''Punctelia graminicola'' is a species of

foliose Foliose lichen is one of the morphological classes of lichens, which are complex organisms that arise from the symbiotic relationship between fungi and a photosynthetic partner, typically algae. This partnership allows lichen to live in diverse ...

(leafy)

lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.Parmeliaceae The Parmeliaceae is a large and diverse family of Lecanoromycetes. With over 2700 species in 71 genera, it is the largest family of lichen-forming fungi. The most speciose genera in the family are the well-known groups: '' Xanthoparmelia'' ( 822 ...

. It grows on rocks, and, less frequently, on bark in North America, South America, and East Africa. It has a blue-grey

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 ...

measuring up to about , covered with tiny pores called

pseudocyphella Pseudocyphellae (singular ''pseudocyphella'') are structures in lichens that appear as tiny pores on the outer surface (the cortex of the lichen. They are caused when there is a break in the cortex of the lichen, and the medullary hyphae extend t ...

e. Sometimes the lichen forms small lobes that project out from the surface.

Fruiting bodies The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...

are uncommon in this species; if present, they resemble small cups with a brown internal disc measuring in diameter. A lookalike species, '' Punctelia hypoleucites'', is not readily distinguishable from ''Punctelia graminicola'' by appearance or habitat alone; these species can only be reliably differentiated by examining the length of their

conidia A conidium ( ; ), sometimes termed an asexual chlamydospore or chlamydoconidium (), is an asexual, non-motile spore of a fungus. The word ''conidium'' comes from the Ancient Greek word for dust, ('). They are also called mitospores due to the ...

(asexual

spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...

s). First described in 1942 from specimens collected in

New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...

by Brother Arsène Brouard, the

type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...

, stored in

Dunkirk Dunkirk (french: Dunkerque ; vls, label=French Flemish, Duunkerke; nl, Duinkerke(n) ; , ;) is a commune in the department of Nord in northern France.synonymy A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...

with ''

Punctelia subrudecta ''Punctelia subrudecta'' is a species of lichen in the family Parmeliaceae. It was described as a new species by Finnish lichenologist William Nylander as ''Parmelia subrudecta''. Hildur Krog Hildur Krog (22 March 1922 – 25 August 2014) was a ...

''. In 1980, the species became known as ''Parmelia semansiana'' after studies published by

William William is a male given name of Germanic origin.Hanks, Hardcastle and Hodges, ''Oxford Dictionary of First Names'', Oxford University Press, 2nd edition, , p. 276. It became very popular in the English language after the Norman conquest of Engl ...

and Chicita Culberson; a couple of years later it was transferred to the new genus ''

Punctelia ''Punctelia'' is a genus of foliose lichens belonging to the large family Parmeliaceae. The genus, which contains about 50 species, was segregated from genus ''Parmelia'' in 1982. Characteristics that define ''Punctelia'' include the presence of ...

'', created to contain ''

Parmelia Parmelia may refer to: * Parmelia (barque), the vessel that in 1829 transported the first settlers of the British colony of Western Australia * ''Parmelia'' (fungus), a genus of lichens with global distribution * Parmelia, Western Australia Pa ...

''-like species with pseudocyphellae on the thallus surface. Its name was changed again after, in 2001, some of Brouard's collections were rescued from disposal at a local

landfill A landfill site, also known as a tip, dump, rubbish dump, garbage dump, or dumping ground, is a site for the disposal of waste materials. Landfill is the oldest and most common form of waste disposal, although the systematic burial of the waste ...

. Among this material were duplicates of the original collection that used the original

epithet An epithet (, ), also byname, is a descriptive term (word or phrase) known for accompanying or occurring in place of a name and having entered common usage. It has various shades of meaning when applied to seemingly real or fictitious people, di ...

''graminicola''.

Taxonomy

The lichen was first formally described in 1942 as a new species by French lichenologist

Maurice Bouly de Lesdain Maurice Léopold Joseph Bouly de Lesdain (20 September 1869–3 January 1965) was a French botanist and lichenologist. Early life and career Bouly de Lesdain was born in Dunkirk on 20 September 1869. He did his classical studies at the Jesui ...

, as ''Parmelia graminicola''. The type specimens were collected by Brother Arsène Brouard, a

Lasallian french: Frères des Écoles Chrétiennes , image = Signum Fidei.jpg , image_size = 175px , caption = , abbreviation = FSC , nickname = Lasallians , named_after = , formation ...

Catholic The Catholic Church, also known as the Roman Catholic Church, is the largest Christian church, with 1.3 billion baptized Catholics worldwide . It is among the world's oldest and largest international institutions, and has played a ...

monk, in 1935 from two sites near

Las Vegas, New Mexico Las Vegas is a city in and the county seat of San Miguel County, New Mexico, United States. Once two separate municipalities (one a city and the other a town), both were named Las Vegas—West Las Vegas ("Old Town") and East Las Vegas ("New Town" ...

, at an altitude of . The

specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

''graminicola'' suggests a close association with grasses, as ''gramini'' refers to the grass family

Gramineae 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 and ...

, while the ending ''-cola'' indicates a dweller or inhabitant, and is usually used to imply the habitat. However, Bouly de Lesdain noted that the lichen was found growing on

moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...

es and the

spikemoss ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...

Selaginella ''Selaginella'' is the sole genus of vascular plants in the family Selaginellaceae, the spikemosses or lesser clubmosses. This family is distinguished from Lycopodiaceae (the clubmosses) by having scale-leaves bearing a ligule and by having ...

''. Bouly de Lesdain kept this material, and other

collections Collection or Collections may refer to: * Cash collection, the function of an accounts receivable department * Collection (church), money donated by the congregation during a church service * Collection agency, agency to collect cash * Collection ...

sent to him by Brouard, at his private

herbarium A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...

Dunkirk Dunkirk (french: Dunkerque ; vls, label=French Flemish, Duunkerke; nl, Duinkerke(n) ; , ;) is a commune in the department of Nord in northern France. The herbarium was destroyed during the bombing of Dunkerque in

World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposin ...

, and his collections were destroyed. After the loss of the type material, the name ''Parmelia graminicola'' was often later relegated to

synonymy A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...

with ''

''. In his original description of ''Parmelia graminicola'', Bouly de Lesdain had noted its similarity to ''P. subrudecta''. In 1980,

and Chicita Culberson reported their observations on the differences in the length of the

in populations of '' Parmelia hypoleucites'' collected from

Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...

and Mexico. They noted that the long-form conidia morphs (''P. hypoleucites'') grew on bark and had a range restricted to woodlands of the Mexican highlands, while the short-form conidia morphs grew on rocks and were widespread in south-central North America, with few occurrences in regions with the long-form morph. They used this dimorphism to distinguish the short-form morph as a distinct species, ''P. semansiana'', using the larger of Müller's specimens as the type of this new species.

Hildur Krog Hildur Krog (22 March 1922 – 25 August 2014) was a Norwegian botanist. She was born in Modum. She took the dr.philos. degree in 1968 with a thesis on Alaskan lichens, was hired as a curator at the Botanical Museum of Oslo in 1971 and served as ...

transferred ''Parmelia semansiana'' and 21 other ''Parmelia'' species with rounded (''punctate'') pseudocyphellae to the newly

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

genus ''

'' in 1982. In 2001, after Robert Shaw Egan discovered that some of Brouard's collections had been rescued from being disposed at a local (New Mexico)

, the specimens were transferred to

Arizona State University Arizona State University (Arizona State or ASU) is a public research university in the Phoenix metropolitan area. Founded in 1885 by the 13th Arizona Territorial Legislature, ASU is one of the largest public universities by enrollment in the ...

and curated. Among the lichens were specimens collected in 1935 close to the Chimayó dam, thus at the same time and place as some of the specimens on which the species description of ''Parmelia graminicola'' was based; Egan suggested that they are isotypes (i.e. duplicates of the

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

) or isosyntypes (i.e., duplicates of

syntype In biological nomenclature, a syntype is any one of two or more biological types that is listed in a description of a taxon where no holotype was designated. Precise definitions of this and related terms for types have been established as part of ...

s) of the original specimens. ''P. semansiana'' was found to be identical with ''P. graminicola''.

Description

The

of ''Punctelia graminicola'' measures in diameter. It comprises irregularly branched lobes, often crowded or overlapping, measuring wide. The upper thallus surface is blue-grey to greenish-grey, often with brown margins, and usually smooth and without

pruina Pruinescence , or pruinosity, is a "frosted" or dusty-looking coating on top of a surface. It may also be called a pruina (plural: ''pruinae''), from the Latin word for hoarfrost. The adjectival form is pruinose . Entomology In insects, a "blo ...

. Herbarium specimens tend to turn brown.

Pseudocyphella Pseudocyphellae (singular ''pseudocyphella'') are structures in lichens that appear as tiny pores on the outer surface (the cortex of the lichen. They are caused when there is a break in the cortex of the lichen, and the medullary hyphae extend t ...

e on the thallus surface range in number from occasional to abundant. They are circular to irregularly shaped, measuring 0.03–0.6 by 0.03–0.4 mm. The

medulla Medulla or Medullary may refer to: Science * Medulla oblongata, a part of the brain stem * Renal medulla, a part of the kidney * Adrenal medulla, a part of the adrenal gland * Medulla of ovary, a stroma in the center of the ovary * Medulla of th ...

is white.

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 such as

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 ...

soralia 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 o ...

are absent in this species, but sometimes the lichen forms lobules – small lobes (''lacinulae'') on the lobe margins that project out from the surface. Adriano Spielmann and Marcelo Marcelli have noted that ''Punctelia graminicola'' has a broad

species concept The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...

, as it include individuals both with and without lacinulae. The thallus undersurface ranges in color from pale

tan Tan or TAN may refer to: Businesses and organisations * Black and Tans, a nickname for British special constables during the Irish War of Independence. By extension "Tans" can now also colloquially refer to English or British people in general, es ...

to light to medium brown, with sparse, light-colored

rhizine In lichens, rhizines are multicellular root-like structures, arising mostly from the lower surface. A lichen with rhizines is termed rhizinate, while a lichen lacking rhizines is termed erhizinate. Rhizines serve only to anchor the lichen to their ...

s. These rhizines are completely covered by a cortex. The upper

cortex Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...

is paraplenctenchymatous; this means that it is made of a type of tissue in which the

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 are oriented in all directions, analogous to the

parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word π ...

of plants. Depending on the dimensions of the thallus, the upper cortex is between 3 and 7 cells thick. It comprises three distinct layers: the top layer is made of small rounded cells that contact the epicortex; the lower layer is itself organised into two layers of larger, thicker-walled cells with a gelatinous appearance. Covering the upper cortex is a thin epicortex, which is continuous even over the pseudocyphellae. The pseudocyphellae are formed from the inside to outside of the thallus. Hyphae in the medulla are organized in circular groups at specific sites of the thallus; these groups of hyphae push the

alga Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mi ...

l cells towards the upper cortex—rupturing both cortex and epicortex—and exposing the medulla. The size of the pore gradually increases as the cortical cells around the inside perimeter disintegrate and the medullary hyphae grow into the pore area. Punctelia hypoleucites 152719

''Punctelia graminicola'' occasionally makes

apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...

, although they are absent or immature in some individuals. They are cup shaped with a brown disc measuring in diameter. There are often abundant pseudocyphellae on the folded margins of the apothecia. The

ascospore An ascus (; ) is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or s ...

s are more or less spherical to broadly

ellipsoid An ellipsoid is a surface that may be obtained from a sphere by deforming it by means of directional scalings, or more generally, of an affine transformation. An ellipsoid is a quadric surface; that is, a surface that may be defined as the ...

, translucent, and measure 7–14 by 6–9

μm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...

. Depending on the individual lichen,

pycnidia A pycnidium (plural pycnidia) is an asexual fruiting body produced by mitosporic fungi, for instance in the order Sphaeropsidales ( Deuteromycota, Coelomycetes) or order Pleosporales (Ascomycota, Dothideomycetes). It is often spherical or inve ...

can be rare or abundant. They are immersed in the thallus surface, resembling brown to black dots. They produce

that are usually like short, translucent rods and have a length of 5–6 by 1 μm; in rare instances the conidia are hooklike (''unciform''). The expected results of standard

lichen spot test A spot test in lichenology is a spot analysis used to help identify lichens. It is performed by placing a drop of a chemical on different parts of the lichen and noting the colour change (or lack thereof) associated with application of the chemical ...

s for ''Punctelia graminicola'' are K+ (yellow), C−, KC−, P−, and UV− in the upper cortex; the cortex contains

atranorin Atranorin is a chemical substance produced by some species of lichen. It is a secondary metabolite belonging to a group of compounds known as depsides. Atranorin has analgesic, anti-inflammatory, antibacterial, antifungal, cytotoxic, antioxidant, ...

as a minor or trace component, and chloroatranorin in trace amounts. In the medulla, spot test results are K−, KC+ (red), C+ (red), P−, and UV−, indicating the presence of

lecanoric acid Lecanoric acid is a chemical produced by several species of lichen.ubChem - Lecanoric acid"> Lecanoric acid is classified as a polyphenol and a didepside and it functions as an antioxidant. The acid is named after the lichen ''Lecanora''. The acid ...

. ''Punctelia hypoleucites'' is quite similar in appearance, and can be reliably distinguished from ''Punctelia graminicola'' by the length of its conidia (11–12 μm). Unlike ''Punctelia graminicola'', which grows on both rock and bark (the latter much less frequently), ''Punctelia hypoleucites'' only grows on bark.

Habitat and distribution

''Punctelia graminicola'' grows on rocks, or on

es that are on rocks. It has been recorded growing on a wide variety of rocks:

basalt Basalt (; ) is an aphanite, aphanitic (fine-grained) extrusive igneous rock formed from the rapid cooling of low-viscosity lava rich in magnesium and iron (mafic lava) exposed at or very near the planetary surface, surface of a terrestrial ...

conglomerate Conglomerate or conglomeration may refer to: * Conglomerate (company) * Conglomerate (geology) * Conglomerate (mathematics) In popular culture: * The Conglomerate (American group), a production crew and musical group founded by Busta Rhymes ** Co ...

granite Granite () is a coarse-grained (phaneritic) intrusive igneous rock composed mostly of quartz, alkali feldspar, and plagioclase. It forms from magma with a high content of silica and alkali metal oxides that slowly cools and solidifies undergro ...

limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...

rhyolite Rhyolite ( ) is the most silica-rich of volcanic rocks. It is generally glassy or fine-grained (aphanitic) in texture, but may be porphyritic, containing larger mineral crystals (phenocrysts) in an otherwise fine-grained groundmass. The mineral ...

sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...

schist Schist ( ) is a medium-grained metamorphic rock showing pronounced schistosity. This means that the rock is composed of mineral grains easily seen with a low-power hand lens, oriented in such a way that the rock is easily split into thin flakes o ...

, and

volcanic A volcano is a rupture in the crust of a planetary-mass object, such as Earth, that allows hot lava, volcanic ash, and gases to escape from a magma chamber below the surface. On Earth, volcanoes are most often found where tectonic plates a ...

. Much less frequently, it grows on bark. Recorded bark substrates include ''

Fouquieria ''Fouquieria'' is a genus of 11 species of desert plants, the sole genus in the Family (biology), family Fouquieriaceae. The genus includes the ocotillo (''Fouquieria splendens, F. splendens'') and the Boojum tree or cirio (''Fouquieria columnar ...

'', ''

Cupressus ''Cupressus'' is one of several genera of evergreen conifers within the family Cupressaceae that have the common name cypress; for the others, see cypress. It is considered a polyphyletic group. Based on genetic and morphological analysis, the ge ...

'', and ''

Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...

''. In the United States, ''Punctelia graminicola'' occurs in the central US east to the

Appalachians The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...

. It is common in the

Great Plains The Great Plains (french: Grandes Plaines), sometimes simply "the Plains", is a broad expanse of flatland in North America. It is located west of the Mississippi River and east of the Rocky Mountains, much of it covered in prairie, steppe, an ...

region, extending to the southwestern part of the country. The Culbersons note that in the US, the lichen is only commonly found on trees in the centre of its range, while being largely confined to rock in the peripheries of its range. They suggest that the lichen "is primarily adapted to life on rock, extending substrate tolerance to bark only under the most favorable ecological conditions". In Mexico ''Punctelia graminicola'' is widely distributed across the northern and central parts of the country. Its range continues south into Central America. In South America, its distribution includes Argentina and Brazil. In

East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historical ...

(where it was recorded as ''P. semansiana''), it is rare; it is known from

Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...

Kenya ) , national_anthem = "Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...

, and

Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...

Notes

References

{{Taxonbar, from=Q10646746

graminicola ''Graminicola'' is a genus of passerine birds in the family Pellorneidae The jungle babblers are a family, Pellorneidae, of mostly Old World passerine birds belonging to the superfamily Sylvioidea. They are quite diverse in size and coloratio ...

Lichen species Lichens described in 1942 Lichens of Africa Lichens of Mexico Lichens of the United States Lichens of Argentina Lichens of Brazil Taxa named by Maurice Bouly de Lesdain Fungi without expected TNC conservation status