Parmelia Subrudecta
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''Punctelia subrudecta'' is a species of
foliose lichen A foliose lichen is a lichen with flat, leaf-like , which are generally not firmly bonded to the substrate on which it grows. It is one of the three most common growth forms of lichens. It typically has distinct upper and lower surfaces, each o ...
in the family
Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species 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'' ( 82 ...
. Originally described in 1886 as ''Parmelia subrudecta'', it was transferred to the genus ''
Punctelia ''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 presenc ...
'' in 1982. For much of the twentieth century, North American specimens were misidentified as this species, until
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies in 2010 revealed that they represented three distinct species, restricting the true ''P. subrudecta'' to
Old World The "Old World" () is a term for Afro-Eurasia coined by Europeans after 1493, when they became aware of the existence of the Americas. It is used to contrast the continents of Africa, Europe, and Asia in the Eastern Hemisphere, previously ...
populations. The lichen forms loosely attached rosettes with narrow, pale grey-green and is distinguished by its pale underside, fine powdery reproductive structures (
soredia Soredia are common reproduction, reproductive structures of lichens. Lichens asexual reproduction, reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungus, fung ...
), and distinctive hook-shaped
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
. It is found primarily in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
and
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
woodlands of Europe,
Macaronesia Macaronesia (; ) is a collection of four volcanic archipelagos in the Atlantic Ocean#North Atlantic, North Atlantic Ocean, off the coast of North Africa and Europe. Each archipelago is made up of a number of list of islands in the Atlantic Oc ...
, and parts of North and East Africa, where it grows on tree bark and occasionally on rock surfaces.


Taxonomy

The Finnish lichenologist
William Nylander William Andrew Michael Junior Nylander Altelius (born 1 May 1996) is a Swedish professional ice hockey Forward (ice hockey), forward for the Toronto Maple Leafs of the National Hockey League (NHL). Nylander was selected by the Maple Leafs in t ...
described the species in 1886 as ''Parmelia subrudecta''. In his succinct Latin Nylander wrote that the thallus "resembles ''P. rudecta'' Ach., agreeing even in the CaCl2 reaction, yet differs by its sublageniform (flask-shaped) , which bear 4.5 × 0.6 
μm The micrometre (Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a unit of length in the International System ...
; apothecia were not seen, and the type material was rupicolous (rock-dwelling)".
Hildur Krog Hildur Krog (22 March 1922 – 25 August 2014) was a Norwegian lichenologist and university professor who made contributions to the field of lichenology, particularly in the areas of lichen taxonomy (biology), taxonomy, chemotaxonomy, and floris ...
transferred the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
to the newly erected genus ''
Punctelia ''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 presenc ...
'' in 1982, retaining Nylander's
epithet An epithet (, ), also a byname, is a descriptive term (word or phrase) commonly accompanying or occurring in place of the name of a real or fictitious person, place, or thing. It is usually literally descriptive, as in Alfred the Great, Suleima ...
and placing the species within
Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species 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'' ( 82 ...
. For much of the twentieth century North-American material with a pale lower surface and a
medulla Medulla (Latin for "marrow") or medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ...
containing
lecanoric acid Lecanoric acid is a chemical produced by several species of lichen.ubChem - Lecanoric acid"> Lecanoric acid is classified as a polyphenol and a didepside, and it functions as an antioxidant. It is an ester of orsellinic acid with itself. The acid ...
was routinely identified as ''P. subrudecta''. A continent-wide molecular and morphological study by James Lendemer and Brendan Hodkinson published in 2010 demonstrated that these specimens represent three distinct species—'' P. caseana'', '' P. jeckeri''; (≡ ''P. ulophylla''), and '' P. perreticulata''—none of which is
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organism ...
with the European ''P. subrudecta'' ''
sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
'' (in the strict sense). Their
ITS ITS, its or it's may refer to: Language * It's, an English contraction of ''it is'' or ''it has'' * Its (pronoun), the possessive form of the pronoun it * Itsekiri language (ISO 639 language code its), a language found in Nigeria and the Niger De ...
phylogeny recovered three well-supported
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s that correlated with traits previously dismissed as unreliable, namely
conidia A conidium ( ; : conidia), sometimes termed an asexual chlamydospore or chlamydoconidium (: chlamydoconidia), is an asexual, non- motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also ...
l type and length ( vs long ), the presence or absence of on the upper , and the development of on lobe tips. By integrating these characters with sequence data, the authors resurrected conidial morphology as a useful diagnostic feature and restricted the application of ''P. subrudecta'' to European and African populations that bear unciform (hook-like) conidia 4–6 Î¼m long. Within ''Punctelia'', ''P. subrudecta'' now represents the sorediate, lecanoric-acid-producing lineage with a pale undersurface and unciform conidia, distinct from the closely related ''P. perreticulata'' (short filiform conidia, scrobiculate upper surface), ''P. caseana'' (long filiform conidia, epruinose lobes), and ''P. jeckeri'' (long filiform conidia, pruinose lobes). The clarified circumscription means that confirmed records of ''P. subrudecta'' are confined to Europe, Macaronesia and parts of North Africa; all purported North-American reports have been reassigned. The study also provided a key to North-American ''Punctelia'' species and formally described ''P. caseana'', underscoring the importance of coupling DNA evidence with careful re-evaluation of traditional morphological markers when resolving species complexes within the Parmeliaceae.


Description

The
thallus Thallus (: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. A thallus usually names the entir ...
of ''Punctelia subrudecta'' is foliose and usually forms loosened rosettes several centimetres across. Lobes are rather narrow (mostly 2–4 mm wide), flat to slightly ridged, and display a pale grey-green colour that darkens when wet. The upper surface is smooth and lacks the shallow pits () that give ''P. perreticulata'' its look; it is also free of the frost-like bloom () that dusts the lobe tips of ''P. jeckeri''. Minute white pores (
pseudocyphella Pseudocyphellae (singular ''pseudocyphella'') are structures in lichens that appear as tiny pores on the outer surface (the cortex) of the lichen. They are caused when there is a break in the cortex of the lichen, and the medullary hyphae extend ...
e) are scattered but never abundant.
Vegetative reproduction Vegetative reproduction (also known as vegetative propagation, vegetative multiplication or cloning) is a form of asexual reproduction occurring in plants in which a new plant grows from a fragment or cutting of the parent plant or specializ ...
dominates in ''P. subrudecta''. Powdery
propagule In biology, a propagule is any material that functions in propagating an organism to the next stage in its life cycle, such as by dispersal. The propagule is usually distinct in form from the parent organism. Propagules are produced by organisms ...
s called
soredia Soredia are common reproduction, reproductive structures of lichens. Lichens asexual reproduction, reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungus, fung ...
—tiny balls of algal cells wrapped in fungal threads—break through the cortex in discrete patches known as
soralia Soredia are common reproductive structures of lichens. Lichens reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungal hyphae wrapped around cyanobacteria or g ...
. In ''P. subrudecta'' the soralia first appear along lobe edges and secondary lobes; older parts of the thallus may also develop scattered soralia, but the soredia remain fine and never coarsen into granules. The lower surface is characteristically pale off-white to light tan (never black) and bears sparse, similarly pale anchoring
rhizine In lichens, rhizines are multicellular root-like structures arising mainly from the lower surface. A lichen with rhizines is termed rhizinate, while a lichen lacking rhizines is termed erhizinate. Rhizines serve only to anchor the lichen to their s ...
s, a combination that helps separate the species from dark-undersided members of the genus. Microscopically the
pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...
produce (gently hook-shaped) conidia only 4–6 Î¼m long—one of the most reliable diagnostic characters, because related North-American taxa have straight, much longer conidia. Apothecia are rare; when present they are (thallus-coloured rim) and may develop sorediate margins. The
ascospore In fungi, an ascospore is the sexual spore formed inside an ascus—the sac-like cell that defines the division Ascomycota, the largest and most diverse Division (botany), division of fungi. After two parental cell nucleus, nuclei fuse, the ascu ...
s have been reported as broadly
ellipsoid An ellipsoid is a surface that can be obtained from a sphere by deforming it by means of directional Scaling (geometry), scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface;  that is, a Surface (mathemat ...
, 14–17 × 10–12 Î¼m, though most populations reproduce purely by soredia. Adler and Ahti measured 25 % of their worldwide collections and found that the unciform conidia of ''P. subrudecta'' have a modal length of 4–5.5 Î¼m (range 3.5–6.5 Î¼m) and are sometimes intermixed with a few straight conidia, whereas lobe width spans (1–)2–4(–8) mm, overlapping completely with that of ''P. perreticulata''; they therefore dismissed lobe width as a diagnostic trait. Chemical spot tests are distinctive: the cortex contains atranorin (K+ yellow), while the medulla is rich in lecanoric acid and turns pale pink to
salmon Salmon (; : salmon) are any of several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the genera ''Salmo'' and ''Oncorhynchus'' of the family (biology), family Salmonidae, native ...
in the C test—a subtler hue than the deep crimson produced by ''P. borreri''. These reactions, together with the pale underside, fine soredia and hooked conidia, provide a concise field and laboratory profile that separates ''P. subrudecta'' from superficially similar grey sorediate lichens. Trace-level variation occurs in European material:
thin-layer chromatography Thin-layer chromatography (TLC) is a chromatography technique that separates components in non-volatile mixtures. It is performed on a TLC plate made up of a non-reactive solid coated with a thin layer of adsorbent material. This is called the sta ...
of 216 Polish specimens detected atranorin only in about 30 % of thalli, even though lecanoric acid was consistently present in the medulla. Microscope studies show that the is the green alga '' Trebouxia gelatinosa''. At high magnification the algal cells appear irregular in outline and have a thick wall; each cell contains a multi-lobed
chloroplast A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
with a single, ''decolorans''-type
pyrenoid Pyrenoids are sub-cellular phase-separated micro-compartments found in chloroplasts of many algae,Giordano, M., Beardall, J., & Raven, J. A. (2005). CO2 concentrating mechanisms in algae: mechanisms, environmental modulation, and evolution. ''An ...
—a carbon-fixing body that is pierced by parallel pyrenotubules and surrounded by starch grains called pyrenoglobules. The fungal
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
e either press directly against the algal wall or extend short, type-1 intraparietal
haustoria In botany and mycology, a haustorium (plural haustoria) is a rootlike structure that grows into or around another structure to absorb water or nutrients. For example, in mistletoe or members of the broomrape family, the structure penetrates th ...
into it.
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...
of Iberian material shows that ''P. subrudecta'' teams up with only a few, uncommon genetic lineages of ''T. gelatinosa'', two of which it shares with ''P. borreri''; this pattern points to a symbiosis that is selective yet locally adaptable.


Habitat and distribution

''Punctelia subrudecta'' is predominantly
corticolous This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found ...
, forming tightly attached rosettes on the bark of trees in temperate and Mediterranean woodlands; specimens examined from the
Iberian Peninsula The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, comprisin ...
were taken almost exclusively from trunks and branches, with only occasional collections on siliceous rock faces that mimic bark microclimates. Populations recorded in Atlantic oak forests and in the drier, eastern Mediterranean mountains show that the species tolerates a broad moisture gradient, provided atmospheric pollution remains low. Thalli from heavily trafficked coastal sites in eastern Spain show pink central
necrosis Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. The term "necrosis" came about in the mid-19th century and is commonly attributed to German pathologist Rudolf Virchow, who i ...
and marginal bleaching, symptoms the authors link to chronic oxidant and nitrogen pollution. Polish
herbarium A herbarium (plural: herbaria) is a collection of preserved plant biological specimen, specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sh ...
data show the species colonises a wide range of
phorophyte In botany, phorophytes are plants on which epiphytes grow. The term is composed of ''phoro'', meaning bearer or carrier and ''phyte'', meaning plant. Commensalistic interactions The epiphytes benefit from the habitats provided, but the phoroph ...
s, most frequently oak (''
Quercus An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
'', 26 % of records) but also alder (''
Alnus Alders are trees of the genus ''Alnus'' in the birch family Betulaceae. The genus includes about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species ext ...
'', 13 %), European ash (''
Fraxinus excelsior ''Fraxinus excelsior'', known as the ash, or European ash or common ash to distinguish it from other types of ash, is a flowering plant species in the olive family Oleaceae. It is native throughout mainland Europe east to the Caucasus and Alb ...
'', 11 %), small-leaved linden (''
Tilia cordata ''Tilia cordata'', the small-leaved lime or small-leaved linden, is a species of tree in the family Malvaceae, native to much of Europe. Other common names include little-leaf or littleleaf linden, or traditionally in South East England, pry or p ...
'', 11 %), willow (''
Salix Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...
'', 10 %) and a suite of other broad-leaved hosts; rare saxicolous thalli have even been found on wood and concrete. Sequence-verified collections confirm an essentially Old-World range stretching from
Fennoscandia __NOTOC__ Fennoscandia (Finnish language, Finnish, Swedish language, Swedish and ; ), or the Fennoscandian Peninsula, is a peninsula in Europe which includes the Scandinavian Peninsula, Scandinavian and Kola Peninsula, Kola peninsulas, mainland ...
south to the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
, eastwards into Kenya and at least parts of China and Australasia; records from the Americas are now considered misidentifications. Within Europe the species is widespread, having been documented in at least thirty countries, but it shows marked regional bias: in Poland, for example, it is largely confined to the
Carpathian The Carpathian Mountains or Carpathians () are a range of mountains forming an arc across Central Europe and Southeast Europe. Roughly long, it is the third-longest European mountain range after the Urals at and the Scandinavian Mountains at ...
and Sudeten foothills and is scarce in the north and west. Because of continuing declines linked to air pollution and roadside tree removal, ''P. subrudecta'' is listed as vulnerable on the Polish
Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is an inventory of the global conservation status and extinction risk of biological sp ...
and enjoys full legal protection there. National assessments echo a broader European trend: the lichen is now legally protected in Poland and Norway and is flagged as expanding northwards in areas where
sulphur dioxide Sulfur dioxide (IUPAC-recommended spelling) or sulphur dioxide (traditional Commonwealth English) is the chemical compound with the formula . It is a colorless gas with a pungent smell that is responsible for the odor of burnt matches. It is r ...
levels have fallen, yet remains threatened where mature roadside trees are felled for road modernisation.


References

{{Taxonbar, from=Q1454559 subrudecta Lichen species Taxa named by William Nylander (botanist) Lichens described in 1886 Lichens of Australasia Lichens of China Lichens of Europe Lichens of Macaronesia Lichens of Kenya