Vermilacinia

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Vermilacinia
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Ramalinaceae
Genus: Vermilacinia
Spjut & Hale (1995)
Type species
Vermilacinia combeoides
(Nyl.) Spjut & Hale (1995)

Vermilacinia, a genus of lichenized fungi in the family Ramalinaceae, [1] is a yellow-green fruticose type of lichen with about 30 species that grow on rocks, trees, and soil within the fog zone along the Pacific Coast of North America and South America. [2] The genus name refers to the thallus being divided into narrow worm-like (vermis) branches (lacinia);the latter part of the name (lacinia) generally applied in descriptions and taxonomic keys such as exemplified in a key to Sonoran Desert species of Ramalina . [3]

The species of Vermilacinia are classified in two subgenera: subgenus Cylindricaria and subgenus Vermilacinia. Subgenus Cylindricaria, which includes 10+ species (type: Vermilacinia corrugata Spjut) that grow mostly on shrubs, is distinguished by a thin membranous pliable cortex (an outer skin-like layer composed of gelatinized hyphal cells) that expands as the thallus imbibes moisture, usually from fog, and shrinks upon drying (as fog dissipates). Subgenus Vermilacinia, which includes 20 species (type: same as for the genus: Vermilacinia combeoides (Nyl.) Spjut & Hale) that grow on rocks (saxicolous) and soil (terricolous), is distinguished by a relatively thick crusty cortex (like a dry pie crust) overlying a medulla with rigid lattice of hyphae. [4] The firmer cortex of subgenus Vermilacinia undoubtedly adds support to the thallus that allows its branches to grow upright from the substrate.

Niebla and Ramalina are related genera distinguished by the hyphae forming multiple longitudinal gelatinized cords (chondroid stands), within the medulla. In many species of Ramalina they form a network interconnected with the lower cortex and medulla, in contrast to a uniform lattice or a single central cord in the Vermilacinia medulla. [4] Also, both Niebla and Vermilacinia have been included in Ramalina, which is distinguished by the pale pycnidia, the presence of pseudocyphellae, by the absence of the diterpene (−)-16 α-hydroxykaurane, and by the absence of the triterpene zeorin, in most species. [4]

Vermilacinia is also unique in the Ramalinaceae for producing the diterpene (−)-16 α-hydroxykaurane (common in nonlichen fungi) [5] among other lichen substances, notably zeorin and bourgeanic acid; depsidones are frequently present as accessory substances. The diterpene is found in all species of subgenus Vermilacinia and in most species of subgenus Cylindricaria. [2] Exceptions are two North American species endemic to Baja California that only contain zeorin, or appearing deficient in lichen substances, although usnic acid may be present in the cortex (V. corrugata, V. howei). [2]

This diterpene, known also as ceruchdiol [6] or ceruchinol, [7] has been attributed to cause a mildew-like appearance [8] that develops gradually on herbarium specimens. The thallus structure in some specimens collected more than 100 years ago is not discernible, making it difficult to determine whether or not soredia are present, which is necessary for species identification. [2] The mold-like appearance is the result of efflorescence—a chemical change associated with the breakdown of the cortex from which the internal (medulla) hyphae erupt. [2] Efflorescence can be slowed or prevented by storing specimens in a frost-free refrigerator below 40 °F, although there may be a trade-off in that long-term storage causes the cortex to become brittle over time (nine yrs). [9]

Not all species of Vermilacinia that produce the diterpene exhibit efflorescence. The presence or absence of this condition has also been referred to in the literature as "deposit" and "no deposit". For example, Gerd Bendz, Johan Santesson and Carl Wachtmeister reported on two specimens of Ramalina homalea (=Niebla homalea) with the diterpene that were collected in 1874, [6] [7] one with "deposit", the other without "deposit"; however, it was not clear as to how their specimens were identified to species. In this case the taxonomy was probably based on R. Heber Howe. [10] The presence of divaricatic acid noted in the one specimen to have the deposit included reference to Mason E Hale Jr., Lichen Handbook. [11] Although one of the two specimens was probably Vermilacinia laevigata, [2] the chemotypes appear incorrectly reported as generally recognized for the species now known as Niebla homalea, which does not produce the diterpene. [2] The same chemical data for Ramalina homalea were also reported by Gerhard Follmann, [7] which again appear to include misidentifications for other lichen substances, without accounting for type specimens. [2] Additionally, no reference was made to zeorin being present; the reported presence of stictic acid was likely a misidentification for salazinic acid and that of atranorin a misidentification of usnic acid. The mold-like condition (“deposit”) has been associated with the occurrence of an unknown terpene ("T3"), [2] which may be related to bourgeanic acid. [5]

The "Ceruchis-Group", which was defined by Gerhard Follmann [7] by the presence (−)-16 α-hydroxykaurane, included species subsequently classified by Peter Bowler in three groups within the genus Niebla (1) N. homalea, (2) N. combeoides, and (3) N. ceruchis. [12] The first two groups are generally recognized to grow on rocks, the third group (N. ceruchis) on trees and shrubs; however, the type for the species in the “Niebla ceruchis group” is one that grows on sand and rock; [2] the type compares closely to specimens that lack a holdfast, as might be expected for thalli growing on soil. A specimen collected by Charles Darwin [2] was noted by him to have been found at "Iquique, 2,000 to 3,000 ft high, where clouds often hang, lying without adhesion on bare sand ... through the coast mountains, no other plant on the coast for 16 leagues inward" (Taylor Herbarium within the Farlow Herbarium, Harvard University. [4] Vermilacinia ceruchis was described to have a cortex 75–125 microns thick, to lack apothecia, and to contain three characteristic terpenes of the N. combeoides group, [2] [12] described by Peter Bowler to have a thick cortex up to 200+ microns, [12] in contrast to the corticolous lichen species, not more than 60 microns thick. [2] [5] [7] [12] There seems to be no disagreement on the morphological differences between the corticolous and saxicolous groups, only a lack of applying the appropriate types to the names, [2] according to the International Code of Botanical Nomenclature. Despite the continued reports indicating that Niebla ceruchis, which is a synonym of Vermilacinia ceruchis, to occur on shrubs and trees in North America, it does not occur in North America. Vermilacinia ceruchis is endemic to South America, and may be extinct. [9]

The South America species of subgenus Cylindricaria often differ from the North American species by containing methyl 3,5-dichlorolecanorate, also known as tumidulin. This was made in reference to the compound having allegedly been discovered in Ramalina ceruchis var. tumidula (basionym Usnea tumidula); [13] [14] However, the type specimen does not contain this compound. [2] The references to the material from which the compound was isolated include a figure showing thalli growing on a cactus, Eulychnia acida, not on sand or rock. The report of lichen substances being present in species—without reference to type specimens—has led to considerable confusion. [2] Many specimens of Vermilacinia in the United States National Herbarium (Smithsonian Institution, Department of Botany)—annotated by species names in the genus Niebla collected in North America—were erroneously reported to contain methyl 3,5-dichlorolecanorate, which may have been deduced from the previous erroneous reports mentioned above; the compound in question appears to be unknown, T3. [2]

Species

The following are species in the genus Vermilacinia according to their classification in subgenera.

Subgenus Vermilacinia

Subgenus Cylindricaria

Related Research Articles

<i>Niebla</i> (lichen) Genus of lichen

Niebla is a genus of yellow-green fruticose lichens that grow on rocks, trees, and shrubs within the fog zone of coastal North America, or more narrowly defined to occur on rocks and soil along the Pacific Coast from Mendocino County in California south to Baja California Sur.

<i>Niebla homalea</i> Species of lichen

Niebla homalea is a species of fruticose lichen that grows on rocks in foggy areas along the Pacific Coast of North America, from Mendocino County, California south to Bahía de San Quintín on the main peninsula of Baja California, with an isolated occurrence further south on vertical rock faces above Punta Camachos, and other occurrences in the Channel Islands and on Guadalupe Island. The epithet homalea, given by Acharius, suggests it was in regard to the branches appearing flattened.

Vermilacinia acicularis is a fruticose lichen that occurs in the Channel Islands of California. The type specimen was collected from San Clemente Island. Other specimens have been collected from Santa Barbara Island and Anacapa Island.

Vermilacinia cedrosensis is a species of pale fruticose lichen that is endemic to Baja California, Mexico.

Vermilacinia ceruchis is a fruticose lichen found on sand and rock in coastal fog areas of Peru and Chile.

Vermilacinia combeoides is a fruticose lichen found on rocks along the Pacific Coast of North America from Sonoma County, California south to San Quintín, Baja California, and also on Santa Catalina Island and Guadalupe Island.

Vermilacinia laevigata is a fruticose lichen that occurs in the fog zone along the Pacific Coast of the California Floristic Province from Marin County, California to 15 miles south of Misión San Vicente Ferrer in Baja California

<i>Vermilacinia procera</i> Species of lichen

Vermilacinia procera is a fruticose lichen of local occurrences on rocks near the sea along the Pacific Coast from San Francisco California to the Channel Islands, and to Punta Escarpada in Baja California. The species is also reported to occur further south to the Vizcaíno Peninsula and on Cedros Island, but these reports are controversial in view of different interpretations of the species that include V. pumila and V. paleoderma that were not recognized at the time V. procera was described ; for example, a specimen collected on Guadalupe Island by Weber and MCoy (L-3605, COLO that was cited by Phillip Rundel and Peter Bowler in 1994 as belonging to Niebla procera, whereas in a revision of the genus by Richard Spjut in 1996, it was cited as belonging to Vermilacinia paleoderma. Both authorities generally agree to some extent on the description of the species and its geographical range within the California Floristic Province.

Vermilacinia tuberculata is a fruticose lichen known only from Morro Bay along the Pacific Coast of California The epithet tuberculata is a reference to the tuberculate surface of the lichen.

Vermilacinia cephalota is a fruticose lichen usually found on trees, shrubs and wooden fences in the fog regions along the Pacific Coast of North America from southeastern Alaska to the Vizcaíno Peninsula of Baja California.

Vermilacinia cerebra is a fruticose lichen that grows on trees and shrubs in the fog regions along the Pacific Coast of North America from the Channel Islands and mainland California near Los Angeles to southern Baja California, also occurring in South America in the Antofagasta Province of northern Chile. The epithet is in reference to the apical swollen lobes that resemble the cerebrum of the brain.

Vermilacinia corrugata is a fruticose lichen that grows on trees and shrubs in the fog regions along the Pacific Coast of North America from the Channel Islands and San Diego in California to southern Baja California. The epithet is in reference to corrugated cortex of the species.

Vermilacinia flaccescens is a fruticose lichen that grows on cacti and shrubs in the fog regions along the Pacific Coast of South America, Peru, Chile and in the Juan Fernandez Islands The epithet is in reference to the flaccid thallus but some specimens have been interpreted to have rigid branches.

Vermilacinia howei is a fruticose lichen that grows on trees and shrubs in the fog regions along the Pacific Coast of North America in the coastal scrub region of the Channel Islands of California, and around Bahía de San Quintín, Baja California and further south in the Vizcaíno Desert. The epithet is in honor of Reginald Heber Howe, Jr. for his contributions to lichenology, especially acknowledged for providing images of the type (biology) specimens in his revision of the genus Ramalina.

Vermilacinia leonis is a fruticose lichen usually found on branches of shrubs in the fog regions along the Pacific Coast of North America and South America; in North America it is found on the southern half of the main peninsula of Baja California north to the southern coast of the Vizcaíno Peninsula. In South America, it occurs on bushes and rocks in Chile; reported from Colchaqua (Valley) and Santiago The epithet is in regard to absence of the black transverse bands often seen in other species such as V. leopardina, V. tigrina and V. zebrina.

Vermilacinia leopardina is a fruticose lichen usually that grows abundantly on the branches of shrubs in the fog regions along the Pacific Coast of North America, in the Channel Islands and on the mainland of California from Santa Barbara County south to the Vizcaíno Peninsula of Baja California. The species is also reported to occur in Chile, based on a single specimen mounted on a large index card off to one corner with the type (biology) of Usnea tumidula in the center and bottom ; it is possible that the specimen of V. leopardina was from North America and placed on the card for the purpose of making a comparison to the type for Usnea tumidula, which was annotated Ramalina ceruchis var. gracilior Muell.Arg., a name of uncertain status. The epithet, leopardina, is in reference to the black transverse bands and irregularly shaped black spots commonly seen on the thallus branches that obviously imply a similarity to the leopard animal, while also making comparative distinctions to other black banded species: V. tigrina and V. zebrina, obviously to a tiger and zebra, and to V. leonis, obviously a lion, which has no black stripes.

Vermilacinia nylanderi is a fruticose lichen that grows on branches of shrubs in the fog regions along the Pacific Coast of North America in the Channel Islands and in Baja California from near El Rosario south to the Vizcaíno Peninsula The epithet honors William Nylander who published a monograph on the related genus Ramalina in 1870.

Vermilacinia tigrina is a fruticose lichen that grows on branches of shrubs and on earth among cactus and shrubs in the fog regions along the Pacific Coast of North America and South America; in North America found in the Channel Islands and near San Diego, California, and in central and southern Baja California; and in South America from Peru to Chile. The epithet is in reference to the irregular black spots or bands on the thallus branches.

Vermilacinia zebrina is a fruticose lichen that grows on bark of trees and shrubs, occasionally on rocks, in the fog regions along the Pacific Coast of North America, from Puget Sound in northern Washington to near El Rosario in Baja California. The epithet, zebrina, is in reference to the black transverse bands on the thallus branches; however, the species is interpreted to include thalli without black bands.

Niebla testudinaria is a fruticose lichen that grows on rocks along the foggy Pacific Coast of North America, from Marin County, California to just south of Tijuana in Baja California, in the Channel Islands in California, and Isla San Martín and Guadalupe Island in Baja California The epithet, testudinaria given by William Nylander in 1870, is probably in reference to the dilated branch with a reticulated surface, similar to a tortoise shell.

References

  1. ↑ Lumbsch TH, Huhndorf SM. (December 2007). "Outline of Ascomycota – 2007". Myconet. Chicago, USA: The Field Museum, Department of Botany. 13: 1–58. Archived from the original on March 18, 2009.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Spjut, R. W. Niebla and Vermilacinia (Ramalinaceae) from California and Baja California. Sida Miscellany 14
  3. ↑ Kashiwadani, H. and T. H. Nash III. 2004. Ramalina. Lichen Flora of the Greater Sonoran Desert Region 2:440–456
  4. 1 2 3 4 Spjut, R. W. 1995. Vermilacinia (Ramalinaceae, Lecanorales), a new genus of lichens. In: Flechten Follmann; Contr. Lichen in honor of Gerhard Follmann; F. J. A. Daniels, M. Schulz & J. Peine, eds., Koeltz Scientific Books: Koenigstein, pp. 337–351.
  5. 1 2 3 Enzien, M. and L. Margulis. 1988. Niebla ceruchis from Laguna Figueroa: dimorphic spore morphology and secondary compounds localized in pycnidia and apothecia. Microbios 55:75–83.
  6. 1 2 , G., J. Santesson & C. W. Wachtmeister. 1965. Studies on the chemistry of lichens. 20. The chemistry of the Ramalina ceruchis group. Acta Chem. Scand. 19:1185–1187.
  7. 1 2 3 4 5 , Follmann, G. 1966. Eine neue Ramalina-Art uas der Ceruches-Gruppe. Willdenowia 4: 225–233.
  8. ↑ Howe, R.H., Jr. 1913. North American species of the genus Ramalina. The Bryologist 16: 65–74.
  9. 1 2 "Vermilacinia Subgenus Vermilacinia".
  10. ↑ Howe, R.H., Jr. 1913. North American species of the genus Ramalina. The Bryologist 16: 65–74.
  11. ↑ Hale, M. E., Jr. 1961. Lichen Handbook, Washington D.C.
  12. 1 2 3 4 Bowler, P.A. 1981. Cortical diversity in the Ramalinaceae. Can. J. Bot. 59: 437–453.
  13. ↑ Huneck, S. 1966. Die Struktur von Tumidulin, einem neuen chlorhaltigen Depsid. Chem. Ber. 99:1106–1110
  14. ↑ Huneck, S. and G. Follmann. 1965. Zur Chemie chilenischer Flechtern XVI. Über die Inhaltasstoffe von Ramalina ceruchis (ACH.) DE NOT. var. tumidula (TAYL.) NYL Zeitschrift für Naturforschung 20b(6): 611–612.
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