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 and often with black hair- 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
Historical taxonomy 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 () texture, and a black underside, adorned with slightly scalloped, brown, stalked fruiting bodies. The taxon was later transferred to the genus Parmelia by Erik Acharius in 1803, becoming Parmelia perlata. The name was well-established in scientific literature, being cited extensively in works like Alexander Zahlbruckner's popular 1929 catalogue.
In 1952, Maurice Choisy reclassified it under the current name, Parmotrema perlatum. 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, 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.
The name Parmelia perlata was widely accepted until Hale and Ahti (1986) encountered the designation Lichen chinensis, introduced by Pehr Osbeck in 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.
Impact of the Tokyo Code (1993) 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.
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.
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.
Phylogeny 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. 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.
Common names 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. In North America, vernacular names used for the species include "powdered ruffle lichen", "powdered scatter-rug", and "queen ruffle". 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"; this name later morphed into "pearly parmelia" in some 19th-century British accounts of lichen flora.
Description
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. Individual thalli typically measure up to in diameter. The upper thallus surface is greenish-grey, blue-grey, or yellowish-grey in colour, lacking and either free of spots (), or with few maculae. This species develops soredia, a type of asexual reproductive structure, aiding in its propagation. The of this lichen vary from 1.5 to 10 mm in width, with a wave-like () or ruffled pattern and overlapping () arrangement. The tips and edges of these lobes are generally smooth and round, sometimes notched () or incised, often curling up or inward, revealing the paler brown to black underside adorned with hair-like structures () up to 2.5 mm in length. Rhizines are common on the underside of the thallus, except for a brown border near the edges.
The soredia found in this species are 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 presence of soredia causes the lobe margins to curl back and form soralia. The upper surface of the lichen is typically whitish grey to pale greenish-grey, and can be either smooth or slightly wrinkled, without spots (), featuring scattered, shallow cracks. Isidia are absent in this species.
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 . 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 () of the thallus, with the conidia being thread-like and straight, measuring 6–8 by 1 μm.
Photobiont
The photobiont partner of Parmotrema perlatum is from Trebouxia, a green algal genus belonging to the order Trebouxiales (order Chlorophyta). It has been identified as an undescribed species within a clade containing Trebouxia arboricola.
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.
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.
Chemistry Chemically, Parmotrema perlatum contains atranorin and chloroatranorin, alongside a predominant stictic acid 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. The cortical layer, in contrast, is K+ (yellow), KC−, and P−.