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''Parmelia pinnatifida'' 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 ...
, first recognised as a distinct species in 1976. Originally classified as a variety of '' P. omphalodes'' in 1803, it is characterised by its small, circular body (
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 ...
) with narrow, highly branched, overlapping , and its grey to brown upper surface contrasting with a black underside. While its status as a separate species has been debated, modern genetic studies support this classification, though some uncertainty remains. The lichen typically grows on
siliceous Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , commonly found in nature as quartz. In many parts of the world, silica is the major constituent of sand. Silica is one of the most complex and abundant ...
(silicon-rich) rocks, and occasionally on
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
, plant debris, or tree bark, showing a preference for humid environments. It has been documented across Europe, North America, and Antarctica, with populations in mountainous regions, though its reported presence in the Southern Hemisphere requires further verification.


Taxonomy

''Parmelia pinnatifida'' was originally described as ''Parmelia omphalodes'' var. ''panniformis'' by
Erik Acharius Erik Acharius (10 October 1757 – 14 August 1819) was a Swedish botanist who pioneered the Taxonomy (biology), taxonomy of lichens and is known as the "father of lichenology". Acharius was famously the last pupil of Carl Linnaeus. Life Ac ...
in 1803. However, its unique characteristics led Syo Kurokawa, in 1976, to recognise it as a distinct species. Kurokawa's reclassification was based on detailed studies of the lichen's
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
and chemistry, which revealed differences from closely related taxa such as '' P. omphalodes'' and '' P. discordans''. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''pinnatifida'' highlights the species' hallmark repeatedly branched , which form narrow, overlapping . Within the genus '' Parmelia'' (in the strict sense), ''P. pinnatifida'' belongs to a
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group known as the '' P. saxatilis'' group. This clade includes several closely related species such as '' P. discordans'', '' P. ernstiae'', '' P. hygrophila'', '' P. imbricaria'', '' P. mayi'', ''P. omphalodes'', ''P. saxatilis'', '' P. serrana'', '' P. submontana'', and '' P. sulymae''. The group has its centre of diversity 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 ...
regions of the Northern Hemisphere but has achieved a
cosmopolitan distribution In biogeography, a cosmopolitan distribution is the range of a taxon that extends across most or all of the surface of the Earth, in appropriate habitats; most cosmopolitan species are known to be highly adaptable to a range of climatic and en ...
. The taxonomic status of ''P. pinnatifida'' has been subject to debate over the years. In 1987,
Mason Hale Mason Ellsworth Hale, Jr. (September 23, 1929 – April 23, 1990) was one of the most prolific American lichenologists of the 20th century. Many of his scholarly articles focused on the taxonomy of the family Parmeliaceae. Hale was one of the fi ...
, in his
monograph A monograph is generally a long-form work on one (usually scholarly) subject, or one aspect of a subject, typically created by a single author or artist (or, sometimes, by two or more authors). Traditionally it is in written form and published a ...
of the genus ''Parmelia'', treated it as a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of ''P. omphalodes'', suggesting the morphological differences were insufficient to warrant species status. However, this classification was later challenged by
molecular A molecule is a group of two or more atoms that are held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions that satisfy this criterion. In quantum physics, organic chemistry, ...
studies. Modern phylogenetic analyses, using both ITS (
internal transcribed spacer Internal transcribed spacer (ITS) is the spacer DNA situated between the small-subunit ribosomal RNA (rRNA) and large-subunit rRNA genes in the chromosome or the corresponding transcribed region in the polycistronic rRNA precursor transcript. ...
) and β-
tubulin Tubulin in molecular biology can refer either to the tubulin protein superfamily of globular proteins, or one of the member proteins of that superfamily. α- and β-tubulins polymerize into microtubules, a major component of the eukaryotic cytosk ...
gene sequences, have definitively demonstrated that ''P. pinnatifida'' and ''P. omphalodes'' are genetically distinct taxa, supporting Kurokawa's original classification as separate species. The development of molecular phylogenetic techniques has been crucial in resolving relationships within the ''P. saxatilis'' group, though some taxonomic uncertainties remain. While the status of ''P. pinnatifida'' as a distinct species is supported by a combination of morphological, chemical, and molecular evidence, some authors still consider ''P. pinnatifida'' to be merely a
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
of ''P. omphalodes''. This taxonomic uncertainty, combined with its morphological similarity to ''P. omphalodes'', has historically led to ''P. pinnatifida'' being overlooked or misidentified in field surveys, potentially affecting our understanding of its true distribution.


Description

''Parmelia pinnatifida'' is characterised by a small, orbicular thallus that attaches loosely to its substrate. The thallus can vary in growth form from flat to cushion-forming. The species displays a grey to brown upper surface and a contrasting black lower surface. The thallus features narrow, highly branched lobes that are (overlapping like roof tiles) and somewhat ascending. These lobes measure 1–2 mm in width, with lobules measuring 0.1–0.3 mm in width. The species lacks both
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
isidia An isidium (plural: isidia) is a tiny, wart- or finger-like outgrowth on the thallus surface of certain lichen species. It is one of two principal types of vegetative reproduction, vegetative reproductive structures in lichens, the other being ...
, reproductive structures common in many other lichens. The upper surface of the thallus has
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, small pore-like structures, which are white and linear to branched in form. These pseudocyphellae are predominantly located along the margins of the lobes, distinguishing it from ''P. omphalodes'', where pseudocyphellae are found both marginally and laminally (on the flat upper surface). The
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, root-like structures on the underside of the thallus, are simple or forked and help anchor the lichen to its . Internally, the lichen has a white
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 ...
. The chemical composition of ''P. pinnatifida'' includes several characteristic substances. The major
lichen product Lichen products, also known as lichen substances, are organic compounds produced by a lichen. Specifically, they are secondary metabolites. Lichen products are represented in several different chemical classes, including terpenoids, orcinol deri ...
s are
atranorin Atranorin is a chemical substance produced by some species of lichen. It is a secondary metabolite belonging to a group of compounds known as depsides. Atranorin has analgesic, anti-inflammatory, antibacterial, antifungal, cytotoxic, antioxidant, ...
, salazinic acid, and consalazinic acid, while
protolichesterinic acid Protolichesterinic acid is a naturally occurring γ-lactone compound found in various lichen species. Its structure consists of a combination of a lactone ring with a carboxylic acid group and a long aliphatic side chain. First isolated in the ...
is present as a minor substance. Some populations have been reported to contain trace amounts of galbinic and
fumarprotocetraric acid Fumarprotocetraric acid is a chemical compound with the chemical formula . It is a secondary metabolite produced by a variety of lichens. Occurrence Fumarprotocetraric acid is mainly known for its occurrence in Iceland moss (''Cetraria islandica ...
, though these compounds are not consistently present across all specimens. These chemical compounds, particularly the combination and concentrations of different acids, help differentiate ''P. pinnatifida'' from other species in the ''P. omphalodes'' group.
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 ''P. pinnatifida'' reveals
fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an ...
s similar to those in '' Parmelia expallida'' and '' Parmelia grayana''.


Similar species

''Parmelia pinnatifida'' can be confused with several morphologically similar species within the genus. ''P. discordans'' appears superficially similar but can be distinguished by its chemical composition, particularly the presence of protocetraric and lobaric acids, which are absent in ''P. pinnatifida''. ''Parmelia omphalodes'' shares some morphological features but differs in several key characteristics. While both species have pseudocyphellae, in ''P. omphalodes'' these structures occur both on the margins and across the upper surface (laminally), whereas in ''P. pinnatifida'' they are primarily marginal. The lobe structure also differs significantly between the two species, with ''P. pinnatifida'' having much narrower, more extensively branched lobes that are distinctively imbricated (overlapping) in multiple layers. This creates a more complex, layered appearance compared to ''P. omphalodes''. Additionally, ''P. omphalodes'' lacks the distinctive marginal lobules characteristic of ''P. pinnatifida'' and contains lobaric acid as part of its chemical profile. Despite these morphological differences, recent phylogenetic analyses using ITS sequence data have been unable to clearly separate ''P. pinnatifida'' and ''P. omphalodes'' as distinct evolutionary lineages. This suggests that their taxonomic relationship may be more complex than previously thought and might require genome-wide data to fully resolve. The arctic species '' P. skultii'' presents another challenge for identification, as it shares similar morphological and chemical traits with ''P. pinnatifida''. However, ''P. skultii'' can be distinguished by its broader lobes, which can reach up to 8 mm in width, compared to the narrower lobes of ''P. pinnatifida''. Furthermore, ''P. skultii'' has a distinctive chemical signature, containing equal concentrations of norstictic and salazinic acids, a feature not found in ''P. pinnatifida''.


Habitat and distribution

''Parmelia pinnatifida'' has been recorded in various parts of Europe, including Sweden, Germany, and Switzerland. The species has also been documented in Austria, Finland, Norway, Romania, Slovakia, and Spain. Beyond Europe, it occurs in North America, with populations confirmed in Canada, Greenland, and the United States. While the species has been documented in Antarctica with molecular confirmation, its reported presence in southern South America and New Zealand requires additional studies to verify these distributions. The species was first discovered in Italy in 2019, where it was found in Val Brandet, a narrow secondary valley in north-western
Val Camonica Val Camonica or Valcamonica (), also Valle Camonica and anglicized as Camonica Valley, is one of the largest valleys of the central Alps, in eastern Lombardy, Italy. It extends about from the Tonale Pass to Corna Trentapassi, in the c ...
in the
Orobic Alps The Bergamasque Alps or Bergamo Alps (; sometimes translated into English as Orobic Alps) are a mountain range in the Italian Alps. They are located in northern Lombardy and named after the city Bergamo, south of the mountains. Within the Eastern ...
. Prior to this discovery, the species was only known from the French and Austrian
Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...
. The species shows a strong preference for siliceous rocks, making it primarily an
epilithic Lithophytes are plants that grow in or on rocks. They can be classified as either epilithic (or epipetric) or endolithic; epilithic lithophytes grow on the surfaces of rocks, while endolithic lithophytes grow in the crevices of rocks (and are als ...
lichen. While it predominantly colonises rock surfaces, specimens have been found growing on
bryophyte Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s and plant debris. In rare instances, it has been documented growing on tree bark, with one notable record on Norway spruce (''
Picea abies ''Picea abies'', the Norway spruce or European spruce, is a species of spruce native to Northern Europe, Northern, Central Europe, Central and Eastern Europe. It has branchlets that typically hang downwards, and the largest cones of any spruce, ...
''). In its Italian location, the species was found growing on a
schist Schist ( ) is a medium-grained metamorphic rock generally derived from fine-grained sedimentary rock, like shale. It shows pronounced ''schistosity'' (named for the rock). This means that the rock is composed of mineral grains easily seen with a l ...
boulder at the edge of a pathway in a shady, moist and cool
montane Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
coniferous forest near a stream, demonstrating its preference for humid microhabitats. In mountainous regions, ''P. pinnatifida'' can be particularly scarce. For example, in Poland, where it was first documented in 2016, the species is found exclusively in the
Carpathian Mountains 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 Ural Mountains, Urals at and the Scandinav ...
and the Góry Świętokrzyskie Mountains, where it maintains small, isolated populations.


References

{{Taxonbar, from=Q22106274 pinnatifida Lichen species Lichens described in 1976 Lichens of Europe Taxa named by Syo Kurokawa