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 . However, recent studies have revised this understanding, now suggesting Diplosphaera 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: "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] William Nylander transferred the taxon to the genus Normandina in 1861. [6]

Classification

Recent advancements have clarified the classification of Normandina pulchella, addressing ambiguities and refining its placement within the systematic taxonomy. Initially, perithecia found within the thallus led to confusion, as these were later attributed to the parasitic fungus Sphaerulina chlorococca rather than to Normandina pulchella itself. This misunderstanding contributed to significant taxonomic challenges regarding the lichen's classification. Historically, N. pulchella was often classified as a basidiolichen, largely due to its morphological similarities with Coriscium viride, associated with the basidiomycete Omphalina hudsoniana. However, the discovery that Normandina lacks dolipore septa — key features of basidiomycetes — cast doubt on its classification as a basidiolichen. Despite this, the exact systematic position of Normandina remained unresolved. [7]

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 Normandina pulchella has a complex structure with separate fungal and algal layers, a characteristic of heteromerous thalli. 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. [7]

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. The exclusive presence of Woronin bodies and the characteristics of the algal partner align with traits typically found in ascomycetes and their associated algae in ascolichens. This evidence collectively shifted Normandina pulchella away from being classified as a basidiolichen, as some previous studies suggested, firmly placing it within the ascolichens. [7]

In 2010, Lucia Muggia and colleagues employed molecular phylogenetics analysis to conclusively classify Normandina pulchella within the family Verrucariaceae, resolving longstanding classification debates. [8]

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", [9] and "elf ear lichen". [10]

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. [11] The thallus is closely appressed to the substrate . [10]

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. [11] Rhizines do not occur in this species. [10]

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, 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. Chemical analysis, particularly thin-layer chromatography, identify zeorin as a secondary metabolite (lichen product), yet N. pulchella remains unresponsive to standard chemical spot tests. [11]

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. [11] [10]

Photobiont

In 1981, Elisabeth Tschermak-Woess identified Nannochloris normandinae as the photobiont partner, the algae associated with Normandina pulchella. [12] Subsequent studies, however, have been less definitive about the role of Nannochloris. [13] By 2011, research by Thüs and colleagues revealed that Diplosphaera , not Nannochloris, was present in ten examined Normandina specimens. [14] More recently, Pröschold & Darienko (2020) reclassified Nannochloris normandinae as synonymous with Diplosphaera chodatii of the order Prasiolales, [15] further complicating the identification of the true photobiont in N. pulchella and suggesting that previous attributions to Nannochloris may be incorrect. [14]

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. [11] 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 Austria, Bavaria, France, Czechoslovakia, and select Mediterranean locales. [16] 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. [17] Its range in North America extends north to Alaska, [18] although in 2022 Alan Orange showed that some collections from there represent a different species found in the Americas, provisionally named N. 'americana'. [19]

Ecologically, Normandina pulchella favours moss-covered deciduous trees and rocks within woodlands and parks, often colonising over mosses, bryophytes, and occasionally bare bark. [11] It also often grows on other lichens, particularly those that contain cyanobacteria, [20] such as Fuscopannaria , Pannaria , Parmeliella , Pectenia , and Peltigera . Its presence is increasing in southern and western Britain and throughout Ireland, reflecting a broadening distribution. [11]

It is listed as a vulnerable in the Finnish Regional Red List because of its small known population. [21]

Species interactions

Among the lichenicolous (lichen-dwelling) fungi specifically associated with Normandina 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. Moreover, Lawreymyces pulchellae has an endolichenic relationship, residing within the lichen's structure. [11] [22]

Related Research Articles

<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.

<i>Normandina</i> Genus of lichens

Normandina is a genus of lichen-forming fungi in the family Verrucariaceae. It has five species of crustose and squamulose (scaly) lichens.

<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>Parmelia barrenoae</i> Species of lichen

Parmelia barrenoae is a species of foliose lichen in the large family Parmeliaceae. It was formally described as a new species in 2005. Before this, it was lumped together as one of several lichens in the Parmelia sulcata group—a species complex of genetically distinct lookalikes. Parmelia barrenoae is widely distributed, occurring in Europe, western North America, Africa, and Asia.

Acantholichen pannarioides is a species of basidiolichen in the family Hygrophoraceae, and the type species of genus Acantholichen. The lichen has a bluish-tinged, gelatinous thallus with a surface texture that has a powdery to hairy texture. It is found in montane regions of Central America and northern South America, where it grows on forest litter, bark, on bryophytes, and on other lichens.

Wilketalia is a fungal genus in the family Teloschistaceae. It is monotypic, containing the single species Wilketalia citrinoides, a saxicolous (rock-dwelling) crustose lichen found in the Bolivian Andes.

<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.

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.

Borinquenotrema is a single-species fungal genus in the family Graphidaceae. It contains the species Borinquenotrema soredicarpum, a corticolous (bark-dweling) lichen. Found in Puerto Rico, this lichen is characterized by its carbonizedascomata, which develop from within soralia, and its distinctive distoseptate, violet-blue ascospores. Borinquenotrema soredicarpum grows on tree trunks in shaded understory environments of Tabonuco forests in El Yunque National Forest.

Verrucaria kowenensis is a species of terricolous (ground-dwelling), crustose lichen in the family Verrucariaceae. It is found in the Australian Capital Territory of Australia, where it grows on silica-rich soil.

<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.

Anaptychia ethiopica is a species of lichen in the family Teloschistaceae. Found in East Africa, China, and Russia, it was formally described as a new species in 1976 by lichenologists Thomas Douglas Victor Swinscow and Hildur Krog. The type specimen was collected from Mount Bwahit, where it was found growing on moss.

<i>Kuettlingeria soralifera</i> Species of lichen

Kuettlingeria soralifera is a saxicolous (rock-dwelling), crustose lichen species in the family Teloschistaceae, first described in 2006. It is similar to Kuettlingeria xerica but distinguished by the presence of soredia on its thallus.

Austroplaca soropelta is a species of saxicolous and muscicolous, crustose lichen in the family Teloschistaceae. It has a bipolar distribution, meaning it occurs in polar areas of both the Northern and Southern Hemispheres.

<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.

Knightiellastrum is a single-species fungal genus in the family Icmadophilaceae. This monotypic genus the contains the corticolous (bark-dwelling), squamulose lichen species Knightiellastrum eucalypti, found in Tasmania, Australia.

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, W. (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. ↑ 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, W. (1861). "Additamentum ad lichenographiam Andium Boliviensium". Annales des Sciences Naturelles. Botanique. 4 (in Latin). 15: 365–382 [382].
  7. 1 2 3 Mares, Donatello; Fasulo, Maria P.; Bruni, Alessandro (1993). "Contribution to the study of Normandina pulchella: a cytological approach". Orsis. 8: 33–40.
  8. ↑ 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.
  9. ↑ 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.
  10. 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.
  11. 1 2 3 4 5 6 7 8 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.
  12. ↑ 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.
  13. ↑ 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.
  14. 1 2 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.
  15. ↑ 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.
  16. ↑ 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.
  17. ↑ 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.
  18. ↑ 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.
  19. ↑ Orange, Alan (2022). "The crustose species of Normandina Verrucariaceae)". The Lichenologist. 54 (6): 371–378. doi:10.1017/s0024282922000317.
  20. ↑ Sharnoff, Stephen (2014). A Field Guide to California Lichens. New Haven/London: Yale University Press. p. 357. ISBN   978-0-300-19500-2.
  21. ↑ 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.
  22. ↑ 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.
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