''Niebla'' is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

of yellow-green fruticose

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.Brodo, I. M., S. D. Sharnoff, and S. Sharnoff. 2001. Lichens of North America. Yale University Press: New Haven. or more narrowly defined to occur on rocks and soil along the Pacific Coast from

Mendocino County Mendocino County (; ''Mendocino'', Spanish language, Spanish for "of Antonio de Mendoza, Mendoza) is a County (United States), county located on the North Coast (California), North Coast of the U.S. state of California. As of the 2020 United Sta ...

California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...

south to

Baja California Baja California (; 'Lower California'), officially the Free and Sovereign State of Baja California ( es, Estado Libre y Soberano de Baja California), is a state in Mexico. It is the northernmost and westernmost of the 32 federal entities of Mex ...

Sur.Spjut, R. W. 1996. ''Niebla'' and ''Vermilacinia'' (Ramalinaceae) from California and Baja California. Sida Miscellany 14, pp. 1-208. Botanical Research Institute of Texas, Inc., Fort Worth.

Taxonomy

The genus name is a substitute nameRundel, P.W. and P.A. Bowler, 1978. ''Niebla'', a new generic name for ''Desmazieria'' (Ramalinaceae). Mycotaxon 6: 497–499 for ''Desmazieria'', a

homonym In linguistics, homonyms are words which are homographs (words that share the same spelling, regardless of pronunciation), or homophones (equivocal words, that share the same pronunciation, regardless of spelling), or both. Using this definition, ...

created by

Camille Montagne Jean Pierre François Camille Montagne (15 February 1784 – 5 December 1866) was a French military physician and botanist who specialized in the fields of bryology and mycology. He was born in the commune of Vaudoy in the department of Seine-et- ...

in 1852 who recognized only one species in the genus (''D. homalea''); however, the name ''

Desmazeria ''Desmazeria'' is a genus of Mediterranean plants in the grass family, related to ''Catapodium''. The genus is named for John Baptiste Henri Joseph Desmazières, (1796-1862) a French merchant, amateur botanist, and editor of scientific journals. ...

'', although not spelled exactly the same but nevertheless considered to be the same (homonym), had been given to a genus of grass (

Poaceae Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...

) by Barthélemy Dumortier in 1822. The earliest name is the one that must be retained unless the later name is conserved, according to the

International Code of Botanical Nomenclature The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "trad ...

. Additionally, the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

name for ''Niebla'' is ''Niebla homalea'', based on the name given to the one species that had been recognized by Montagne for the genus, not ''Niebla ceruchis''.Sérusiaux, E., P. van den Boom, and D. Ertz. 2010. A two-gene phylogeny shows the lichen genus ''Niebla'' (Lecanorales) is endemic to the New World and does not occur in Macaronesia nor in the Mediterranean basin. Fungal Biology 114: 528–37. The genus ''Niebla'' has also been interpreted broadly to include the genus ''

Vermilacinia ''Vermilacinia'', a genus of lichenized fungi in the family Ramalinaceae, 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 ...

'',Bowler, P. and J. Marsh. 2004. ''Niebla''. ‘Lichen Flora of the Greater Sonoran Desert 2’: 368–380 although both classifications distinguish species by the presence or absence of chondroid strands. In the broad classification, only three species of ''Niebla'' with chondroid strands are recognized under two arbitrary groups of lichen substances based on their reaction to chemical spot tests of which the most useful in ''Niebla'' appears to be ''para''-phenylenediamine (pd); the nine species in the narrow classification that contain

depsidone Depsidones (+ " depside" + "one") are chemical compounds that are sometimes found as secondary metabolites in lichens. They are esters that are both depsides and cyclic ethers. An example is norstictic acid Norstictic acid is a depsidone produc ...

s (pd+) are all placed under ''N. josecuervoi'' in the broad classification. A second group that contains depsides but also includes the acid-deficient chemotype (''N. homaleoides''), collectively treats 32 species under the name ''N. homalea''. The inclusion of the acid-deficient species in ''N. homalea'' disregards the chemotaxonomic attribute of a

pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...

associated with species that contain

salazinic acid Salazinic acid is a depsidone with a lactone ring. It is found in some lichens, and is especially prevalent in ''Parmotrema'' and ''Bulbothrix'', where its presence or absence is often used to help classify species in those genera. History I ...

. Both classifications recognize ''N. isidiaescens''. Eleven other species that are treated in ''

'' are distinguished by thallus morphology.

Molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioche ...

studies on the North American species of ''Niebla'' and ''Vermilacinia'' have been limited and contradictory. An unpublished study mentioned in a 1995 communication indicated that ''Niebla'' was

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

and that by recognizing ''Vermilacinia'' rendered ''Niebla'' monophyletic. A later study—that reported largely on Old World species of ''Ramalina'' and which included collections of ''Niebla'' and ''Vermilacinia'' species—concluded that ''Niebla'' was

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

. However, source material (voucher specimens) cited in the later study for the basis of the DNA analysis of ''Niebla'' was reported to be from California and identified ''N. homalea'' with protocetraric acid and triterpenes. But this

chemotype A chemotype (sometimes chemovar) is a chemically distinct entity in a plant or microorganism, with differences in the composition of the secondary metabolites. Minor genetic and epigenetic changes with little or no effect on morphology or anatomy m ...

had not been known in ''Niebla'', nor are depsidones known to be present in ''N. homalea'', nor are they present in any species of ''Niebla'' in California; protocetraric acid is found only in ''N. pulchribarbara'', which lacks triterpenes, a rare species that occurs in northern Baja California. In the ''Lichen Flora of the Greater Sonoran Desert Region'', the description given for ''N. homalea'' erroneously described the species to have depsidones, including protocetraric acid, which may be confused with salazinic acid when conducting

thin-layer chromatography Thin-layer chromatography (TLC) is a chromatography technique used to separate non-volatile mixtures. Thin-layer chromatography is performed on a sheet of an inert substrate such as glass, plastic, or aluminium foil, which is coated with a t ...

. Salazinic acid is a common lichen substance accessory to triterpenes in species of ''Vermilacinia'', while it may also be noted that the morphology of ''V. laevigata'' is easily confused with that of ''N. homalea''. In view of the controversial classification of this ''Ramalina'' complex of genera, and that many of the species are common along the Pacific coast of North America, it may be helpful to conservationists to have molecular studies be conducted to correlate the different views of the species concepts with their DNA, which should include representative specimens from the entire geographic range of the species before drawing definitive conclusions about their synonymy.

Description

''Niebla'' has 42 species distinguished by

secondary metabolites Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the nor ...

(lichen substances) and by morphology. ''Niebla'' lichen substances fall into two groups. One group has triterpenes, the other lacks triterpenes. Species of ''Niebla'' that contain triterpenes also contain

depside A depside is a type of polyphenolic compound composed of two or more monocyclic aromatic units linked by an ester bond. Depsides are most often found in lichens, but have also been isolated from higher plants, including species of the Ericaceae, L ...

s as major lichen substances, which in a species may be either divaricatic acid or sekikaic acid. The group lacking

triterpene Triterpenes are a class of chemical compounds composed of three terpene units with the molecular formula C30H48; they may also be thought of as consisting of six isoprene units. Animals, plants and fungi all produce triterpenes, including squale ...

s contains

s as major lichen substances— protocetraric acid, hyprotocetraric acid,

, or none of the acids are present in ''N. homaleoides''.

Usnic acid Usnic acid is a naturally occurring dibenzofuran derivative found in several lichen species with the formula C18H16O7. It was first isolated by German scientist W. Knop in 1844 and first synthesized between 1933-1937 by Curd and Robertson. Usnic a ...

, pigments, and other unknown secondary substances are also produced but have little taxonomic value except for one pigment associated with species that produce salazinic acid. The same pigment occurs in the acid-deficient ''N. homaleoides''. The morphological characters employed to distinguish species of ''Niebla'' include the orientation of ridges in the

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.propagule In biology, a propagule is any material that functions in propagating an organism to the next stage in its life cycle, such as by dispersal. The propagule is usually distinct in form from the parent organism. Propagules are produced by organisms ...

s referred to as

soredia Soredia are common reproductive structures of lichens. Lichens reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungal hyphae wrapped around cyanobacteria or gr ...

and

isidia An isidium is a vegetative reproductive structure present in some lichens. Isidia are outgrowths of the thallus surface, and are corticated (i.e., containing the outermost layer of the thallus), usually with a columnar structure, and consisting o ...

. Most species occur in Baja California, Mexico. ''Niebla'' is related to ''

Ramalina ''Ramalina'' is a genus of greenish fruticose lichens that grow in the form of flattened, strap-like branches. Members of the genus are commonly called strap lichensField Guide to California Lichens, Stephen Sharnoff, Yale University Press, 201 ...

'' and ''

'', distinguished from them by the presence of thread-like

hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...

e intertwined into gelatinized cords called chondroid strands. These strands run lengthwise within the central axis (medulla) of the ''Niebla''

thallus Thallus (plural: thalli), from Latinized Greek (), meaning "a green shoot" or "twig", is the vegetative tissue of some organisms in diverse groups such as algae, fungi, some liverworts, lichens, and the Myxogastria. Many of these organisms wer ...

, and are visible to the naked eye when a branch of the thallus is sliced or torn. Chondroid strands also protrude into the cortical surface in various network arrangements in which species are recognized by their particular network structure. Some ''Ramalina'' species in the

Mediterranean Region In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...

have medullary chondroid stands unattached to the medulla; however, they differ from ''Niebla'' in having pale pycnidia instead of black pycnidia, and often contain pseudocyphellae and more than one depside or depsidone accompanied by triterpenes and/or bourgeanic acid. Their classification in ''Ramalina'' is supported by molecular phylogeny data. ''

'' differs from both ''

'' and ''Niebla'' by the absence of chrondroid strands, and by presence of other lichens substances, predominantly terpenes that differ from those in ''Niebla''. Of taxonomic significance is the diterpene (-)-16 α-hydroxykaurane, a rare

secondary metabolite Secondary metabolites, also called specialised metabolites, toxins, secondary products, or natural products, are organic compounds produced by any lifeform, e.g. bacteria, fungi, animals, or plants, which are not directly involved in the norm ...

in lichens. This diterpene is found only in ''Vermilacinia'' within the Ramalinaceae, and also in ''

Roccella Roccella may refer to: * Roccella Valdemone, a municipality in Sicily, Italy * Roccella Ionica (or Roccella Jonica), a municipality in Calabria, Italy * The lichen genus ''Roccella'', known for the species ''Roccella tinctoria ''Roccella tincto ...

'' (Roccellaceae).

Species

The following lists all the species in the genus ''Niebla'' that have been recognized. In bold are those generally accepted. Listed separately are those also treated in the genus ''Vermilacinia''. Species rare or isolated in occurrence are also noted. *'' Niebla arenaria'' *'' Niebla brachyura'' Rare *'' Niebla caespitosa'' *'' Niebla contorta'' *'' Niebla cornea'' *'' Niebla dactylifera'' Rare *'' Niebla dilatata'' *'' Niebla disrupta'' *'' Niebla dissecta'' *'' Niebla eburnea'' *'' Niebla effusa'' *'' Niebla fimbriata'' *'' Niebla flabellata'' *'' Niebla flagelliforma'' *''

Niebla halei ''Niebla halei'' is a fruticose lichen that grows on rocks along the foggy Pacific Coast of northern California on San Bruno MountainSpjut, R. W. 1996. ''Niebla'' and ''Vermilacinia'' (Ramalinaceae) from California and Baja California. Sida Bot ...

'' Rare *''

Niebla homalea ''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 ...

'' *'' Niebla homaleoides'' Rare *'' Niebla infundibula'' Rare *'' Niebla isidiaescens'' *'' Niebla isidiosa'' Rare *'' Niebla josecuervoi'' *'' Niebla juncosa'' var. ''juncosa'' *'' Niebla juncosa'' var. ''spinulifera'' *'' Niebla laminaria'' *'' Niebla limicola'' *'' Niebla lobulata'' *'' Niebla marinii'' *'' Niebla palmeri'' Rare *'' Niebla podetiaforma'' *'' Niebla pulchribarbara'' Rare *'' Niebla ramosissima'' Rare *'' Niebla rugosa'' *'' Niebla siphonoloba'' *'' Niebla sorediata'' Rare *'' Niebla sorocarpia'' *'' Niebla spatulata'' Rare *'' Niebla suffnessii'' *'' Niebla tesselata'' Rare *'' Niebla testudinaria'' *'' Niebla turgida'' *'' Niebla undulata'' *'' Niebla usneoides'' *'' Niebla versiforma'' Rare Species classified in ''Niebla'' also recognized in ''Vermilacinia *'' Niebla cephalota'' *'' Niebla ceruchis'' *'' Niebla cedrosensis'' *''

Niebla ceruchoides Niebla, a Spanish word (from Latin ''nebula'', also meaning) "mist, fog", may refer to : Places and jurisdiction * Niebla, Spain, a town, former bishopric and titular see in Andalusia, southern Spain ** Taifa of Niebla The Taifa of Niebla () ...

'' *'' Niebla combeoides'' *'' Niebla flaccescens'' *'' Niebla granulans''Sipman, H.J.M. 2011. New and notable species of ''Enterographa'', ''Niebla'', and ''Sclerophyton s. lat.'' from coastal Chile. Bibliotheca Lichenologica 106: 297-308. *'' Niebla laevigata'' *'' Niebla polymorpha'' *'' Niebla procera'' *'' Niebla robusta'' *'' Niebla tigrina'' *'' Niebla tuberculata'' Rare

References

External links

*
Information on ''Vermilacinia'' subgenus ''Vermilacin''
{{Authority control Lecanorales genera Ramalinaceae Lichen genera Taxa described in 1978