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| status = G4 | | status = G4 | ||
| status_system = TNC | | status_system = TNC | ||
| status_ref = | | status_ref = <ref name=NS>{{cite NatureServe |id=2.123193 |title=''Parmotrema perlatum'' |access-date=24 October 2025}}</ref> | ||
| taxon = Parmotrema perlatum | | taxon = Parmotrema perlatum | ||
| authority = ([[William Hudson (botanist)|Huds.]]) [[M.Choisy]] (1952) | | authority = ([[William Hudson (botanist)|Huds.]]) [[M.Choisy]] (1952) | ||
| synonyms_ref = | | synonyms_ref = <ref name="Species Fungorum synonymy"/> | ||
| synonyms = {{Collapsible list|bullets=on | | synonyms = {{Collapsible list|bullets=on | ||
|''Lichen perlatus'' {{au|Huds. (1762)}} | |''Lichen perlatus'' {{au|Huds. (1762)}} | ||
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===Historical taxonomy=== | ===Historical taxonomy=== | ||
The [[taxonomy (biology)|taxonomy]] of ''Parmotrema perlatum'' has a rich history marked by periods of confusion and clarification that typify the dynamic nature of botanical classification. It was originally [[species description|described]] as ''Lichen perlatus'' by [[William Hudson (botanist)|William Hudson]] in his 1762 work ''Flora Anglica''. Hudson described it as a foliaceous (leafy) lichen with creeping, lobed, and smooth characteristics, having a pearly edge, a farinaceous ({{lichengloss|pruinose}}) texture, and a black underside, adorned with slightly scalloped, brown, stalked fruiting bodies. | The [[taxonomy (biology)|taxonomy]] of ''Parmotrema perlatum'' has a rich history marked by periods of confusion and clarification that typify the dynamic nature of botanical classification. It was originally [[species description|described]] as ''Lichen perlatus'' by [[William Hudson (botanist)|William Hudson]] in his 1762 work ''Flora Anglica''. Hudson described it as a foliaceous (leafy) lichen with creeping, lobed, and smooth characteristics, having a pearly edge, a farinaceous ({{lichengloss|pruinose}}) texture, and a black underside, adorned with slightly scalloped, brown, stalked fruiting bodies.<ref name="Hudson 1762"/> The [[taxon]] was later transferred to the genus ''[[Parmelia (fungus)|Parmelia]]'' by [[Erik Acharius]] in 1803, becoming ''Parmelia perlata''.<ref name="Acharius 1803"/> The name was well-established in scientific literature, being cited extensively in works like [[Alexander Zahlbruckner]]'s popular 1929 catalogue.<ref name="Hawksworth 2004"/> | ||
In 1952, [[Maurice Choisy]] reclassified it under the current name, ''Parmotrema perlatum''. | In 1952, [[Maurice Choisy]] reclassified it under the current name, ''Parmotrema perlatum''.<ref name="Choisy 1952"/> The nomenclature of ''Parmotrema perlatum'' was revisited in the late 20th century, amid a broader effort to clarify the [[type (biology)|typification]] and application of early lichen names. [[Mason Hale]], in 1961, undertook a detailed restudy of the species,<ref name="Hale 1962"/> selecting a [[lectotype]] from the Dillenian collections—the [[herbarium]] and associated works of [[Johann Jacob Dillenius]] housed at the [[University of Oxford]]. This solidified the application of Hudson's name and was part of a larger trend in lichenology to fix historical names to specific [[herbarium]] specimens to stabilise nomenclature.<ref name="Hawksworth 2004"/> | ||
The name ''Parmelia perlata'' was widely accepted until Hale and Ahti (1986) encountered the designation ''Lichen chinensis'', | The name ''Parmelia perlata'' was widely accepted until Hale and Ahti (1986) encountered the designation ''Lichen chinensis'',<ref name="Hale & Ahti 1986"/> introduced by [[Pehr Osbeck]] in 1757.<ref name="Osbeck 1757"/> They proposed the name ''Parmotrema chinense'', based on the assumption that Osbeck's specimen corresponded to the well-known species ''Parmotrema perlatum''. However, this proposal was not universally adopted due to the lack of valid typification and the name's absence in the literature between 1757 and 1986.<ref name="Hawksworth 2004"/> | ||
====Impact of the Tokyo Code (1993)==== | ====Impact of the Tokyo Code (1993)==== | ||
The Tokyo Code of 1993 extended the provisions for [[conserved name|conserving names]] to all species, not just those of major economic importance. This change in the [[International Code of Botanical Nomenclature]] allowed for the conservation of names that would promote nomenclatural stability. Despite this provision, no formal proposal was made to conserve the name ''Parmotrema chinense'', and thus it did not gain widespread acceptance. | The Tokyo Code of 1993 extended the provisions for [[conserved name|conserving names]] to all species, not just those of major economic importance. This change in the [[International Code of Botanical Nomenclature]] allowed for the conservation of names that would promote nomenclatural stability. Despite this provision, no formal proposal was made to conserve the name ''Parmotrema chinense'', and thus it did not gain widespread acceptance.<ref name="Hawksworth 2004"/> | ||
[[David Leslie Hawksworth|David Hawksworth's]] 2004 study brought significant clarity to the taxonomic confusion. He rediscovered Osbeck's original material in Linnaeus' herbarium and identified it as belonging to ''[[Parmotrema tinctorum]]'', not ''Parmotrema perlatum''. Hawksworth demonstrated that ''Lichen chinensis'' was not [[validly published name|validly published]] because it lacked a proper description and was linked with an expression of doubt by Osbeck. Hawksworth's work led to the reinstatement of the name ''Parmotrema perlatum'', confirming that Hudson's name was legitimate and should continue to be used. This resolution was based on the original typification by Hale and the invalid publication status of ''Lichen chinensis''. | [[David Leslie Hawksworth|David Hawksworth's]] 2004 study brought significant clarity to the taxonomic confusion. He rediscovered Osbeck's original material in Linnaeus' herbarium and identified it as belonging to ''[[Parmotrema tinctorum]]'', not ''Parmotrema perlatum''. Hawksworth demonstrated that ''Lichen chinensis'' was not [[validly published name|validly published]] because it lacked a proper description and was linked with an expression of doubt by Osbeck. Hawksworth's work led to the reinstatement of the name ''Parmotrema perlatum'', confirming that Hudson's name was legitimate and should continue to be used. This resolution was based on the original typification by Hale and the invalid publication status of ''Lichen chinensis''.<ref name="Hawksworth 2004"/> | ||
Recent studies suggest that the circumscription of ''Parmotrema perlatum'' may need to be revised. Research utilising [[DNA sequencing]] has uncovered [[cryptic species|cryptic]] diversity within the genus ''Parmotrema'', indicating that traditional phenotype-based identification methods may underestimate species diversity. Specifically, the genetic analysis of ''P.&nbsp;perlatum'' and related species revealed multiple distinct [[lineage (evolution)|lineages]] that were previously grouped under a single nominal taxon. These findings highlight the need for a comprehensive taxonomic re-evaluation of ''P.&nbsp;perlatum'' to accurately delineate species boundaries and account for hidden genetic diversity. | Recent studies suggest that the circumscription of ''Parmotrema perlatum'' may need to be revised. Research utilising [[DNA sequencing]] has uncovered [[cryptic species|cryptic]] diversity within the genus ''Parmotrema'', indicating that traditional phenotype-based identification methods may underestimate species diversity. Specifically, the genetic analysis of ''P.&nbsp;perlatum'' and related species revealed multiple distinct [[lineage (evolution)|lineages]] that were previously grouped under a single nominal taxon. These findings highlight the need for a comprehensive taxonomic re-evaluation of ''P.&nbsp;perlatum'' to accurately delineate species boundaries and account for hidden genetic diversity.<ref name="Del-Prado et al. 2019"/> | ||
===Phylogeny=== | ===Phylogeny=== | ||
In [[molecular phylogenetics]] analysis, ''Parmotrema perlatum'' has a [[sister group|sister]] relationship with ''[[Parmotrema crinitum]]''. These two species form a [[clade]] that itself is sister to a clade with ''[[Parmotrema austrosinense|P.&nbsp;austrosinense]]'' and ''[[Parmotrema tinctorum|P.&nbsp;tinctorum]]''. | In [[molecular phylogenetics]] analysis, ''Parmotrema perlatum'' has a [[sister group|sister]] relationship with ''[[Parmotrema crinitum]]''. These two species form a [[clade]] that itself is sister to a clade with ''[[Parmotrema austrosinense|P.&nbsp;austrosinense]]'' and ''[[Parmotrema tinctorum|P.&nbsp;tinctorum]]''.<ref name="Crespo et al. 2010"/> In a comprehensive phylogenetic analysis by Stelate and colleagues (2022), ''P.&nbsp;perlatum'' and ''P&nbsp;crinitum'' were found to form a well-supported [[monophyletic]] group using [[internal transcribed spacer]] sequences and several analytical methods. This study highlights the complexity of species boundaries within the genus and the need for further research incorporating additional molecular markers to confirm these findings.<ref name="Stelate et al. 2022"/> | ||
===Common names=== | ===Common names=== | ||
[[Common name|Vernacular]] names used for this species include black stone flower, stone lichen, sea lichen, kalpasi, kalpas, kalpashi, and kalpash. The latter name and its variations, however, have been used as a crude drug in Indian medicines for more than one species, including ''Parmotrema perlatum'', ''Parmotrema tinctorum'', and ''[[Everniastrum cirrhatum]]''. | [[Common name|Vernacular]] names used for this species include black stone flower, stone lichen, sea lichen, kalpasi, kalpas, kalpashi, and kalpash. The latter name and its variations, however, have been used as a crude drug in Indian medicines for more than one species, including ''Parmotrema perlatum'', ''Parmotrema tinctorum'', and ''[[Everniastrum cirrhatum]]''.<ref name="Ravindran 2017"/> In North America, vernacular names used for the species include "powdered ruffle lichen",<ref name="Brodo et al. 2001"/><ref name="McMullin 2023"/> "powdered scatter-rug",<ref name="Goward 1994"/> and "queen ruffle".<ref name="gov.bc.ca"/> The [[botanical name|species epithet]] ''perlatum'' refers to the [[pearl]]-like margins of the lobes, which are directly referenced in Hudson's original 1762 description of the species. He proposed the English name "pearl lichen";<ref name="Hudson 1762"/> this name later morphed into "pearly parmelia" in some 19th-century British accounts of lichen flora.<ref name="Mosley 1863"/><ref name="Smith 1844"/> | ||
==Description== | ==Description== | ||
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| caption2 = ... and marginal cilia. | | caption2 = ... and marginal cilia. | ||
}} | }} | ||
''Parmotrema perlatum'' has a [[thallus]] that ranges from loosely to tightly attached to the surface it grows on, forming expansive, spreading colonies that often merge together. | ''Parmotrema perlatum'' has a [[thallus]] that ranges from loosely to tightly attached to the surface it grows on, forming expansive, spreading colonies that often merge together.<ref name="Kantvilas 2023"/> Individual thalli typically measure up to {{cvt|10|–|15|cm|0}} in diameter. The upper thallus surface is greenish-grey, blue-grey,<ref name="McMullin 2023"/> or yellowish-grey in colour,<ref name="Brodo et al. 2001"/> lacking {{lichengloss|pruina}} and either free of spots ({{lichengloss|maculae}}),<ref name="McMullin 2023"/> or with few maculae.<ref name="Brodo et al. 2001"/> This species develops soredia, a type of asexual reproductive structure, aiding in its propagation. The {{lichengloss|lobes}} of this lichen vary from 1.5 to 10&nbsp;mm in width, with a wave-like ({{lichengloss|undulate}}) or ruffled pattern and overlapping ({{lichengloss|imbricate}}) arrangement. The tips and edges of these lobes are generally smooth and round, sometimes notched ({{lichengloss|crenate}}) or incised, often curling up or inward, revealing the paler brown to black underside adorned with hair-like structures ({{lichengloss|cilia}}) up to 2.5&nbsp;mm in length.<ref name="Kantvilas 2023"/> [[Rhizine]]s are common on the underside of the thallus, except for a brown border near the edges.<ref name="McMullin 2023"/> | ||
The [[soredia]] found in this species are {{lichengloss|granular}} and appear white or may become grey due to wear. They are located within specifically structured groups called soralia, which can be linear to oval in shape, often positioned at the edges of the lobes. | The [[soredia]] found in this species are {{lichengloss|granular}} and appear white or may become grey due to wear. They are located within specifically structured groups called soralia, which can be linear to oval in shape, often positioned at the edges of the lobes.<ref name="Kantvilas 2023"/> The presence of soredia causes the lobe margins to curl back and form {{lichengloss|capitate}} soralia.<ref name="McMullin 2023"/> The upper surface of the lichen is typically whitish grey to pale greenish-grey, and can be either smooth or slightly wrinkled, without spots ({{lichengloss|immaculate}}), featuring scattered, shallow cracks.<ref name="Kantvilas 2023"/> [[Isidia]] are absent in this species.<ref name="McMullin 2023"/> | ||
[[Apothecia]] (fruiting bodies) are rare in ''Parmotrema perlatum''. When present, they measure 4–8&nbsp;mm across and are somewhat stalked and funnel-shaped with a brown, concave {{lichengloss|disc}}. The edges of these structures curl inward, becoming thick with soredia as they mature. Its spores are [[ellipsoid]] in shape and typically measure between 20 and 28&nbsp;[[μm]] in length and 11 to 17&nbsp;μm in width, with a wall thickness of 2–3&nbsp;μm. [[Pycnidia]], which are structures that produce asexual spores called [[conidia]], appear sporadically on the surface ({{lichengloss|laminal}}) of the thallus, with the conidia being thread-like and straight, measuring 6–8 by 1&nbsp;μm. | [[Apothecia]] (fruiting bodies) are rare in ''Parmotrema perlatum''. When present, they measure 4–8&nbsp;mm across and are somewhat stalked and funnel-shaped with a brown, concave {{lichengloss|disc}}. The edges of these structures curl inward, becoming thick with soredia as they mature. Its spores are [[ellipsoid]] in shape and typically measure between 20 and 28&nbsp;[[μm]] in length and 11 to 17&nbsp;μm in width, with a wall thickness of 2–3&nbsp;μm. [[Pycnidia]], which are structures that produce asexual spores called [[conidia]], appear sporadically on the surface ({{lichengloss|laminal}}) of the thallus, with the conidia being thread-like and straight, measuring 6–8 by 1&nbsp;μm.<ref name="Kantvilas 2023"/> | ||
===Photobiont=== | ===Photobiont=== | ||
The photobiont partner of ''Parmotrema perlatum'' is from ''[[Trebouxia]]'', a [[green alga]]l genus belonging to the order [[Trebouxiales]] (order [[Chlorophyta]]). | The photobiont partner of ''Parmotrema perlatum'' is from ''[[Trebouxia]]'', a [[green alga]]l genus belonging to the order [[Trebouxiales]] (order [[Chlorophyta]]).<ref name="Carniel 2015"/> It has been identified as an undescribed species within a [[clade]] containing ''[[Trebouxia arboricola]]''.<ref name="Bertuzzi et al. 2017"/> | ||
A study compared the desiccation tolerance and physiological responses of lichenised ''Trebouxia'' to isolated cultures of the same alga. Both forms can survive extended [[desiccation]], but with differing responses to photo-oxidative stress. Lichenisation enhances the [[photoprotection|photoprotective]] mechanisms of ''Trebouxia'', improving [[Quenching (fluorescence)|quenching]] of excess light energy, particularly under high [[relative humidity]], and controlling [[reactive oxygen species]] production under light exposure. However, isolated cultures showed better photosynthetic performance after desiccation recovery. This research demonstrates the mutual benefits of the lichen-photobiont partnership, where the alga gains a sheltered environment boosting its resilience to environmental stressors. | A study compared the desiccation tolerance and physiological responses of lichenised ''Trebouxia'' to isolated cultures of the same alga. Both forms can survive extended [[desiccation]], but with differing responses to photo-oxidative stress. Lichenisation enhances the [[photoprotection|photoprotective]] mechanisms of ''Trebouxia'', improving [[Quenching (fluorescence)|quenching]] of excess light energy, particularly under high [[relative humidity]], and controlling [[reactive oxygen species]] production under light exposure. However, isolated cultures showed better photosynthetic performance after desiccation recovery. This research demonstrates the mutual benefits of the lichen-photobiont partnership, where the alga gains a sheltered environment boosting its resilience to environmental stressors.<ref name="Carniel 2015"/> | ||
Further studies on ''Parmotrema perlatum'' revealed specific [[antioxidant]] mechanisms supporting its photobiont under stress. The lichen shows high levels of [[reactive oxygen species]] scavenging enzymes such as [[superoxide dismutase]] and [[ascorbate peroxidase]], protecting the photobiont from [[oxidative damage]] during dehydration and rehydration cycles. This enhanced antioxidant system provides not only physical shelter but also biochemical protection, increasing the photobiont's resilience to environmental fluctuations. | Further studies on ''Parmotrema perlatum'' revealed specific [[antioxidant]] mechanisms supporting its photobiont under stress. The lichen shows high levels of [[reactive oxygen species]] scavenging enzymes such as [[superoxide dismutase]] and [[ascorbate peroxidase]], protecting the photobiont from [[oxidative damage]] during dehydration and rehydration cycles. This enhanced antioxidant system provides not only physical shelter but also biochemical protection, increasing the photobiont's resilience to environmental fluctuations.<ref name="Bertuzzi et al. 2017"/> | ||
==Chemistry== | ==Chemistry== | ||
Chemically, ''Parmotrema perlatum'' contains [[atranorin]] and [[chloroatranorin]], alongside a predominant stictic acid {{lichengloss|chemosyndrome}} that includes [[stictic acid|stictic]] as a major [[secondary metabolite]] and smaller amounts of [[constictic acid]] and other related substances. Testing the [[medulla (lichenology)|medulla]] (the inner layer beneath the upper cortex) with [[spot test (lichen)|spot tests]] results in K+ (yellow), KC–, and P+ (orange) reactions. | Chemically, ''Parmotrema perlatum'' contains [[atranorin]] and [[chloroatranorin]], alongside a predominant stictic acid {{lichengloss|chemosyndrome}} that includes [[stictic acid|stictic]] as a major [[secondary metabolite]] and smaller amounts of [[constictic acid]] and other related substances. Testing the [[medulla (lichenology)|medulla]] (the inner layer beneath the upper cortex) with [[spot test (lichen)|spot tests]] results in K+ (yellow), KC–, and P+ (orange) reactions.<ref name="Kantvilas 2023"/> The cortical layer, in contrast, is K+ (yellow), KC–, and P–.<ref name="McMullin 2023"/> | ||
The secondary metabolites of ''Parmotrema perlatum'' have been studied using [[gas chromatography–mass spectrometry]] (GC–MS) and [[liquid chromatography–mass spectrometry]] (LC–MS/MS). The lichen produces several notable compounds, including [[orcinol]], [[atraric acid]], [[benzoic acid]], 2,4-dihydroxy-3,6-dimethyl-, methyl ester, and [[palmitic acid]], methyl ester. GC–MS analysis revealed the presence of orcinol (63%) and atraric acid (21%) in the methanol extract, while benzoic acid was predominant in the chloroform extract. The [[hexane]] extract contained significant amounts of benzoic acid, 2,4-dihydroxy-3,6-dimethyl-, methyl ester (62%). | The secondary metabolites of ''Parmotrema perlatum'' have been studied using [[gas chromatography–mass spectrometry]] (GC–MS) and [[liquid chromatography–mass spectrometry]] (LC–MS/MS). The lichen produces several notable compounds, including [[orcinol]], [[atraric acid]], [[benzoic acid]], 2,4-dihydroxy-3,6-dimethyl-, methyl ester, and [[palmitic acid]], methyl ester. GC–MS analysis revealed the presence of orcinol (63%) and atraric acid (21%) in the methanol extract, while benzoic acid was predominant in the chloroform extract. The [[hexane]] extract contained significant amounts of benzoic acid, 2,4-dihydroxy-3,6-dimethyl-, methyl ester (62%).<ref name="Dwarakanath et al. 2022"/> A more recent study using liquid chromatography-[[electrospray ionization]]-mass spectrometry/mass spectrometry as an analysis technique tentatively identified a total of twenty-five lichen products, including 5 [[depside]]s, 12 [[depsidone]]s, 2 [[diphenyl ether]]s, 1 [[aromatic]] considered as possible artifact, 1 [[dibenzofuran]], 1 [[carbohydrate]], 1 [[organic acid]], and 2 undefined compounds.<ref name="Castañeta et al. 2024"/> | ||
==Similar species== | ==Similar species== | ||
The distinguishing features of ''Parmotrema perlatum'', such as the presence of soredia and stictic acid, facilitate its easy identification. In mature specimens, the appearance of scattered, fine cracks on the upper surface may resemble the cracked maculae seen in ''[[Parmotrema reticulatum|P.&nbsp;reticulatum]]'', which shares similar habitats. However, the two species can be differentiated chemically, as ''P.&nbsp;reticulatum'' contains [[salazinic acid]], unlike ''P.&nbsp;perlatum''. | The distinguishing features of ''Parmotrema perlatum'', such as the presence of soredia and stictic acid, facilitate its easy identification. In mature specimens, the appearance of scattered, fine cracks on the upper surface may resemble the cracked maculae seen in ''[[Parmotrema reticulatum|P.&nbsp;reticulatum]]'', which shares similar habitats. However, the two species can be differentiated chemically, as ''P.&nbsp;reticulatum'' contains [[salazinic acid]], unlike ''P.&nbsp;perlatum''.<ref name="Kantvilas 2023"/> | ||
''Parmotrema perlatum'' and ''[[Parmotrema stuppeum]]'' are two morphologically similar species that can be found in similar habitats. Both species have a loosely attached thallus with revolute, wavy lobes and sparsely ciliate lobe tips. Their upper cortex is continuous and not finely reticulately cracked, while the lower surface is black and rhizinate. Both species also feature linear soralia. However, there are several key differences that can help distinguish between the two. While earlier descriptions suggested that ''P.&nbsp;stuppeum'' has a matte, olive-green to brownish-green upper surface and ''P.&nbsp;perlatum'' has a slightly shiny, whitish-grey to greyish-green upper surface, recent observations have shown that both species have a distinctly matte upper surface with similar colouration. | ''Parmotrema perlatum'' and ''[[Parmotrema stuppeum]]'' are two morphologically similar species that can be found in similar habitats. Both species have a loosely attached thallus with revolute, wavy lobes and sparsely ciliate lobe tips. Their upper cortex is continuous and not finely reticulately cracked, while the lower surface is black and rhizinate. Both species also feature linear soralia. However, there are several key differences that can help distinguish between the two. While earlier descriptions suggested that ''P.&nbsp;stuppeum'' has a matte, olive-green to brownish-green upper surface and ''P.&nbsp;perlatum'' has a slightly shiny, whitish-grey to greyish-green upper surface, recent observations have shown that both species have a distinctly matte upper surface with similar colouration.<ref name="Tsurykau et al. 2015"/> | ||
The most reliable morphological difference in the field is the location of the soralia: ''P.&nbsp;stuppeum'' has strictly terminal soralia, whereas ''P.&nbsp;perlatum'' has submarginal soralia. Additionally, the two species can be distinguished by their chemical composition. ''P.&nbsp;stuppeum'' contains salazinic acid, while ''P.&nbsp;perlatum'' has a stictic acid complex. Although both acids cause a Pd+ orange to orange-red medulla reaction, a potassium (K) spot test can separate the species: the medulla of ''P.&nbsp;perlatum'' turns yellow (K+ yellow), whereas in ''P.&nbsp;stuppeum'', the yellow colour turns red (K+ yellow turning red). | The most reliable morphological difference in the field is the location of the soralia: ''P.&nbsp;stuppeum'' has strictly terminal soralia, whereas ''P.&nbsp;perlatum'' has submarginal soralia. Additionally, the two species can be distinguished by their chemical composition. ''P.&nbsp;stuppeum'' contains salazinic acid, while ''P.&nbsp;perlatum'' has a stictic acid complex. Although both acids cause a Pd+ orange to orange-red medulla reaction, a potassium (K) spot test can separate the species: the medulla of ''P.&nbsp;perlatum'' turns yellow (K+ yellow), whereas in ''P.&nbsp;stuppeum'', the yellow colour turns red (K+ yellow turning red).<ref name="Tsurykau et al. 2015"/> | ||
''Parmotrema perlatum'' can be distinguished from other sorediate and marginally ciliate species like ''[[Parmotrema arnoldii|P.&nbsp;arnoldii]]'' and ''[[Parmotrema robustum|P.&nbsp;robustum]]'' by the presence of the stictic acid chemosyndrome. | ''Parmotrema perlatum'' can be distinguished from other sorediate and marginally ciliate species like ''[[Parmotrema arnoldii|P.&nbsp;arnoldii]]'' and ''[[Parmotrema robustum|P.&nbsp;robustum]]'' by the presence of the stictic acid chemosyndrome.<ref name="Jabłońska et al. 2009"/> Also, the medulla of ''P.&nbsp;arnoldii'' fluoresces strongly when lit with an [[ultraviolet]] lamp.<ref name="McCune & Geiser 2009"/> ''Parmotrema perlatum'' is similar to ''[[Parmotrema crinitum|P.&nbsp;crinitum]]'' due to both species having a brown to tan, erhizinate marginal zone and the presence of the stictic acid chemosyndrome in the medulla. However, ''P.&nbsp;crinitum'' can be distinguished by its [[isidia]]te upper surface.<ref name="Jayalal et al. 2013"/> Another potential lookalike, ''[[Parmotrema margaritatum|P.&nbsp;margaritatum]]'', is distinguished from ''P.&nbsp;perlatum'' by the K+ (red) reaction of its medulla.<ref name="McMullin 2023"/> ''[[Cetrelia cetrarioides]]'' has been documented as a lookalike, presumably because of the cilia on its thallus margin, and the presence of atranorin and the stictic acid chemosyndrome.<ref name="Kukwa et al. 2011"/> | ||
{| class="center toccolours" | {| class="center toccolours" | ||
|+ '''Lookalikes''' | |+ '''Lookalikes''' | ||
| | |<gallery mode="packed" heights="140" style="line-height:130%"> | ||
File:Parmotrema arnoldii 236323.jpg |alt= | ''[[Parmotrema arnoldii]]'' | File:Parmotrema arnoldii 236323.jpg |alt= | ''[[Parmotrema arnoldii]]'' | ||
File:Parmotream crinitum - Flickr - pellaea.jpg |alt= | ''[[Parmotrema crinitum]]'' | File:Parmotream crinitum - Flickr - pellaea.jpg |alt= | ''[[Parmotrema crinitum]]'' | ||
| Line 119: | Line 119: | ||
File:Parmotrema reticulatum - Flickr - pellaea.jpg |alt= | ''[[Parmotrema reticulatum]]'' | File:Parmotrema reticulatum - Flickr - pellaea.jpg |alt= | ''[[Parmotrema reticulatum]]'' | ||
File:Parmotrema robustum group 202849.jpg |alt= | ''[[Parmotrema robustum]]'' | File:Parmotrema robustum group 202849.jpg |alt= | ''[[Parmotrema robustum]]'' | ||
</gallery> | |||
|- | |- | ||
|} | |} | ||
| Line 125: | Line 125: | ||
==Habitat and distribution== | ==Habitat and distribution== | ||
[[File:Powdered Ruffle Lichen (982620239).jpg|thumb|right|''Parmotrema perlatum'' in Northwestern California growing on a hardwood limb]] | [[File:Powdered Ruffle Lichen (982620239).jpg|thumb|right|''Parmotrema perlatum'' in Northwestern California growing on a hardwood limb]] | ||
''Parmotrema perlatum'' typically grows in areas with ample light, favouring neutral to slightly acidic-barked broad-leaved trees. It is commonly found on [[siliceous rock]]s and walls, as well as mossy coastal rocks, generally growing in places with moderate to strong sunlight. | ''Parmotrema perlatum'' typically grows in areas with ample light, favouring neutral to slightly acidic-barked broad-leaved trees. It is commonly found on [[siliceous rock]]s and walls, as well as mossy coastal rocks, generally growing in places with moderate to strong sunlight.<ref name="Cannon et al. 2023"/> In the [[Great Smoky Mountains National Park]] in the United States, ''Parmotrema perlatum'' is especially abundant on branches in humid, high-elevation habitats.<ref name="Tripp & Lendemer 2020"/> Similarly, in East Africa, it grows in the misty environments of [[inselberg]]s, [[montane]] forests, and ''[[Erica (plant)|Erica]]''-dominated habitats, typically found between {{cvt|1400|and|3100|m}} above sea level.<ref name="Swinscow & Krog 1988"/> | ||
The species is globally distributed, found in both [[temperate]] and [[tropical regions]]. It has been reported across numerous European countries including Austria, Belgium, the Czech Republic, France, Germany, Great Britain, Ireland, Italy, Luxembourg, the Netherlands, Portugal, Scandinavia, Slovakia, Spain, and Ukraine. | The species is globally distributed, found in both [[temperate]] and [[tropical regions]]. It has been reported across numerous European countries including Austria, Belgium, the Czech Republic, France, Germany, Great Britain, Ireland, Italy, Luxembourg, the Netherlands, Portugal, Scandinavia, Slovakia, Spain, and Ukraine.<ref name="Jabłońska et al. 2009"/> Although it is rare in Eastern Europe, it is widely distributed in both the Asian and European parts of Russia.<ref name="Tsurykau et al. 2015"/> Beyond Europe, it is also present in Macaronesia, Africa, Australia, North America, and South America.<ref name="Jabłońska et al. 2009"/> Its Asian distribution includes India, Japan, Taiwan, and South Korea.<ref name="Jayalal et al. 2013"/> Although it has historically been recorded in Nepal and Sri Lanka, these reports are considered tentative due to shifting species concepts and possible confusion with the lookalike ''[[Parmotrema pseudonilgherrense]]''.<ref name="Awasthi 2007"/> | ||
''Parmotrema perlatum'' is globally widespread lichen found on all continents except Antarctica and predominantly in oceanic areas in Europe, primarily grows on bark and occasionally on siliceous rocks amongst mosses. While it is seeing an increase in the Netherlands due to global warming, it is [[critically endangered]] in the Czech Republic, | ''Parmotrema perlatum'' is globally widespread lichen found on all continents except Antarctica and predominantly in oceanic areas in Europe, primarily grows on bark and occasionally on siliceous rocks amongst mosses. While it is seeing an increase in the Netherlands due to global warming, it is [[critically endangered]] in the Czech Republic,<ref name="Liška et al. 2008"/> Slovakia, and Poland<ref name="Cieśliński et al. 2003"/> due to susceptibility to air pollution, and is listed as extinct in certain regional Red Data Books due to a lack of recent findings.<ref name="Vondrák & Liška 2010"/> In contrast, it has been increasing in sightings in the Netherlands, a phenomenon attributed to both global warming decreases in the levels or air pollution in recent decades.<ref name="van Herk et al. 2002"/><ref name="Aptroot & van Herk 2007"/> Its recent recurrence in Hungary, particularly on some unusual hosts (''[[Catalpa bignonioides]]'', ''[[Prunus serotina]]'', and ''[[Robinia pseudoacacia]]'') have been suggested as a possible consequence of "a recolonisation process, due to the improving air quality".<ref name="Balogh et al. 2021"/> | ||
==Ecology== | ==Ecology== | ||
''Parmotrema perlatum'' is an important species within specific lichen [[community (biology)|communities]] in British woodlands, particularly those in late successional [[Mesotrophic soils|mesotrophic]] settings in oceanic or humid microclimates. It is associated with the Type K Lobaria pulmonaria-Isothecium myosuroides ecological Community. This community type is characterised by its occurrence in mature mesotrophic environments, which are often warmer in winter climates or specific microhabitats. This community includes, in addition to ''P.&nbsp;perlatum'', dominant foliose lichens like ''[[Lobaria pulmonaria]]'', ''[[Hypotrachyna taylorensis]]'', and ''[[Parmotrema crinitum]]'', as well as [[bryophyte]]s such as ''[[Isothecium myosuroides]]''. | ''Parmotrema perlatum'' is an important species within specific lichen [[community (biology)|communities]] in British woodlands, particularly those in late successional [[Mesotrophic soils|mesotrophic]] settings in oceanic or humid microclimates. It is associated with the Type K Lobaria pulmonaria-Isothecium myosuroides ecological Community. This community type is characterised by its occurrence in mature mesotrophic environments, which are often warmer in winter climates or specific microhabitats. This community includes, in addition to ''P.&nbsp;perlatum'', dominant foliose lichens like ''[[Lobaria pulmonaria]]'', ''[[Hypotrachyna taylorensis]]'', and ''[[Parmotrema crinitum]]'', as well as [[bryophyte]]s such as ''[[Isothecium myosuroides]]''.<ref name="Ellis et al. 2015"/> | ||
A 2017 study investigated the physiological responses of ''Parmotrema perlatum'' along an [[aridity]] gradient in Southern Portugal. The researchers transplanted thalli of ''P.&nbsp;perlatum'' to rural and forested sites characterised by varying levels of aridity and measured several physiological parameters, including photosynthetic performance, pigment content, [[ergosterol]] content, and sample viability, both before and after a six-month exposure period. The study found that ''P.&nbsp;perlatum'' showed lower photosynthetic performance (measured as [[Plant stress measurement#FV/FM|FV/FM]] and the performance index on an absorption basis, PIABS) in drier sites compared to more humid sites. In humid environments, the content of photosynthetic pigments increased post-exposure, while in drier sites, this increase was less pronounced. Additionally, ergosterol content was lower in drier sites, indicating a [[stress response]] to arid conditions. These results highlight that ''P.&nbsp;perlatum''{{'}}s physiological responses are significantly influenced by water availability. The ability to maintain higher photosynthetic performance and pigment content in humid conditions suggests that ''P.&nbsp;perlatum'' is better adapted to environments with higher moisture levels. This adaptability makes ''P.&nbsp;perlatum'' useful as a bioindicator for monitoring ecological responses to [[climate change]] and varying moisture conditions in Mediterranean ecosystems. | A 2017 study investigated the physiological responses of ''Parmotrema perlatum'' along an [[aridity]] gradient in Southern Portugal. The researchers transplanted thalli of ''P.&nbsp;perlatum'' to rural and forested sites characterised by varying levels of aridity and measured several physiological parameters, including photosynthetic performance, pigment content, [[ergosterol]] content, and sample viability, both before and after a six-month exposure period. The study found that ''P.&nbsp;perlatum'' showed lower photosynthetic performance (measured as [[Plant stress measurement#FV/FM|FV/FM]] and the performance index on an absorption basis, PIABS) in drier sites compared to more humid sites. In humid environments, the content of photosynthetic pigments increased post-exposure, while in drier sites, this increase was less pronounced. Additionally, ergosterol content was lower in drier sites, indicating a [[stress response]] to arid conditions. These results highlight that ''P.&nbsp;perlatum''{{'}}s physiological responses are significantly influenced by water availability. The ability to maintain higher photosynthetic performance and pigment content in humid conditions suggests that ''P.&nbsp;perlatum'' is better adapted to environments with higher moisture levels. This adaptability makes ''P.&nbsp;perlatum'' useful as a bioindicator for monitoring ecological responses to [[climate change]] and varying moisture conditions in Mediterranean ecosystems.<ref name="Paoli et al. 2017"/> | ||
[[Lichenicolous fungus|Lichenicolous]] (lichen-dwelling) fungi that have been recorded [[parasitism|parasitising]] ''Parmotrema perlatum'' include ''[[Abrothallus parmotrematis]]'', ''[[Briancoppinsia cytospora]]'', ''[[Lichenoconium erodens]]'', and ''[[Spirographa lichenicola]]''. | [[Lichenicolous fungus|Lichenicolous]] (lichen-dwelling) fungi that have been recorded [[parasitism|parasitising]] ''Parmotrema perlatum'' include ''[[Abrothallus parmotrematis]]'', ''[[Briancoppinsia cytospora]]'', ''[[Lichenoconium erodens]]'', and ''[[Spirographa lichenicola]]''.<ref name="BLS"/> | ||
==Conservation== | ==Conservation== | ||
''Parmotrema perlatum'' has been identified as a species of concern in some regions due to its rarity and declining populations. In Hungary, it has been proposed for '[[endangered species|endangered]]' status in the Hungarian lichen [[Regional Red List|red list]], reflecting its limited distribution and the pressures it faces in its natural habitats there. | ''Parmotrema perlatum'' has been identified as a species of concern in some regions due to its rarity and declining populations. In Hungary, it has been proposed for '[[endangered species|endangered]]' status in the Hungarian lichen [[Regional Red List|red list]], reflecting its limited distribution and the pressures it faces in its natural habitats there.<ref name="Balogh et al. 2021"/> Similarly, in Ukraine, the species is listed in the Red Data Book of Ukraine with the status of "Rare".<ref name="Didukh 2009"/> Additionally, ''Parmotrema perlatum'' is red-listed in Sweden<ref name="Thell et al. 2011"/> In northern North America, its [[NatureServe conservation status]] is designated as "G4", meaning "apparently secure" at the global level. In the United States, it has been assessed as secure in [[Kentucky]] and presumed [[extirpated]] in [[Wisconsin]], while in Canada, it is considered as vulnerable in [[British Columbia]] and [[Ontario]], and critically imperiled in [[New Brunswick]].<ref name="NatureServe"/> | ||
==Uses== | ==Uses== | ||
===As a spice=== | ===As a spice=== | ||
[[File:Parmotrema perlatum.jpg|thumb|right|''Parmotrema perlatum'' as the dried spice, kalpaasi]] | [[File:Parmotrema perlatum.jpg|thumb|right|''Parmotrema perlatum'' as the dried spice, kalpaasi]] | ||
''Parmotrema perlatum'' is used as a spice, particularly in the [[Tamil cuisine#Cuisine|cuisine]] of [[Tamil Nadu]]. It is especially prevalent in [[Chettinad cuisine]], being used in the popular rice dish [[biryani]], and also in many meat and vegetarian dishes. | ''Parmotrema perlatum'' is used as a spice, particularly in the [[Tamil cuisine#Cuisine|cuisine]] of [[Tamil Nadu]]. It is especially prevalent in [[Chettinad cuisine]], being used in the popular rice dish [[biryani]], and also in many meat and vegetarian dishes.<ref name="Ravindran 2017"/> | ||
In its raw state, black stone flower does not have much taste or fragrance. However, when put in contact with heat, especially hot [[cooking oil]] or [[ghee]], it releases a distinctive earthy, smoky flavour and aroma. This property of black stone flower is especially valued in the [[Tempering (spices)|tempering]] step of cooking a number of Indian dishes. | In its raw state, black stone flower does not have much taste or fragrance. However, when put in contact with heat, especially hot [[cooking oil]] or [[ghee]], it releases a distinctive earthy, smoky flavour and aroma. This property of black stone flower is especially valued in the [[Tempering (spices)|tempering]] step of cooking a number of Indian dishes.<ref name="Ravindran 2017"/><ref name="Mani 2021"/> | ||
The spice is also integral to various regional [[Masala (spice)|masalas]] throughout the Indian subcontinent. ''Parmotrema perlatum'' is a key ingredient in masalas such as Kala and Goda masala of [[Maharashtra]], Anglo-Indian bottle masala, bhojwar masala from [[Hyderabad]], and potli masala in [[Lucknow]]. It is what many cooks and commercial [[spice blend]] makers believe sets apart accomplished dishes from those made by amateurs. Despite its lack of a specific aroma or describable flavour in its raw form, its contribution to the complex flavour profile of these spice blends is highly valued. | The spice is also integral to various regional [[Masala (spice)|masalas]] throughout the Indian subcontinent. ''Parmotrema perlatum'' is a key ingredient in masalas such as Kala and Goda masala of [[Maharashtra]], Anglo-Indian bottle masala, bhojwar masala from [[Hyderabad]], and potli masala in [[Lucknow]]. It is what many cooks and commercial [[spice blend]] makers believe sets apart accomplished dishes from those made by amateurs. Despite its lack of a specific aroma or describable flavour in its raw form, its contribution to the complex flavour profile of these spice blends is highly valued.<ref name="Mani 2021"/> | ||
===Dyeing=== | ===Dyeing=== | ||
A natural purple dye extracted from ''Parmotrema perlatum'' using [[ammonia fermentation]] showed optimal results, with a notable dye yield and effective application on silk fabric. The study demonstrated the dye's potential as a sustainable alternative to [[synthetic dye]]s, with satisfactory [[colour fastness]] and fabric strength enhancement. | A natural purple dye extracted from ''Parmotrema perlatum'' using [[ammonia fermentation]] showed optimal results, with a notable dye yield and effective application on silk fabric. The study demonstrated the dye's potential as a sustainable alternative to [[synthetic dye]]s, with satisfactory [[colour fastness]] and fabric strength enhancement.<ref name="Roychowdhury et al. 2024"/> | ||
Recent research highlights the antimicrobial, antioxidant, and [[photocatalytic]] capabilities of [[zinc oxide]] [[nanoparticle]]s synthesised using ''Parmotrema perlatum'', marking a significant step towards sustainable dyeing practices and broadening the lichen's applicative horizons. | Recent research highlights the antimicrobial, antioxidant, and [[photocatalytic]] capabilities of [[zinc oxide]] [[nanoparticle]]s synthesised using ''Parmotrema perlatum'', marking a significant step towards sustainable dyeing practices and broadening the lichen's applicative horizons.<ref name="Khan et al. 2024"/> | ||
===Traditional medicines=== | ===Traditional medicines=== | ||
''Parmotrema perlatum'' is used as a component of a herbal mixture in [[Ayurvedic medicine]], one of several {{lichengloss|parmelioid}} lichen species used as ''charila''. Referenced in ancient Ayurvedic texts and first mentioned in the [[Atharvaveda]] around 1500 BCE, charila is a lichen mixture [[traditional medicine|traditionally]] used in India for its purported medicinal properties. It has been employed to treat various ailments, including digestive and respiratory issues, skin conditions, and reproductive health concerns, and it also serves as an ingredient in treatments for infertility. | ''Parmotrema perlatum'' is used as a component of a herbal mixture in [[Ayurvedic medicine]], one of several {{lichengloss|parmelioid}} lichen species used as ''charila''. Referenced in ancient Ayurvedic texts and first mentioned in the [[Atharvaveda]] around 1500 BCE, charila is a lichen mixture [[traditional medicine|traditionally]] used in India for its purported medicinal properties. It has been employed to treat various ailments, including digestive and respiratory issues, skin conditions, and reproductive health concerns, and it also serves as an ingredient in treatments for infertility.<ref name="Crawford 2019"/> For [[chronic ulcer]]s, a powder made from dried lichen, infused in pork suet, is applied externally.<ref name="Ravindran 2017"/> | ||
===Biomonitoring=== | ===Biomonitoring=== | ||
''Parmotrema perlatum'' is sensitive to [[air pollution]], making it a useful bioindicator. This sensitivity is utilised in the "Hawksworth and Rose" scale, which estimates mean winter [[sulphur dioxide]] (SO | ''Parmotrema perlatum'' is sensitive to [[air pollution]], making it a useful bioindicator. This sensitivity is utilised in the "Hawksworth and Rose" scale, which estimates mean winter [[sulphur dioxide]] (SO<sub>2</sub>) levels in England and Wales by observing lichens on acidic and nutrient-poor bark. According to this scale, ''P.&nbsp;perlatum'' is found only in zones 8 to 10, indicating areas with the lowest SO<sub>2</sub> concentrations, less than 35 micrograms per cubic metre (μg/m<sup>3</sup>).<ref name="Hawksworth & Rose 1970"/><ref name="Richardson 1992"/> | ||
A 2022 study analyzed the effects of SO | A 2022 study analyzed the effects of SO<sub>2</sub> and [[nitrogen dioxide]] (NO<sub>2</sub>) fumigation on the chlorophyll content of ''Parmotrema perlatum'' collected from the [[Mount Lawu]] volcano in Indonesia. The results indicated that increased exposure to these pollutants leads to a significant reduction in chlorophyll levels. The study demonstrated that SO<sub>2</sub> and NO<sub>2</sub> negatively impact the physiological processes of the lichen, particularly its [[photosynthetic efficiency]], demonstrating the sensitivity of ''P.&nbsp;perlatum'' to air pollution.<ref name="Roziaty et al. 2023"/> | ||
''Parmotrema perlatum'' has been effectively used in biomonitoring studies to assess environmental radioactivity. Research conducted in [[Turkey]] found that this lichen species retains radioactive [[caesium-137]] ( | ''Parmotrema perlatum'' has been effectively used in biomonitoring studies to assess environmental radioactivity. Research conducted in [[Turkey]] found that this lichen species retains radioactive [[caesium-137]] (<sup>137</sup>Cs) from atmospheric deposition, such as fallout from the [[Chernobyl accident]]. The [[ecological half-life]] of <sup>137</sup>Cs in ''Parmotrema perlatum'' was determined to be approximately 5.5 years, indicating its capability to monitor long-term radioactive contamination in the environment.<ref name="Cevik 2009"/> | ||
==Research== | ==Research== | ||
Research on the bioactive properties of ''Parmotrema perlatum'' has revealed several findings. The methanol extract of this species has been shown to significantly reduce [[blood glucose]] levels in [[streptozotocin]]-induced diabetic rats, attributed to its inhibitory activity on [[alpha-glucosidase]] rather than an effect on [[insulin]] secretion. This extract also has a high [[naturally occurring phenols|phenolic]] content and moderate [[antioxidant]] capacity, which could help prevent secondary complications of diabetes. | Research on the bioactive properties of ''Parmotrema perlatum'' has revealed several findings. The methanol extract of this species has been shown to significantly reduce [[blood glucose]] levels in [[streptozotocin]]-induced diabetic rats, attributed to its inhibitory activity on [[alpha-glucosidase]] rather than an effect on [[insulin]] secretion. This extract also has a high [[naturally occurring phenols|phenolic]] content and moderate [[antioxidant]] capacity, which could help prevent secondary complications of diabetes.<ref name="González-Burgos et al. 2019"/> The antioxidant potential and [[radical (chemistry)|free radical-scavening]] activity of ''P.&nbsp;perlatum'' extracts has been further demonstrated through various chemical assays.<ref name="Dwarakanath et al. 2022"/> | ||
Additionally, ''Parmotrema perlatum'' has some [[antimicrobial]] properties. The crude polysaccharide fraction of this lichen demonstrated [[antibacterial]] activity against ''[[Escherichia coli]]'' and ''[[Staphylococcus aureus]]'', which are common [[pathogen]]s in [[diabetic foot ulcer]]s. Furthermore, extracts from this species showed significant [[antiviral]] activity against the [[yellow fever virus]] envelope. | Additionally, ''Parmotrema perlatum'' has some [[antimicrobial]] properties. The crude polysaccharide fraction of this lichen demonstrated [[antibacterial]] activity against ''[[Escherichia coli]]'' and ''[[Staphylococcus aureus]]'', which are common [[pathogen]]s in [[diabetic foot ulcer]]s. Furthermore, extracts from this species showed significant [[antiviral]] activity against the [[yellow fever virus]] envelope.<ref name="González-Burgos et al. 2019"/> Tests against the [[Gram-negative]] bacteria ''[[Pseudomonas aeruginosa]]'', ''[[Chromobacterium violaceum]]'', and Gram-positive ''[[Lactobacillus plantarum]]'' showed that the [[methanol]] extract had the highest antibacterial activity among the three [[solvent]] extracts evaluated.<ref name="Dwarakanath et al. 2022"/> | ||
In terms of [[cytotoxic]] and anticancer activities, the [[hexane|''n''-hexane]], [[diethyl ether]], and methanol extracts of ''Parmotrema perlatum'' have been studied against various [[cancer cell line]]s, with the ''n''-hexane extract showing the highest cytotoxic effects. The extracts were particularly effective against murine [[Lewis lung carcinoma]] and human [[glioblastoma]] cell lines. | In terms of [[cytotoxic]] and anticancer activities, the [[hexane|''n''-hexane]], [[diethyl ether]], and methanol extracts of ''Parmotrema perlatum'' have been studied against various [[cancer cell line]]s, with the ''n''-hexane extract showing the highest cytotoxic effects. The extracts were particularly effective against murine [[Lewis lung carcinoma]] and human [[glioblastoma]] cell lines.<ref name="González-Burgos et al. 2019"/> | ||
==See also== | ==See also== | ||
| Line 178: | Line 178: | ||
{{Reflist|colwidth=30em|refs= | {{Reflist|colwidth=30em|refs= | ||
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<ref name="NatureServe">{{cite web |title=''Parmotrema perlatum''. Powdered Ruffle Lichen |url=https://explorer.natureserve.org/Taxon/ELEMENT_GLOBAL.2.123193/Parmotrema_perlatum |publisher=[[NatureServe]] |access-date=25 May 2024}}</ref> | |||
<ref name="Osbeck 1757">{{cite book |last=Osbeck |first=P. |year=1757 |title=Dagbok öfwer en ostindisk resa åren 1750, 1751, 1752 |trans-title=Diary of an East Indian journey in the years 1750, 1751, 1752 |location=Stockholm |publisher=Lorenz Ludvig Grefing}}</ref> | |||
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<ref name="Ravindran 2017">{{cite encyclopedia |last=Ravindran |first=P.N. |year=2017 |chapter=104. Kalpashi (Kalpasi) ''Parmotrema perlatum'' |title=The Encyclopedia of Herbs and Spices |publisher=CAB International |isbn=978-1-78064-315-1 |pages=493–496 |url=https://books.google.com/books?id=6pJNDwAAQBAJ&pg=PA493}}</ref> | |||
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}} | }} | ||
Latest revision as of 23:39, 21 March 2026
| Parmotrema perlatum | |
|---|---|
| |
| in Noordwijk, Netherlands | |
Scientific classification 
| |
| : | [[Template:Taxonomy/Parmotrema]] |
| Species: | Template:Taxonomy/ParmotremaP. perlatum
|
| Binomial name | |
| Template:Taxonomy/ParmotremaParmotrema perlatum | |
| Synonyms[2] | |
|
List
| |
Parmotrema perlatum, commonly known as the powdered ruffle lichen, is a common species of foliose lichen in the family Parmeliaceae. The species has a cosmopolitan distribution and occurs throughout the Northern and Southern Hemispheres. Parmotrema perlatum is a prominent and widely recognised species within its genus across primarily temperate zones, preferring humid, oceanic-suboceanic habitats. It is found in diverse geographic areas including Africa, North and South America, Asia, Australasia, Europe, and islands in the Atlantic and Pacific oceans. It usually grows on bark, but occasionally occurs on siliceous rocks, often among mosses.
The thallus of Parmotrema perlatum is large, light-grey to pale-blue patch-shaped with rounded and ruffled Template:Lichengloss and often with black hair-Template:Lichengloss at the edges. Distinguishing features of the lichen include its conspicuous soralia (reproductive structures) near the lobe edges, curled leaf-like lobes, and a narrow, shiny, and sometimes wrinkly area on the underside near the margin. This species is known for producing certain secondary metabolites, namely atranorin and a group of substances known as the stictic acid complex, which includes stictic and constictic acids, among other related compounds. These morphological and chemical characteristics help distinguish P. perlatum from several other potential lookalikes.
Parmotrema perlatum has a complex taxonomic history, having undergone multiple reclassifications since its original description in 1762. Significant efforts in the mid-20th century helped clarify its nomenclature, stabilising its current name. Although there were challenges to this name in the 1980s, it was confirmed as valid in 2004. More recently, DNA studies suggest that there may be hidden diversity within the species, indicating the need for further taxonomic evaluation.
The lichen is used as a spice in Indian cuisine. For this purpose, it is commonly known as black stone flower or kalpasi (among other names). Although nearly tasteless on its own, it releases an earthy fragrance and taste when cooked in with oil or butter.
Systematics[edit | edit source]
Historical taxonomy[edit | edit source]
The taxonomy of Parmotrema perlatum has a rich history marked by periods of confusion and clarification that typify the dynamic nature of botanical classification. It was originally described as Lichen perlatus by William Hudson in his 1762 work Flora Anglica. Hudson described it as a foliaceous (leafy) lichen with creeping, lobed, and smooth characteristics, having a pearly edge, a farinaceous (Template:Lichengloss) texture, and a black underside, adorned with slightly scalloped, brown, stalked fruiting bodies.[3] The taxon was later transferred to the genus Parmelia by Erik Acharius in 1803, becoming Parmelia perlata.[4] The name was well-established in scientific literature, being cited extensively in works like Alexander Zahlbruckner's popular 1929 catalogue.[5]
In 1952, Maurice Choisy reclassified it under the current name, Parmotrema perlatum.[6] The nomenclature of Parmotrema perlatum was revisited in the late 20th century, amid a broader effort to clarify the typification and application of early lichen names. Mason Hale, in 1961, undertook a detailed restudy of the species,[7] selecting a lectotype from the Dillenian collections—the herbarium and associated works of Johann Jacob Dillenius housed at the University of Oxford. This solidified the application of Hudson's name and was part of a larger trend in lichenology to fix historical names to specific herbarium specimens to stabilise nomenclature.[5]
The name Parmelia perlata was widely accepted until Hale and Ahti (1986) encountered the designation Lichen chinensis,[8] introduced by Pehr Osbeck in 1757.[9] They proposed the name Parmotrema chinense, based on the assumption that Osbeck's specimen corresponded to the well-known species Parmotrema perlatum. However, this proposal was not universally adopted due to the lack of valid typification and the name's absence in the literature between 1757 and 1986.[5]
Impact of the Tokyo Code (1993)[edit | edit source]
The Tokyo Code of 1993 extended the provisions for conserving names to all species, not just those of major economic importance. This change in the International Code of Botanical Nomenclature allowed for the conservation of names that would promote nomenclatural stability. Despite this provision, no formal proposal was made to conserve the name Parmotrema chinense, and thus it did not gain widespread acceptance.[5]
David Hawksworth's 2004 study brought significant clarity to the taxonomic confusion. He rediscovered Osbeck's original material in Linnaeus' herbarium and identified it as belonging to Parmotrema tinctorum, not Parmotrema perlatum. Hawksworth demonstrated that Lichen chinensis was not validly published because it lacked a proper description and was linked with an expression of doubt by Osbeck. Hawksworth's work led to the reinstatement of the name Parmotrema perlatum, confirming that Hudson's name was legitimate and should continue to be used. This resolution was based on the original typification by Hale and the invalid publication status of Lichen chinensis.[5]
Recent studies suggest that the circumscription of Parmotrema perlatum may need to be revised. Research utilising DNA sequencing has uncovered cryptic diversity within the genus Parmotrema, indicating that traditional phenotype-based identification methods may underestimate species diversity. Specifically, the genetic analysis of P. perlatum and related species revealed multiple distinct lineages that were previously grouped under a single nominal taxon. These findings highlight the need for a comprehensive taxonomic re-evaluation of P. perlatum to accurately delineate species boundaries and account for hidden genetic diversity.[10]
Phylogeny[edit | edit source]
In molecular phylogenetics analysis, Parmotrema perlatum has a sister relationship with Parmotrema crinitum. These two species form a clade that itself is sister to a clade with P. austrosinense and P. tinctorum.[11] In a comprehensive phylogenetic analysis by Stelate and colleagues (2022), P. perlatum and P crinitum were found to form a well-supported monophyletic group using internal transcribed spacer sequences and several analytical methods. This study highlights the complexity of species boundaries within the genus and the need for further research incorporating additional molecular markers to confirm these findings.[12]
Common names[edit | edit source]
Vernacular names used for this species include black stone flower, stone lichen, sea lichen, kalpasi, kalpas, kalpashi, and kalpash. The latter name and its variations, however, have been used as a crude drug in Indian medicines for more than one species, including Parmotrema perlatum, Parmotrema tinctorum, and Everniastrum cirrhatum.[13] In North America, vernacular names used for the species include "powdered ruffle lichen",[14][15] "powdered scatter-rug",[16] and "queen ruffle".[17] The species epithet perlatum refers to the pearl-like margins of the lobes, which are directly referenced in Hudson's original 1762 description of the species. He proposed the English name "pearl lichen";[3] this name later morphed into "pearly parmelia" in some 19th-century British accounts of lichen flora.[18][19]
Description[edit | edit source]
Closeup of soredious lobe edges ...
... and marginal cilia.
Parmotrema perlatum has a thallus that ranges from loosely to tightly attached to the surface it grows on, forming expansive, spreading colonies that often merge together.[20] Individual thalli typically measure up to 10–15 cm (4–6 in) in diameter. The upper thallus surface is greenish-grey, blue-grey,[15] or yellowish-grey in colour,[14] lacking Template:Lichengloss and either free of spots (Template:Lichengloss),[15] or with few maculae.[14] This species develops soredia, a type of asexual reproductive structure, aiding in its propagation. The Template:Lichengloss of this lichen vary from 1.5 to 10 mm in width, with a wave-like (Template:Lichengloss) or ruffled pattern and overlapping (Template:Lichengloss) arrangement. The tips and edges of these lobes are generally smooth and round, sometimes notched (Template:Lichengloss) or incised, often curling up or inward, revealing the paler brown to black underside adorned with hair-like structures (Template:Lichengloss) up to 2.5 mm in length.[20] Rhizines are common on the underside of the thallus, except for a brown border near the edges.[15]
The soredia found in this species are Template:Lichengloss and appear white or may become grey due to wear. They are located within specifically structured groups called soralia, which can be linear to oval in shape, often positioned at the edges of the lobes.[20] The presence of soredia causes the lobe margins to curl back and form Template:Lichengloss soralia.[15] The upper surface of the lichen is typically whitish grey to pale greenish-grey, and can be either smooth or slightly wrinkled, without spots (Template:Lichengloss), featuring scattered, shallow cracks.[20] Isidia are absent in this species.[15]
Apothecia (fruiting bodies) are rare in Parmotrema perlatum. When present, they measure 4–8 mm across and are somewhat stalked and funnel-shaped with a brown, concave Template:Lichengloss. The edges of these structures curl inward, becoming thick with soredia as they mature. Its spores are ellipsoid in shape and typically measure between 20 and 28 μm in length and 11 to 17 μm in width, with a wall thickness of 2–3 μm. Pycnidia, which are structures that produce asexual spores called conidia, appear sporadically on the surface (Template:Lichengloss) of the thallus, with the conidia being thread-like and straight, measuring 6–8 by 1 μm.[20]
Photobiont[edit | edit source]
The photobiont partner of Parmotrema perlatum is from Trebouxia, a green algal genus belonging to the order Trebouxiales (order Chlorophyta).[21] It has been identified as an undescribed species within a clade containing Trebouxia arboricola.[22]
A study compared the desiccation tolerance and physiological responses of lichenised Trebouxia to isolated cultures of the same alga. Both forms can survive extended desiccation, but with differing responses to photo-oxidative stress. Lichenisation enhances the photoprotective mechanisms of Trebouxia, improving quenching of excess light energy, particularly under high relative humidity, and controlling reactive oxygen species production under light exposure. However, isolated cultures showed better photosynthetic performance after desiccation recovery. This research demonstrates the mutual benefits of the lichen-photobiont partnership, where the alga gains a sheltered environment boosting its resilience to environmental stressors.[21]
Further studies on Parmotrema perlatum revealed specific antioxidant mechanisms supporting its photobiont under stress. The lichen shows high levels of reactive oxygen species scavenging enzymes such as superoxide dismutase and ascorbate peroxidase, protecting the photobiont from oxidative damage during dehydration and rehydration cycles. This enhanced antioxidant system provides not only physical shelter but also biochemical protection, increasing the photobiont's resilience to environmental fluctuations.[22]
Chemistry[edit | edit source]
Chemically, Parmotrema perlatum contains atranorin and chloroatranorin, alongside a predominant stictic acid Template:Lichengloss that includes stictic as a major secondary metabolite and smaller amounts of constictic acid and other related substances. Testing the medulla (the inner layer beneath the upper cortex) with spot tests results in K+ (yellow), KC–, and P+ (orange) reactions.[20] The cortical layer, in contrast, is K+ (yellow), KC–, and P–.[15]
The secondary metabolites of Parmotrema perlatum have been studied using gas chromatography–mass spectrometry (GC–MS) and liquid chromatography–mass spectrometry (LC–MS/MS). The lichen produces several notable compounds, including orcinol, atraric acid, benzoic acid, 2,4-dihydroxy-3,6-dimethyl-, methyl ester, and palmitic acid, methyl ester. GC–MS analysis revealed the presence of orcinol (63%) and atraric acid (21%) in the methanol extract, while benzoic acid was predominant in the chloroform extract. The hexane extract contained significant amounts of benzoic acid, 2,4-dihydroxy-3,6-dimethyl-, methyl ester (62%).[23] A more recent study using liquid chromatography-electrospray ionization-mass spectrometry/mass spectrometry as an analysis technique tentatively identified a total of twenty-five lichen products, including 5 depsides, 12 depsidones, 2 diphenyl ethers, 1 aromatic considered as possible artifact, 1 dibenzofuran, 1 carbohydrate, 1 organic acid, and 2 undefined compounds.[24]
Similar species[edit | edit source]
The distinguishing features of Parmotrema perlatum, such as the presence of soredia and stictic acid, facilitate its easy identification. In mature specimens, the appearance of scattered, fine cracks on the upper surface may resemble the cracked maculae seen in P. reticulatum, which shares similar habitats. However, the two species can be differentiated chemically, as P. reticulatum contains salazinic acid, unlike P. perlatum.[20]
Parmotrema perlatum and Parmotrema stuppeum are two morphologically similar species that can be found in similar habitats. Both species have a loosely attached thallus with revolute, wavy lobes and sparsely ciliate lobe tips. Their upper cortex is continuous and not finely reticulately cracked, while the lower surface is black and rhizinate. Both species also feature linear soralia. However, there are several key differences that can help distinguish between the two. While earlier descriptions suggested that P. stuppeum has a matte, olive-green to brownish-green upper surface and P. perlatum has a slightly shiny, whitish-grey to greyish-green upper surface, recent observations have shown that both species have a distinctly matte upper surface with similar colouration.[25]
The most reliable morphological difference in the field is the location of the soralia: P. stuppeum has strictly terminal soralia, whereas P. perlatum has submarginal soralia. Additionally, the two species can be distinguished by their chemical composition. P. stuppeum contains salazinic acid, while P. perlatum has a stictic acid complex. Although both acids cause a Pd+ orange to orange-red medulla reaction, a potassium (K) spot test can separate the species: the medulla of P. perlatum turns yellow (K+ yellow), whereas in P. stuppeum, the yellow colour turns red (K+ yellow turning red).[25]
Parmotrema perlatum can be distinguished from other sorediate and marginally ciliate species like P. arnoldii and P. robustum by the presence of the stictic acid chemosyndrome.[26] Also, the medulla of P. arnoldii fluoresces strongly when lit with an ultraviolet lamp.[27] Parmotrema perlatum is similar to P. crinitum due to both species having a brown to tan, erhizinate marginal zone and the presence of the stictic acid chemosyndrome in the medulla. However, P. crinitum can be distinguished by its isidiate upper surface.[28] Another potential lookalike, P. margaritatum, is distinguished from P. perlatum by the K+ (red) reaction of its medulla.[15] Cetrelia cetrarioides has been documented as a lookalike, presumably because of the cilia on its thallus margin, and the presence of atranorin and the stictic acid chemosyndrome.[29]
Habitat and distribution[edit | edit source]
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Parmotrema perlatum typically grows in areas with ample light, favouring neutral to slightly acidic-barked broad-leaved trees. It is commonly found on siliceous rocks and walls, as well as mossy coastal rocks, generally growing in places with moderate to strong sunlight.[30] In the Great Smoky Mountains National Park in the United States, Parmotrema perlatum is especially abundant on branches in humid, high-elevation habitats.[31] Similarly, in East Africa, it grows in the misty environments of inselbergs, montane forests, and Erica-dominated habitats, typically found between 1,400 and 3,100 m (4,600 and 10,200 ft) above sea level.[32]
The species is globally distributed, found in both temperate and tropical regions. It has been reported across numerous European countries including Austria, Belgium, the Czech Republic, France, Germany, Great Britain, Ireland, Italy, Luxembourg, the Netherlands, Portugal, Scandinavia, Slovakia, Spain, and Ukraine.[26] Although it is rare in Eastern Europe, it is widely distributed in both the Asian and European parts of Russia.[25] Beyond Europe, it is also present in Macaronesia, Africa, Australia, North America, and South America.[26] Its Asian distribution includes India, Japan, Taiwan, and South Korea.[28] Although it has historically been recorded in Nepal and Sri Lanka, these reports are considered tentative due to shifting species concepts and possible confusion with the lookalike Parmotrema pseudonilgherrense.[33]
Parmotrema perlatum is globally widespread lichen found on all continents except Antarctica and predominantly in oceanic areas in Europe, primarily grows on bark and occasionally on siliceous rocks amongst mosses. While it is seeing an increase in the Netherlands due to global warming, it is critically endangered in the Czech Republic,[34] Slovakia, and Poland[35] due to susceptibility to air pollution, and is listed as extinct in certain regional Red Data Books due to a lack of recent findings.[36] In contrast, it has been increasing in sightings in the Netherlands, a phenomenon attributed to both global warming decreases in the levels or air pollution in recent decades.[37][38] Its recent recurrence in Hungary, particularly on some unusual hosts (Catalpa bignonioides, Prunus serotina, and Robinia pseudoacacia) have been suggested as a possible consequence of "a recolonisation process, due to the improving air quality".[39]
Ecology[edit | edit source]
Parmotrema perlatum is an important species within specific lichen communities in British woodlands, particularly those in late successional mesotrophic settings in oceanic or humid microclimates. It is associated with the Type K Lobaria pulmonaria-Isothecium myosuroides ecological Community. This community type is characterised by its occurrence in mature mesotrophic environments, which are often warmer in winter climates or specific microhabitats. This community includes, in addition to P. perlatum, dominant foliose lichens like Lobaria pulmonaria, Hypotrachyna taylorensis, and Parmotrema crinitum, as well as bryophytes such as Isothecium myosuroides.[40]
A 2017 study investigated the physiological responses of Parmotrema perlatum along an aridity gradient in Southern Portugal. The researchers transplanted thalli of P. perlatum to rural and forested sites characterised by varying levels of aridity and measured several physiological parameters, including photosynthetic performance, pigment content, ergosterol content, and sample viability, both before and after a six-month exposure period. The study found that P. perlatum showed lower photosynthetic performance (measured as FV/FM and the performance index on an absorption basis, PIABS) in drier sites compared to more humid sites. In humid environments, the content of photosynthetic pigments increased post-exposure, while in drier sites, this increase was less pronounced. Additionally, ergosterol content was lower in drier sites, indicating a stress response to arid conditions. These results highlight that P. perlatum's physiological responses are significantly influenced by water availability. The ability to maintain higher photosynthetic performance and pigment content in humid conditions suggests that P. perlatum is better adapted to environments with higher moisture levels. This adaptability makes P. perlatum useful as a bioindicator for monitoring ecological responses to climate change and varying moisture conditions in Mediterranean ecosystems.[41]
Lichenicolous (lichen-dwelling) fungi that have been recorded parasitising Parmotrema perlatum include Abrothallus parmotrematis, Briancoppinsia cytospora, Lichenoconium erodens, and Spirographa lichenicola.[42]
Conservation[edit | edit source]
Parmotrema perlatum has been identified as a species of concern in some regions due to its rarity and declining populations. In Hungary, it has been proposed for 'endangered' status in the Hungarian lichen red list, reflecting its limited distribution and the pressures it faces in its natural habitats there.[39] Similarly, in Ukraine, the species is listed in the Red Data Book of Ukraine with the status of "Rare".[43] Additionally, Parmotrema perlatum is red-listed in Sweden[44] In northern North America, its NatureServe conservation status is designated as "G4", meaning "apparently secure" at the global level. In the United States, it has been assessed as secure in Kentucky and presumed extirpated in Wisconsin, while in Canada, it is considered as vulnerable in British Columbia and Ontario, and critically imperiled in New Brunswick.[45]
Uses[edit | edit source]
As a spice[edit | edit source]
Parmotrema perlatum is used as a spice, particularly in the cuisine of Tamil Nadu. It is especially prevalent in Chettinad cuisine, being used in the popular rice dish biryani, and also in many meat and vegetarian dishes.[13]
In its raw state, black stone flower does not have much taste or fragrance. However, when put in contact with heat, especially hot cooking oil or ghee, it releases a distinctive earthy, smoky flavour and aroma. This property of black stone flower is especially valued in the tempering step of cooking a number of Indian dishes.[13][46]
The spice is also integral to various regional masalas throughout the Indian subcontinent. Parmotrema perlatum is a key ingredient in masalas such as Kala and Goda masala of Maharashtra, Anglo-Indian bottle masala, bhojwar masala from Hyderabad, and potli masala in Lucknow. It is what many cooks and commercial spice blend makers believe sets apart accomplished dishes from those made by amateurs. Despite its lack of a specific aroma or describable flavour in its raw form, its contribution to the complex flavour profile of these spice blends is highly valued.[46]
Dyeing[edit | edit source]
A natural purple dye extracted from Parmotrema perlatum using ammonia fermentation showed optimal results, with a notable dye yield and effective application on silk fabric. The study demonstrated the dye's potential as a sustainable alternative to synthetic dyes, with satisfactory colour fastness and fabric strength enhancement.[47]
Recent research highlights the antimicrobial, antioxidant, and photocatalytic capabilities of zinc oxide nanoparticles synthesised using Parmotrema perlatum, marking a significant step towards sustainable dyeing practices and broadening the lichen's applicative horizons.[48]
Traditional medicines[edit | edit source]
Parmotrema perlatum is used as a component of a herbal mixture in Ayurvedic medicine, one of several Template:Lichengloss lichen species used as charila. Referenced in ancient Ayurvedic texts and first mentioned in the Atharvaveda around 1500 BCE, charila is a lichen mixture traditionally used in India for its purported medicinal properties. It has been employed to treat various ailments, including digestive and respiratory issues, skin conditions, and reproductive health concerns, and it also serves as an ingredient in treatments for infertility.[49] For chronic ulcers, a powder made from dried lichen, infused in pork suet, is applied externally.[13]
Biomonitoring[edit | edit source]
Parmotrema perlatum is sensitive to air pollution, making it a useful bioindicator. This sensitivity is utilised in the "Hawksworth and Rose" scale, which estimates mean winter sulphur dioxide (SO2) levels in England and Wales by observing lichens on acidic and nutrient-poor bark. According to this scale, P. perlatum is found only in zones 8 to 10, indicating areas with the lowest SO2 concentrations, less than 35 micrograms per cubic metre (μg/m3).[50][51]
A 2022 study analyzed the effects of SO2 and nitrogen dioxide (NO2) fumigation on the chlorophyll content of Parmotrema perlatum collected from the Mount Lawu volcano in Indonesia. The results indicated that increased exposure to these pollutants leads to a significant reduction in chlorophyll levels. The study demonstrated that SO2 and NO2 negatively impact the physiological processes of the lichen, particularly its photosynthetic efficiency, demonstrating the sensitivity of P. perlatum to air pollution.[52]
Parmotrema perlatum has been effectively used in biomonitoring studies to assess environmental radioactivity. Research conducted in Turkey found that this lichen species retains radioactive caesium-137 (137Cs) from atmospheric deposition, such as fallout from the Chernobyl accident. The ecological half-life of 137Cs in Parmotrema perlatum was determined to be approximately 5.5 years, indicating its capability to monitor long-term radioactive contamination in the environment.[53]
Research[edit | edit source]
Research on the bioactive properties of Parmotrema perlatum has revealed several findings. The methanol extract of this species has been shown to significantly reduce blood glucose levels in streptozotocin-induced diabetic rats, attributed to its inhibitory activity on alpha-glucosidase rather than an effect on insulin secretion. This extract also has a high phenolic content and moderate antioxidant capacity, which could help prevent secondary complications of diabetes.[54] The antioxidant potential and free radical-scavening activity of P. perlatum extracts has been further demonstrated through various chemical assays.[23]
Additionally, Parmotrema perlatum has some antimicrobial properties. The crude polysaccharide fraction of this lichen demonstrated antibacterial activity against Escherichia coli and Staphylococcus aureus, which are common pathogens in diabetic foot ulcers. Furthermore, extracts from this species showed significant antiviral activity against the yellow fever virus envelope.[54] Tests against the Gram-negative bacteria Pseudomonas aeruginosa, Chromobacterium violaceum, and Gram-positive Lactobacillus plantarum showed that the methanol extract had the highest antibacterial activity among the three solvent extracts evaluated.[23]
In terms of cytotoxic and anticancer activities, the n-hexane, diethyl ether, and methanol extracts of Parmotrema perlatum have been studied against various cancer cell lines, with the n-hexane extract showing the highest cytotoxic effects. The extracts were particularly effective against murine Lewis lung carcinoma and human glioblastoma cell lines.[54]
See also[edit | edit source]
References[edit | edit source]
- ↑ Template:Cite NatureServe
- ↑ "GSD Species Synonymy. Current Name: Parmotrema perlatum (Huds.) M. Choisy, Bull. mens. Soc. linn. Soc. Bot. Lyon 21: 174 (1952)". Species Fungorum. Retrieved 25 May 2024.
- ↑ 3.0 3.1 Hudson, William (1762). Flora Anglica (in Latina). London: Prostant Venales apud J. Nourse, in the Strand. p. 543.
- ↑ Acharius, Erik (1803). Methodus qua omnes detectos Lichenes : secundum organa carpomorpha, ad genera, species et varietates [Method by which all detected lichens: according to carpomorph organs, are classified into genera, species, and varieties] (in Latina). Stockholm: F.D.D. Ulrich. p. 216.
- ↑ 5.0 5.1 5.2 5.3 5.4 Hawksworth, D.L. (2004). "Rediscovery of the original material of Osbeck's Lichen chinensis and the reinstatement of the name Parmotrema perlatum" (PDF). Herzogia. 17: 37–44.
- ↑ Choisy, M. (1952). "Catalogue des lichens de la region lyonnaise. Fasc. 9" [Catalogue of Lichens of the Lyon Region. Fascicle 9]. Bulletin Mensuel de la Société Linnéenne de Lyon (in français). 21: 165–180. doi:10.3406/linly.1952.7511.
- ↑ Hale, M.E. (1961). "The typification of Parmelia perlata (Huds.) Ach". Brittonia. 13 (4): 361–367. Bibcode:1961Britt..13..361H. doi:10.2307/2805414. JSTOR 2805414.
- ↑ Hale, Mason E.; Ahti, Teuvo (1986). "An earlier name for Parmotrema perlatum (Huds.) Choisy (Ascomycotina: Parmeliaceae)". Taxon. 35 (1): 133–134. doi:10.2307/1221050. JSTOR 1221050.
- ↑ Osbeck, P. (1757). Dagbok öfwer en ostindisk resa åren 1750, 1751, 1752 [Diary of an East Indian journey in the years 1750, 1751, 1752]. Stockholm: Lorenz Ludvig Grefing.
- ↑ Del-Prado, Ruth; Buaruang, Kawinnat; Lumbsch, H. Thorsten; Crespo, Ana; Divakar, Pradeep K. (2019). "DNA sequence-based identification and barcoding of a morphologically highly plastic lichen forming fungal genus (Parmotrema, Parmeliaceae) from the tropics". The Bryologist. 122 (2): 281–291. doi:10.1639/0007-2745-122.2.281.
- ↑ Crespo, Ana; Kauff, Frank; Divakar, Pradeep K.; del Prado, Ruth; Pérez-Ortega, Sergio; de Paz, Guillermo Amo; et al. (2010). "Phylogenetic generic classification of parmelioid lichens (Parmeliaceae, Ascomycota) based on molecular, morphological and chemical evidence". Taxon. 59 (6): 1735–1753. doi:10.1002/tax.596008.
- ↑ Stelate, Ayoub; Del-Prado, Ruth; Alors, David; Tahiri, Hikmat; Divakar, Pradeep K.; Crespo, Ana (2022). "Resolving the phylogenetic relationship between Parmotrema crinitum and Parmotrema perlatum populations". The Lichenologist. 54 (3–4): 183–194. doi:10.1017/S0024282922000147.
- ↑ 13.0 13.1 13.2 13.3 Ravindran, P.N. (2017). "104. Kalpashi (Kalpasi) Parmotrema perlatum". The Encyclopedia of Herbs and Spices. CAB International. pp. 493–496. ISBN 978-1-78064-315-1.
- ↑ 14.0 14.1 14.2 Brodo, Irwin M.; Sharnoff, Sylvia Duran; Sharnoff, Stephen (2001). Lichens of North America. Yale University Press. p. 493. ISBN 978-0-300-08249-4.
- ↑ 15.0 15.1 15.2 15.3 15.4 15.5 15.6 15.7 McMullin, R. Troy (2023). Lichens. The Macrolichens of Ontario and the Great Lakes Region of the United States. Firefly Books. p. 332. ISBN 978-0-228-10369-1.
- ↑ Goward, Trevor; McCune, Bruce; Meidinger, Del (1994). The Lichens of British Columbia: Illustrated Keys. Part 1 — Foliose and Squamulose Species. Victoria, B.C.: Ministry of Forests Research Program. p. 94. ISBN 0-7726-2194-2. OCLC 31651418.
- ↑ "BC Conservation Data Centre: Species Summary. Parmotrema perlatum". British Columbia Ministry of Environment. Retrieved 25 May 2024.
- ↑ Mosley, sir Oswald (1863). The Natural History of Tutbury. Together with the Fauna and Flora of the District Surrounding Tutbury and Burton-on-Trent. London: John Van Voorst, Paternoster Row. p. 319.
- ↑ Smith, James Edward (1844). English Botany; Or Coloured Figures of British Plants, with Their Essential Characters, Synonyms, and Places of Growth. Vol. 11 (2 ed.). London: R. Taylor. p. 18.
- ↑ 20.0 20.1 20.2 20.3 20.4 20.5 20.6 Kantvilas, G. (2023). de Salas, M.F. (ed.). "Parmotrema, version 2023:1". Flora of Tasmania Online. Tasmanian Herbarium, Tasmanian Museum and Art Gallery.
- ↑ 21.0 21.1 Candotto Carniel, Fabio; Zanelli, Davide; Stefano, Bertuzzi; Tretiach, Mauro (2015). "Desiccation tolerance and lichenization: a case study with the aeroterrestrial microalga Trebouxia sp. (Chlorophyta)". Planta. 242 (2): 493–505. Bibcode:2015Plant.242..493C. doi:10.1007/s00425-015-2319-z. hdl:11368/2871352. PMID 25998523.
- ↑ 22.0 22.1 Bertuzzi, Stefano; Pellegrini, Elisa; Candotto Carniel, Fabio; Incerti, Guido; Lorenzini, Giacomo; Nali, Cristina; Tretiach, Mauro (2017). "Ozone and desiccation tolerance in chlorolichens are intimately connected: a case study based on two species with different ecology". Environmental Science and Pollution Research. 25 (9): 8089–8103. doi:10.1007/s11356-017-9444-0. PMID 28646314.
- ↑ 23.0 23.1 23.2 Dwarakanath, P.R.; Abinaya, K.; Nagasathya, K.; Meenakumari, S.; Gopinath, Subash C.B.; Raman, Pachaiappan (2022). "Profiling secondary metabolites from lichen Parmotrema perlatum (Huds.) M.Choisy" and antibacterial and antioxidant potentials". Biomass Conversion and Biorefinery. doi:10.1007/s13399-022-03572-0.
- ↑ Castañeta, Grover; Sepulveda, Beatriz; Areche, Carlos (2024). "Liquid chromatography-electrospray ionization-mass spectrometry/mass spectrometry characterization of depsides and depsidones from the Chilean lichen Parmotrema perlatum". European Journal of Mass Spectrometry. 30 (2): 125–132. doi:10.1177/14690667241240477. PMID 38523368.
- ↑ 25.0 25.1 25.2 Tsurykau, Andrei; Golubkov, Vladimir; Belyt, Pavel (2015). "The genera Hypotrachyna, Parmotrema and Punctelia (Parmeliaceae, lichenized Ascomycota) in Belarus". Herzogia. 28 (2): 736–745. doi:10.13158/heia.28.2.2015.736.
- ↑ 26.0 26.1 26.2 Jabłońska, Agnieszka; Oset, Magdalena; Kukwa, Martin (2009). "The lichen family Parmeliaceae in Poland. I. The genus Parmotrema". Acta Mycologica. 44 (2): 211–222. doi:10.5586/am.2009.019.
- ↑ McCune, Bruce; Geiser, Linda (2009). Macrolichens of the Pacific Northwest (2nd ed.). Corvallis: Oregon State University Press. p. 233. ISBN 978-0-87071-394-1.
- ↑ 28.0 28.1 Jayalal, Udeni; Divakar, Pradeep K.; Joshi, Santosh; Oh, Soon-Ok; Koh, Young Jin; Hur, Jae-Seoun (2013). "The lichen genus Parmotrema in South Korea". Mycobiology. 41 (1): 25–36. doi:10.5941/MYCO.2013.41.1.25. PMC 3627967. PMID 23610536.
- ↑ Kukwa, Martin; Pietnoczko, Magdalena; Czyżewska, Krystyna (2011). "The lichen family Parmeliaceae in Poland. II. The genus Cetrelia". Acta Societatis Botanicorum Poloniae. 81 (1): 43–52 [44]. doi:10.5586/asbp.2012.007.
- ↑ Cannon, P.; Divakar, P.; Yahr, R.; Aptroot, A.; Clerc, P.; Coppins, B.; Fryday, A.; Sanderson, N.; Simkin, J. (2023). Lecanorales: Parmeliaceae, including the genera Alectoria, Allantoparmelia, Arctoparmelia, Brodoa, Bryoria, Cetraria, Cetrariella, Cetrelia, Cornicularia, Evernia, Flavocetraria, Flavoparmelia, Hypogymnia, Hypotrachyna, Imshaugia, Melanelia, Melanelixia, Melanohalea, Menegazzia, Montanelia, Nesolechia, Parmelia, Parmelina, Parmeliopsis, Parmotrema, Platismatia, Pleurosticta, Protoparmelia, Pseudephebe, Pseudevernia, Punctelia, Raesaenenia, Tuckermannopsis, Usnea, Vulpicida and Xanthoparmelia (PDF). Revisions of British and Irish Lichens. Vol. 33. pp. 52–53.
- ↑ Tripp, Erin A.; Lendemer, James C. (2020). Field Guide to the Lichens of Great Smoky Mountains National Park. Knoxville: The University of Tennessee Press. p. 328. ISBN 978-1-62190-514-1.
- ↑ Swinscow, Thomas Douglas Victor; Krog, Hildur (1988). Macrolichens of East Africa. London: British Museum (Natural History). p. 170. ISBN 978-0-565-01039-3.
- ↑ Awasthi, Dharani Dhar (2007). A Compendium of the Macrolichens from India, Nepal and Sri Lanka. Dehra Dun, India: Bishen Singh Mahendra Pal Singh. p. 345. ISBN 978-8121106009.
- ↑ Liška, J.; Palice, Z.; Slavíková, Š. (2008). "Checklist and Red List of lichens of the Czech Republic". Preslia. 80 (2): 151–182.
- ↑ Cieśliński, S.; Czyzewska, K.; Fabiszewski, J. (2003). "Czerwona lista porostów wymarlych i zagrozonych w Polsce" [Red List of extinct and threatened lichens in Poland]. Monographiae Botanicae. 91: 13–49. doi:10.5586/mb.2003.001.
- ↑ Vondrák, Jan; Liška, Jiří (2010). "Changes in distribution and substrate preferences of selected threatened lichens in the Czech Republic". Biologia. 65 (4): 595–602. Bibcode:2010Biolg..65..595V. doi:10.2478/s11756-010-0061-3.
- ↑ van Hark, C.M.; Aptroot, A.; van Dobben, H.F. (2002). "Long-term monitoring in the Netherlands suggests that lichens respond to global warming". The Lichenologist. 34 (2): 141–154. Bibcode:2002ThLic..34..141V. doi:10.1006/lich.2002.0378.
- ↑ Aptroot, A.; van Herk, C.M. (2007). "Further evidence of the effects of global warming on lichens, particularly those with Trentepohlia phycobionts". Environmental Pollution. 146 (2): 293–298. Bibcode:2007EPoll.146..293A. doi:10.1016/j.envpol.2006.03.018. PMID 16697507.
- ↑ 39.0 39.1 Aszalósné Balogh, R.; Buczkó, K.; Erzberger, P.; Freytag, Cs.; Homm, Th.; Lőkös, L.; Matus, G.; Nagy, Z.; Papp, B.; Farkas, E. (2021). "Taxonomical and chorological notes 15 (153–163)" (PDF). Studia Botanica Hungarica. 52 (2): 165–184. doi:10.17110/StudBot.2021.52.2.165.
- ↑ Ellis, Christopher J.; Eaton, Sally; Theodoropoulos, Marios; Elliott, Kathryn (2015). Epiphyte Communities and Indicator Species. An Ecological Guide for Scotland's Woodlands (Report). Edinburgh: Royal Botanic Garden Edinburgh. pp. 112–113. ISBN 978-1-910877-01-2.
- ↑ Paoli, Luca; Pinho, Pedro; Branquinho, Cristina; Loppi, Stefano; Munzi, Silvana (2017). "The influence of growth form and substrate on lichen ecophysiological responses along an aridity gradient". Environmental Science and Pollution Research. 24 (34): 26206–26212. Bibcode:2017ESPR...2426206P. doi:10.1007/s11356-017-9361-2. PMID 28664492.
- ↑ Sanderson, N.A. "Parmotrema perlatum". British Lichen Society. Retrieved 25 May 2024.
- ↑ Didukh, Ya. P., ed. (2009). Червона книга України [Red data book of Ukraine. Vegetable kingdom] (in українська). Global Consulting Ukraine.
- ↑ Thell, A.; Thor, G.; Ahti, T. (2011). "Parmelia". In Thell, Arne; Moberg, Roland (eds.). Nordic Lichen Flora. Vol. 4. Svenska Botaniska Föreningen. pp. 96–97. ISBN 978-91-85221-24-0.
- ↑ "Parmotrema perlatum. Powdered Ruffle Lichen". NatureServe. Retrieved 25 May 2024.
- ↑ 46.0 46.1 Mani, Priya (2021). "Stone Curry: Parmotrema perlatum as a secret spice in Indian Food". In McWilliams, Mark (ed.). Herbs and Spices: Proceedings of the Oxford Symposium on Food and Cookery 2020. Prospect Books. pp. 239–249. ISBN 978-1909248724.
- ↑ Roychowdhury, Rupsa; Maiti, Saptarshi; Adivarekarb, Ravindra V.; Singhal, Rekha S. (2024). "Sustainable dyeing of silk using an acetylshikonin-based natural colourant from the lichen Parmotrema perlatum". Green Chemistry. 26 (2): 904–917. doi:10.1039/D3GC03686C.
- ↑ Khan, Parvez Pathan; Talaviya, Guranga; Vadavarita, Samim; Desai, Harshal; Togadiya, Vijay; Rakholiya, Kalpna D.; Kaneria, Mital J.; Padalia, Hemali (2023). "Evaluation of antimicrobial, antioxidant, and photocatalytic activity of zinc oxide nanoparticles synthesized from Parmotrema perlatum". In Kaneria, Mital; Rakholiya, Kalpna; Egbuna, Chukwuebuka (eds.). Nanotechnology and In Silico Tools. Natural Remedies and Drug Discovery. Elsevier. doi:10.1016/B978-0-443-15457-7.00019-8. ISBN 978-0-443-15457-7.
- ↑ Crawford, Stuart (2019). "Lichens Used in Traditional Medicine". In Ranković, Branislav (ed.). Lichen Secondary Metabolites. Bioactive Properties and Pharmaceutical Potential (2 ed.). Springer Nature Switzerland AG. pp. 60–62. ISBN 978-3-030-16813-1.
- ↑ Hawksworth, D. L.; Rose, F. (1970). "Qualitative scale for estimating sulphur dioxide air pollution in England and Wales using epiphytic lichens". Nature. 227 (5254): 145–148. Bibcode:1970Natur.227..145H. doi:10.1038/227145a0. PMID 5428399.
- ↑ Richardson, D.H.S. (1992). Pollution Monitoring with Lichens. Naturalists' Handbook. Vol. 19. Pelagic Publishing. p. 8. ISBN 978-1-78427-211-1.
- ↑ Roziaty, Efri; Sutarno; Suntoro; Sugiyarto (2023). "Short Communication: The effects of SO2 and NO2 fumigation on the chlorophyll of Parmotrema perlatum from Mt. Lawu, Cemoro Sewu, Indonesia". Biodiversitas. 24 (5): 2630–2637. doi:10.13057/biodiv/d240515.
- ↑ Cevik, Ugur; Celik, Necati (2009). "Ecological half-life of 137Cs in mosses and lichens in the Ordu province, Turkey by Cevik and Celik". Journal of Environmental Radioactivity. 100 (1): 23–38. doi:10.1016/j.jenvrad.2008.09.010. PMID 19013696.
- ↑ 54.0 54.1 54.2 González-Burgos, Elena; Fernández-Moriano, Carlos; Gómez-Serranillos, M. Pilar (2019). "Current knowledge on Parmelia genus: Ecological interest, phytochemistry, biological activities and therapeutic potential". Phytochemistry. 165. Bibcode:2019PChem.165k2051G. doi:10.1016/j.phytochem.2019.112051. PMID 31234093.