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Parmelia submontana
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Parmelia submontana
Parmelia submontana is a species of corticolous (bark-dwelling), foliose lichen in the family Parmeliaceae. First described in 1987, it is characterised by a loosely attached, greenish-grey body (thallus) reaching diameters of 10–15 cm (4–6 in), with elongated linear lobes and distinctive powdery structures (isidia-like soredia) for reproduction. The species has a complex taxonomic history, having been independently discovered twice – first in Greece in 1832 and later in eastern Bohemia in 1951 – and was long misidentified as related species before being recognised as distinct. It differs from related species in having simple to forked rhizines—root-like holdfasts that attach it to bark—rather than the bushy, bottlebrush-like squarrose rhizines found in similar species, and its blistered rounded soralia that develop from pseudocyphellae.
Though historically considered a Mediterranean and south-central European species, it has been expanding its range northward since the 1990s, particularly in Fennoscandia. The species shows considerable habitat flexibility, growing primarily on deciduous tree bark in areas with high precipitation and humid, sheltered environments, typically near lakes, wetlands, or along coasts. Molecular studies indicate it evolved as a distinct species during the mid-Pleistocene around 1.3 million years ago, and despite appearing morphologically intermediate between some species, it belongs firmly within the Parmelia saxatilis species group. Like many lichens, it produces several secondary metabolites including atranorin, salazinic acid, and consalazinic acid, despite its chemical profile being identical to that of P. saxatilis.
Early lichenologists struggled to classify Parmelia submontana, often treating it as a form or variety of related lichens instead of recognising it as distinct species. The lichen was treated by the American lichenologist Mason Hale in his 1987 monograph on the genus Parmelia. He described Parmelia submontana as a relatively rare European lichen species that had historically been poorly understood. In 1860, William Nylander, who likely examined the type specimen in Paris, regarded it as a narrow-lobed form of P. saxatilis var. sulcata and suggested that it also occurred in North America, although he did not cite any specific specimens. These North American reports were never subsequently confirmed with verified specimens. When the original Bory specimen was examined at the Muséum National d'Histoire Naturelle in Paris, it was found filed under P. saxatilis in the Thuret and Bory herbarium, likely due to Nylander's annotation marking it as "P. saxatilis var. angustifolia Nyl." For instance, Alexander Zahlbruckner placed it under both P. saxatilis and P. sulcata at different times, describing it as P. saxatilis (L.) Ach. var. contorta (Bory) Zahlbr. and P. sulcata Tayl. var. contortoides Zahlbr. The confusion persisted into the 20th century, with many specimens initially misidentified and only recognised later as P. submontana upon re-examination.
Collections made in Austria and Corsica reinforced the distinctiveness of P. submontana. Hale confirmed this based on the species' long, sparsely branched lobes, numerous orbicular soralia, small pseudocyphellae, and sparse, mostly simple (unbranched) rhizines, characteristics initially described by Josef Poelt in 1974. Schindler's 1975 summary provided an up-to-date account of the species' distribution and ecology in Europe, noting that P. submontana is primarily found in Mediterranean and South European montane regions.
Early molecular phylogenetics studies from 2004 examining the relationships within Parmelia s. str. found that P. submontana appeared morphologically intermediate between P. sulcata and P. saxatilis, but molecular evidence showed it was most closely related to P. saxatilis. Analysis of nuclear ITS ribosomal DNA and β-tubulin sequences placed P. submontana within a well-supported clade containing species with simple to furcate rhizines rather than with species having squarrose rhizines like P. sulcata. This rhizine morphology proved to be an important taxonomic character aligning with molecular phylogenetic groupings, helping to resolve some of the historical confusion around the species' relationships. The molecular evidence conclusively demonstrated that despite some morphological similarities, P. submontana belonged firmly within the P. saxatilis group rather than with superficially similar sorediate species like P. sulcata that possess squarrose rhizines.
The species has a peculiar nomenclatural history, having been discovered twice independently. It was first found and described by Bory de Saint-Vincent from Greece in 1832 as Parmelia contorta (type locality: Peloponnese, Taygetus Pass). It was later independently discovered in Eastern Bohemia by Josef Nádvorník, who initially described it under a different name as P. bohemica in 1951. This name was later determined to be illegitimate under Article 53.1 of the International Code of Botanical Nomenclature because P. bohemica Gyeln. (1932) was already in use for a different species (now known as Xanthoparmelia conspersa). To resolve this homonymy, Nádvorník proposed the replacement name P. submontana in 1957, but the name was not validly published under the botanical code. The name was finally validated by Mason Hale in 1987, making him the recognised authority for the species.
Parmelia submontana is a lichen with a thallus that is loosely attached to the bark of trees, often trailing in well-developed specimens. It has a firm, greenish, mineral-grey thallus that can reach a width of 10–15 cm (4–6 in). The lobes are elongated and linear, measuring up to 30 mm in length and 2–5 mm in width. These lobes are only slightly branched and sometimes weakly channeled. The upper surface is shiny and unbroken, ranging from smooth to slightly wrinkled. It features small, round to irregularly shaped pseudocyphellae (tiny openings on the surface), which are located both on the lamina (the main, leafy part of the thallus) and along the margins. These pseudocyphellae, measuring 0.3–1 mm long, quickly develop soralia (powdery patches used for asexual reproduction) that are initially orbicular to linear, often becoming eroded with age. The soredia—tiny grain-like clusters that reproduce the lichen vegetatively—are granular, sometimes appearing almost like isidia (small outgrowths).
The lower surface of the thallus is black and sparsely covered with rhizines (root-like structures), which are simple to sparsely branched and measure 1–1.5 mm in length. The species has simple to furcate rhizines, where secondary branches arise obliquely from the axis, unlike the squarrose rhizines found in some other Parmelia species where secondary branches arise perpendicularly. Apothecia (sexual reproductive structures) are rare in this species. When present, they are round, 1–2 mm in diameter, disc-shaped with a slightly crenulate margin, and have a reddish-brown disc. The spores are colourless, single-celled, measuring 10–11 by 14–15 μm, with thick walls (2 μm). The hymenium (55–70 μm) and excipulum are colourless, with asci 25–55 μm in length.
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Parmelia submontana
Parmelia submontana is a species of corticolous (bark-dwelling), foliose lichen in the family Parmeliaceae. First described in 1987, it is characterised by a loosely attached, greenish-grey body (thallus) reaching diameters of 10–15 cm (4–6 in), with elongated linear lobes and distinctive powdery structures (isidia-like soredia) for reproduction. The species has a complex taxonomic history, having been independently discovered twice – first in Greece in 1832 and later in eastern Bohemia in 1951 – and was long misidentified as related species before being recognised as distinct. It differs from related species in having simple to forked rhizines—root-like holdfasts that attach it to bark—rather than the bushy, bottlebrush-like squarrose rhizines found in similar species, and its blistered rounded soralia that develop from pseudocyphellae.
Though historically considered a Mediterranean and south-central European species, it has been expanding its range northward since the 1990s, particularly in Fennoscandia. The species shows considerable habitat flexibility, growing primarily on deciduous tree bark in areas with high precipitation and humid, sheltered environments, typically near lakes, wetlands, or along coasts. Molecular studies indicate it evolved as a distinct species during the mid-Pleistocene around 1.3 million years ago, and despite appearing morphologically intermediate between some species, it belongs firmly within the Parmelia saxatilis species group. Like many lichens, it produces several secondary metabolites including atranorin, salazinic acid, and consalazinic acid, despite its chemical profile being identical to that of P. saxatilis.
Early lichenologists struggled to classify Parmelia submontana, often treating it as a form or variety of related lichens instead of recognising it as distinct species. The lichen was treated by the American lichenologist Mason Hale in his 1987 monograph on the genus Parmelia. He described Parmelia submontana as a relatively rare European lichen species that had historically been poorly understood. In 1860, William Nylander, who likely examined the type specimen in Paris, regarded it as a narrow-lobed form of P. saxatilis var. sulcata and suggested that it also occurred in North America, although he did not cite any specific specimens. These North American reports were never subsequently confirmed with verified specimens. When the original Bory specimen was examined at the Muséum National d'Histoire Naturelle in Paris, it was found filed under P. saxatilis in the Thuret and Bory herbarium, likely due to Nylander's annotation marking it as "P. saxatilis var. angustifolia Nyl." For instance, Alexander Zahlbruckner placed it under both P. saxatilis and P. sulcata at different times, describing it as P. saxatilis (L.) Ach. var. contorta (Bory) Zahlbr. and P. sulcata Tayl. var. contortoides Zahlbr. The confusion persisted into the 20th century, with many specimens initially misidentified and only recognised later as P. submontana upon re-examination.
Collections made in Austria and Corsica reinforced the distinctiveness of P. submontana. Hale confirmed this based on the species' long, sparsely branched lobes, numerous orbicular soralia, small pseudocyphellae, and sparse, mostly simple (unbranched) rhizines, characteristics initially described by Josef Poelt in 1974. Schindler's 1975 summary provided an up-to-date account of the species' distribution and ecology in Europe, noting that P. submontana is primarily found in Mediterranean and South European montane regions.
Early molecular phylogenetics studies from 2004 examining the relationships within Parmelia s. str. found that P. submontana appeared morphologically intermediate between P. sulcata and P. saxatilis, but molecular evidence showed it was most closely related to P. saxatilis. Analysis of nuclear ITS ribosomal DNA and β-tubulin sequences placed P. submontana within a well-supported clade containing species with simple to furcate rhizines rather than with species having squarrose rhizines like P. sulcata. This rhizine morphology proved to be an important taxonomic character aligning with molecular phylogenetic groupings, helping to resolve some of the historical confusion around the species' relationships. The molecular evidence conclusively demonstrated that despite some morphological similarities, P. submontana belonged firmly within the P. saxatilis group rather than with superficially similar sorediate species like P. sulcata that possess squarrose rhizines.
The species has a peculiar nomenclatural history, having been discovered twice independently. It was first found and described by Bory de Saint-Vincent from Greece in 1832 as Parmelia contorta (type locality: Peloponnese, Taygetus Pass). It was later independently discovered in Eastern Bohemia by Josef Nádvorník, who initially described it under a different name as P. bohemica in 1951. This name was later determined to be illegitimate under Article 53.1 of the International Code of Botanical Nomenclature because P. bohemica Gyeln. (1932) was already in use for a different species (now known as Xanthoparmelia conspersa). To resolve this homonymy, Nádvorník proposed the replacement name P. submontana in 1957, but the name was not validly published under the botanical code. The name was finally validated by Mason Hale in 1987, making him the recognised authority for the species.
Parmelia submontana is a lichen with a thallus that is loosely attached to the bark of trees, often trailing in well-developed specimens. It has a firm, greenish, mineral-grey thallus that can reach a width of 10–15 cm (4–6 in). The lobes are elongated and linear, measuring up to 30 mm in length and 2–5 mm in width. These lobes are only slightly branched and sometimes weakly channeled. The upper surface is shiny and unbroken, ranging from smooth to slightly wrinkled. It features small, round to irregularly shaped pseudocyphellae (tiny openings on the surface), which are located both on the lamina (the main, leafy part of the thallus) and along the margins. These pseudocyphellae, measuring 0.3–1 mm long, quickly develop soralia (powdery patches used for asexual reproduction) that are initially orbicular to linear, often becoming eroded with age. The soredia—tiny grain-like clusters that reproduce the lichen vegetatively—are granular, sometimes appearing almost like isidia (small outgrowths).
The lower surface of the thallus is black and sparsely covered with rhizines (root-like structures), which are simple to sparsely branched and measure 1–1.5 mm in length. The species has simple to furcate rhizines, where secondary branches arise obliquely from the axis, unlike the squarrose rhizines found in some other Parmelia species where secondary branches arise perpendicularly. Apothecia (sexual reproductive structures) are rare in this species. When present, they are round, 1–2 mm in diameter, disc-shaped with a slightly crenulate margin, and have a reddish-brown disc. The spores are colourless, single-celled, measuring 10–11 by 14–15 μm, with thick walls (2 μm). The hymenium (55–70 μm) and excipulum are colourless, with asci 25–55 μm in length.