Normandina pulchella.

Its spore-producing structures, or ascomata, appear as semi-immersed globular to slightly conical perithecia with moderate orange-brown pigmentation in the wall of textura angularis. They are characterised by abundant periphyses , absence of paraphyses and hymenial algae, and an I+ red/KI+ blue hymenial gel. [7] They resemble those in related species but are distinguished by their uniformly pigmented, cohesive nature under microscopic analysis. The ascospores are typically 29–37 by 6–7 μm, mostly with seven internal partitions, known as septa, [12] though some specimens show predominantly (4–5)-septate spores measuring 17–29 by 5–7 μm. [7] Chemical analysis, particularly thin-layer chromatography, identify zeorin as a secondary metabolite (lichen product), yet N. pulchella remains unresponsive to standard chemical spot tests. [12]

Similar species

The squamules of Lichenomphalina hudsoniana share the neat, sharply defined edges characteristic of N. pulchella. However, they can be distinguished by their lack of soralia, the presence of both upper and lower cortices, and adaptation to arctic-alpine environments, typically growing on peaty soil or decaying wood. In contrast, N. pulchella tends to develop more pronounced soralia in shaded, humid settings, diverging from the preferred habitats of Lichenomphalina hudsoniana. Additionally, the fruiting bodies of N. pulchella are found to be more prevalent and larger in tropical regions compared to temperate ones, and they often do not contain soredia. This suggests that environmental factors or underlying taxonomic differences may influence the observed variations between these species in different locales. [12] [11]

Photobiont

In 1981, Elisabeth Tschermak-Woess identified Nannochloris normandinae as the photobiont partner, the algae associated with Normandina pulchella. [13] Subsequent studies, however, have been less definitive about the role of Nannochloris. [14] By 2011, research by Thüs and colleagues revealed that Diplosphaera , not Nannochloris, was present in ten examined Normandina specimens. [15] In 2020, Pröschold and Darienko demonstrated that Nannochloris normandinae is actually synonymous with Diplosphaera chodatii of the order Prasiolales, resolving the apparent conflict in previous studies and confirming D. chodatii as the photobiont of N. pulchella. [16] [17]

Habitat and distribution

Normandina pulchella growing amongst moss on tree bark Normandina pulchella - Flickr - pellaea.jpg
Normandina pulchella growing amongst moss on tree bark

Normandina pulchella has a cosmopolitan distribution, growing across diverse climates and regions. [12] The Swedish lichenologist Gunnar Degelius, in his 1934 phytogeographical study, highlights the species' oceanic distribution in Europe. It predominantly occupies coastal areas in Northern Europe, including Scandinavia and the British Isles, and extends to both coastal and montane regions in Austria, Bavaria, France, Czechoslovakia (now the Czech Republic and Slovakia), and select Mediterranean locales. [18] In contrast, William Louis Culberson and Mason Hale's 1966 analysis of its North American presence noted its prevalence in the mountainous western regions and the Appalachian foothills, without indicating an oceanic distribution pattern. [19] Its range in North America extends north to Alaska, [20] although in 2022 Alan Orange showed that some collections from there represent a different species found in the Americas, provisionally named N. 'americana'. [21]

Ecologically, Normandina pulchella favours moss-covered deciduous trees and rocks within woodlands and parks, often colonising over mosses, bryophytes, and occasionally bare bark. [12] It also often grows on other lichens, particularly those that contain cyanobacteria, [22] such as Fuscopannaria , Pannaria , Parmeliella , Pectenia , and Peltigera . Its presence is increasing in southern and western Britain and throughout Ireland, reflecting a broadening distribution. [12]

It is listed as a vulnerable species in the Finnish Regional Red List because of its small known population. [23]

Species interactions

Among the lichenicolous (lichen-dwelling) fungi specifically associated with N. pulchella are several species that have unique interactions. Capronia normandinae is characterised by its black, superficial, hair-like structures known as setose perithecia. Cladophialophora normandinae is distinguished by its black fruiting bodies, termed sporodochia, which play a role in its reproductive cycle. Additionally, Tremella normandinae is noted for producing pale, swollen growths, referred to as galls, indicative of its parasitic relationship with the lichen. Another parasite, Globosphaeria jamesii , also interacts with Normandina pulchella, further contributing to the diversity of its ecological associations. [12] [24]

Lawreymyces pulchellae is a fungus that lives hidden within Normandina pulchella tissues, one of several recently discovered cryptic basidiomycete-lichen associations. Described in 2017 as part of the family Corticiaceae, it differs from typical lichenicolous fungi by leaving no visible signs of its presence, being detectable only through molecular methods. It belongs to a group of Lawreymyces species that specifically associate with lichens in the family Verrucariaceae, particularly Normandina and Agonimia . This relationship parallels that of Cyphobasidium in Parmeliaceae lichens. The species is identifiable only through its unique internal transcribed spacer DNA sequence. [25]

Related Research Articles

<span class="mw-page-title-main">Basidiolichen</span> Lichen with a Basidiomycota mycobiont

Basidiolichens are lichenized members of the division Basidiomycota within the subkingdom Dikarya of the kingdom Fungi. They form a diverse yet much smaller group of lichens than the far more common ascolichens of the division Ascomycota. Owing to how few described species there are, basidiolichens are generally considered to be poorly researched, and few studies that characterize their natural products exist. Biogeographically, basidiolichen species may be distributed in a cosmopolitan manner or more regionally, ranging from arctic and montane habitats to more temperate and tropical environments. Morphologically, basidiocarp and thallus structures may vary widely within and between basidiolichen genera.

<span class="mw-page-title-main">Verrucariales</span> Order of fungi

Verrucariales is an order of ascomycetous fungi within the subclass Chaetothyriomycetidae of the class Eurotiomycetes. Although most of the Verrucariales are lichenised, the family Sarcopyreniaceae consists of 11 species of lichenicolous (lichen-dwelling) fungi.

<span class="mw-page-title-main">Verrucariaceae</span> Family of mostly lichenised fungi

Verrucariaceae is a family of lichens and a few non-lichenised fungi in the order Verrucariales. The lichens have a wide variety of thallus forms, from crustose (crust-like) to foliose (bushy) and squamulose (scaly). Most of them grow on land, some in freshwater and a few in the sea. Many are free-living but there are some species that are parasites on other lichens, while one marine species always lives together with a leafy green alga.

<span class="mw-page-title-main">Lepidostromatales</span> Order of fungi

Lepidostromatales is an order of fungi in the class Agaricomycetes. It is the only known order of basidiomycete fungi composed entirely of lichenized members. Morphologically, the fruiting bodies of all species are clavarioid. Six species are known, five of which were described within the span of 2007–2013. Due to its morphological similarity to the genus Multiclavula, its isolated phylogenetic position was not understood until quite recently. The photobionts that have been found in association with members of this group are not known to associate with any other types of lichenized fungi.

<i>Hydropunctaria</i> Genus of lichen

Hydropunctaria is a genus of saxicolous (rock-dwelling), crustose lichens in the family Verrucariaceae. The genus includes both aquatic and amphibious species, with members that colonise either marine or freshwater habitats. The type species, Hydropunctaria maura, was formerly classified in the large genus Verrucaria. It is a widely distributed species common to littoral zones. Including the type species, five Hydropunctaria lichens are considered marine species: H. adriatica, H. amphibia, H. aractina, H. orae, and H. oceanica.

<i>Willeya</i> Genus of lichens

Willeya is a genus of saxicolous (rock-dwelling), crustose lichens in the family Verrucariaceae. It has 12 species. Most species are found in southeast Asia, although individual representatives are known from Australia, Europe, and North America.

<i>Placidium arboreum</i> Species of lichen

Placidium arboreum, commonly known as the tree stipplescale, is a species of corticolous (bark-dwelling), squamulose (scaley) lichen in the family Verrucariaceae. Placidium arboreum is primarily found in the southeastern United States, but it also has occurrences in the western and northeastern United States, Mexico, the West Indies, Argentina, and Ontario, Canada. In its habitat, it typically grows at the base of hardwood trees, particularly oak species, and can occasionally be found on other tree genera or even over mosses on limestone. Its preferred substrate is the bark of oak trees, and the lichen usually establishes itself in bark crevices.

<i>Mastodia</i> Genus of lichens

Mastodia is a genus of lichen-forming fungi in the family Verrucariaceae. It has six species.

Verrucaria oulankaensis is a rare species of saxicolous (rock-dwelling) crustose lichen in the family Verrucariaceae. It is found in north-eastern Finland, where it occurs on calcareous rocks on river shores.

Wahlenbergiella tavaresiae is a species of saxicolous (rock-dwelling), crustose lichen in the family Verrucariaceae. Known from several locations in the San Francisco Bay area of the United States, it is a marine lichen that inhabits intertidal zones, and as such is immersed in seawater on a regular basis. Associated algal species include the red algae Hildenbrandia and Mastocarpus papillatus, and the brown algae Pelvetiopsis and Fucus. Petroderma maculiforme, a brown alga, is the photobiont partner in the lichen.

Agonimia flabelliformis is a species of corticolous (bark-dwelling) lichen in the family Verrucariaceae. Found in Europe, it was scientifically described as new to science in 2011 by Josef Halda, Paweł Czarnota, and Beata Guzow-Krzemińska. The type specimen was collected in the Gratzen Mountains at an altitude of 805 m (2,641 ft), where it was found growing on the bark of a beech tree. The thallus of the lichen consists of finger-like (dactyliform) to coral-like (coralloid) aggregations of goniocysts that form a roundish structure. The species epithet flabelliformis refers to the fan-shaped (flabellate) form of the thallus branches. In addition to Europe, the lichen has also been recorded in eastern North America and the Russian Far East.

<i>Verrucaria viridula</i> Species of lichen

Verrucaria viridula is a common and widely distributed species of saxicolous (rock-dwelling), crustose lichen in the family Verrucariaceae. Although it is a somewhat morphologically variable species, two persistent distinguishing characteristics are its relatively large perithecia, which are often curved into a beak, and its large ascospores.

Protothelenella is a genus of fungi in the family Protothelenellaceae. It contains 11 species, some of which form lichens. Protothelenella species have a crustose thallus with spherical to pear-shaped, dark brown to blackish perithecia. Microscopic characteristics of the genus include bitunicate asci with an amyloid tholus, and ascospores that are colourless and contain multiple internal partitions. Some species grow on acidic substrates including rocks, soil, bryophytes, plant detritus or rotten wood. Other species are lichenicolous (lichen-dwelling), growing on species of Solorina, Peltigera, Pseudocyphellaria, or Cladonia.

<i>Hydropunctaria amphibia</i> Species of lichen

Hydropunctaria amphibia is a species of saxicolous (rock-dwelling), crustose lichen in the family Verrucariaceae. One of several marine lichens in the genus Hydropunctaria, is widely distributed across Europe, extending from Norway to the Mediterranean and the Iberian coasts, and has a nearly ubiquitous presence along the Catalan coast of Spain. In North America, it is found along the Atlantic Coast from Nova Scotia to the Boston Harbor islands, where its presence in low-pollution areas indicates its potential as a bioindicator for marine lichen community health, and on the west coast in British Columbia, particularly in the Gwaii Haanas's upper littoral fringe. The black, crust-like thallus grows on seashore rocks – both siliceous rocks and limestone – in the lower supralittoral zone, an area also known as the splash zone. Originally described more than two centuries ago as a species of Verrucaria, Hydropunctaria amphibia sets itself apart from other species in Hydropunctaria through the distinct shape of the perithecium apex, which is either flat-topped or scalloped, in contrast to the typically rounded or immersed apex seen in its relatives.

Placidiopsis hypothallina is a species of saxicolous (rock-dwelling) crustose lichen in the family Verrucariaceae, described as a new species in 2002. It is unique within its genus for its thallus primarily composed of a hyphal hypothallus resembling a subiculum found in some non-lichenised ascomycetes. It is found in Brazil and Taiwan.

Cyanoporina is a fungal genus in the division Ascomycota. The relationship of this taxon to other taxa within the division is unknown, and it has not yet been placed with certainty into any class, order, or family. The genus is monospecific, containing the single species Cyanoporina granulosa, a crustose lichen found in West Java, Indonesia.

<i>Saxiloba</i> Genus of lichens

Saxiloba is a genus of lichen-forming fungi in the family Porinaceae. It comprises three species of saxicolous (rock-dwelling) placodioid lichens, known from disjunct locations in the Caribbean, Hawaii, and Brazil. The genus was circumscribed in 2020 to accommodate species with a distinctive placodioid thallus featuring unique surface patterns and internal crystal structures. Saxiloba lichens are characterised by their flattened, leaf-like thalli with marginal lobes, growing tightly appressed to rock surfaces. They have a complex internal structure, including large crystal clusters embedded within the photobiont layer, which may be an adaptation for light management in their typically shaded habitats.

Pyrenothrix is a small genus of lichen-forming fungi in the family Pyrenotrichaceae. It comprises two species of filamentous lichens, which are organisms formed by a symbiotic relationship between fungi and photosynthetic partners. The genus is characterized by its unique structure, featuring densely arranged filaments composed of cyanobacteria wrapped in fungal threads. Pyrenothrix species form dark greyish-brown growths on various surfaces, with one species found on tree bark and the other on leaves in tropical forests. The genus was circumscribed in 1917 by American scientist Lincoln Ware Riddle, based on specimens collected in Florida. Pyrenothrix is distinguished from other lichens by its intricate cellular structure and reproductive features, including specialized spore-producing structures.

Capronia normandinae is a species of lichenicolous (lichen-dwelling) fungus in the family Herpotrichiellaceae. The fungus was first formally described in 1990 by Rolf Santesson and David Hawksworth. The fungus has been recorded from Papua New Guinea, the Atlantic Ocean, Australasia, Europe, and South America. The fungus parasitises the host lichen Normandina pulchella, after which it is named.

Globosphaeria is a fungal genus that grows on lichens, containing the single species Globosphaeria jamesii. The fungus is characterised by dark, spherical fruiting bodies producing colourless, round spores, and is found growing on the lichen Normandina pulchella. First collected in Tasmania in 1907 but not described until 1990, it was later also discovered in the United States, making it known from just two locations globally. Though initially placed in the Xylariales, the genus is now classified in the order Sordariales, though its exact family placement remains uncertain.

References

  1. ↑ "Synonymy. Current Name: Normandina pulchella (Borrer) Nyl., Annls Sci. Nat., Bot., sér. 4 15: 382 (1861)". Species Fungorum . Retrieved 27 March 2024.
  2. ↑ Nylander, William (1855). "Additamentum in floram cryptogamicam Chilensem". Annales des Sciences Naturelles. Botanique. 4 (in Latin). 3: 145–187 [151].
  3. 1 2 "Record Details: Endocarpon pulchellum Borrer, in Hooker & Sowerby, Suppl. Engl. Bot. 1: tab. 2602 (1831)". Index Fungorum . Retrieved 27 March 2024.
  4. ↑ Bomfleur, Benjamin; Grimm, Guido W.; McLoughlin, Stephen (2015). "Osmunda pulchella sp. nov. from the Jurassic of Sweden—reconciling molecular and fossil evidence in the phylogeny of modern royal ferns (Osmundaceae)". BMC Evolutionary Biology. 15 (1): e15. Bibcode:2015BMCEE..15..126B. doi: 10.1186/s12862-015-0400-7 . PMC   4487210 . PMID   26123220.
  5. 1 2 Borrer, William (1831). Supplement to the English Botany of the late Sir J. E. Smith and Mr. Sowerby. Vol. 1. London: J. D. C. and C. E. Sowerby. tab. 2602, fig. 1.
  6. ↑ Nylander, William (1861). "Additamentum ad lichenographiam Andium Boliviensium" [Supplement to the study of lichens of the Bolivian Andes]. Annales des Sciences Naturelles. Botanique. 4 (in Latin). 15: 365–382 [382].
  7. 1 2 3 4 5 Frisch, Andreas; Ohmura, Yoshihito (2015). "The phylogenetic position of Normandina simodensis (Verrucariaceae, lichenized Ascomycota)". Bulletin of the National Museum of Nature and Science, Series B (Botany). 41 (1): 1–7.
  8. 1 2 3 4 Mares, Donatello; Fasulo, Maria P.; Bruni, Alessandro (1993). "Contribution to the study of Normandina pulchella: a cytological approach". Orsis. 8: 33–40.
  9. ↑ Muggia, Lucia; Gueidan, Cécile; Grube, Martin (2010). "Phylogenetic placement of some morphologically unusual members of Verrucariales". Mycologia. 102 (4): 835–846. doi:10.3852/09-153. PMID   20648751.
  10. ↑ Tripp, Erin A.; Lendemer, James C. (2020). Field Guide to the Lichens of Great Smoky Mountains National Park. Knoxville: The University of Tennessee Press. pp. 308–309. ISBN   978-1-62190-514-1.
  11. 1 2 3 4 McMullin, R. Troy (2023). Lichens. The Macrolichens of Ontario and the Great Lakes Region of the United States. Firefly Books. p. 311. ISBN   978-0-228-10369-1.
  12. 1 2 3 4 5 6 7 8 9 Orange, A.; Cannon, P.; Prieto, M.; Coppins, B.; Sanderson, N.; Simkin, J. (2023). Verrucariales: Verrucariaceae, including the genera Agonimia, Atla, Bagliettoa, Catapyrenium, Dermatocarpon, Endocarpon, Henrica, Heteroplacidium, Hydropunctaria, Involucropyrenium, Merismatium, Nesothele, Normandina, Parabagliettoa, Placidopsis, Placidium, Placopyrenium, Polyblastia, Psoroglaena, Sporodictyon, Staurothele, Thelidium, Trimmatothele, Verrucaria, Verrucula, Verruculopsis and Wahlenbergiella (PDF). Revisions of British and Irish Lichens. Vol. 31. British Lichen Society. p. 43.
  13. ↑ Tschermak-Woess, E. (1981). "Zur Kenntnis der Phycobionten von Lobaria linita und Normandina pulchella" [On the knowledge of the phycobionts of Lobaria linita and Normandina pulchella]. Nova Hedwigia (in German). 35: 63–73.
  14. ↑ Lohtander, Katileena; Oksanen, Ilona; Rikkinen, Jouko (2003). "Genetic diversity of green algal and cyanobacterial photobionts in Nephroma (Peltigerales)". The Lichenologist. 35 (4): 325–339. doi:10.1016/S0024-2829(03)00051-3.
  15. ↑ Thüs, Holger; Muggia, Lucia; Pérez-Ortega, Sergio; Favero-Longo, Sergio E.; Joneson, Suzanne; O’Brien, Heath; Nelsen, Matthew P.; Duque-Thüs, Rhinaixa; Grube, Martin; Friedl, Thomas; Brodie, Juliet; Andrew, Carrie J.; Lücking, Robert; Lutzoni, François; Gueidan, Cécile (2011). "Revisiting photobiont diversity in the lichen family Verrucariaceae (Ascomycota)". European Journal of Phycology. 46 (4): 399–415. Bibcode:2011EJPhy..46..399T. doi:10.1080/09670262.2011.629788.
  16. ↑ Pröschold, Thomas; Darienko, Tatyana (2020). "The green puzzle Stichococcus (Trebouxiophyceae, Chlorophyta): new generic and species concept among this widely distributed genus". Phytotaxa. 441 (1): 113–142. doi:10.11646/phytotaxa.441.2.2.
  17. ↑ Veselá, Veronika; Malavasi, Veronica; Škaloud, Pavel (2024). "A synopsis of green-algal lichen symbionts with an emphasis on their free-living lifestyle". Phycologia. 63 (3): 317–338. doi: 10.1080/00318884.2024.2325329 . hdl: 11577/3511125 .
  18. ↑ Degelius, Gunnar (1935). Das oceanische Element der Strauch- und Laubflechtenflora von Skandinavien[The oceanic element of the shrub and leaf lichen flora of Scandinavia]. Acta Phytogeographica Suecica (in German). Vol. 7.
  19. ↑ Culberson, William Louis; Mason E., Hale (1966). "The range of Normandina pulchella in North America". The Bryologist. 69 (3): 365–367. doi:10.2307/3240836. JSTOR   3240836.
  20. ↑ Spribille, Toby; Fryday, Alan M.; Hampton-Miller, Celia J.; Ahti, Teuvo; Dillman, Karen; Thor, Göran; Tonsberg, Tor; Schirokauer, Dave, eds. (2023). Compendium of the Lichens and Associated Fungi of Alaska. Bibliothecia Lichenologica. J. Cramer. p. 278. doi: 10.1127/bibl_lich/2023/112 . ISBN   978-3-443-58093-3.
  21. ↑ Orange, Alan (2022). "The crustose species of Normandina Verrucariaceae)". The Lichenologist. 54 (6): 371–378. doi:10.1017/s0024282922000317.
  22. ↑ Sharnoff, Stephen (2014). A Field Guide to California Lichens. New Haven/London: Yale University Press. p. 357. ISBN   978-0-300-19500-2.
  23. ↑ Hyvärinen, Esko; Juslén, Aino; Kemppainen, Eija; Uddström, Annika; Liukko, Ulla-Maija, eds. (2019). Suomen lajien uhanalaisuus – Punainen kirja 2019. The 2019 Red List of Finnish Species (Report). Ministry of the Environment. Finnish Environment Institute. p. 292. ISBN   978-952-11-4974-0.
  24. ↑ Diederich, Paul; Lawrey, James D.; Ertz, Damien (2018). "The 2018 classification and checklist of lichenicolous fungi, with 2000 non-lichenized, obligately lichenicolous taxa". The Bryologist. 121 (3): 385, 392. doi:10.1639/0007-2745-121.3.340.
  25. ↑ Lücking, Robert; Moncada, Bibiana (2017). "Dismantling Marchandiomphalina into Agonimia (Verrucariaceae) and Lawreymyces gen. nov. (Corticiaceae): setting a precedent to the formal recognition of thousands of voucherless fungi based on type sequences". Fungal Diversity. 84 (1): 119–138. doi:10.1007/s13225-017-0382-4.
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