Lobaria Spathulata
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''Lobaria'' 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
foliose lichen 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 ...
s, formerly classified in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Lobariaceae, but now placed in the
Peltigeraceae The Peltigeraceae are a family of lichens in the order Peltigerales. The Peltigeraceae, which contains 15 genera and about 600 species, has recently (2018) been emended to include the families Lobariaceae and Nephromataceae. Many Peltigeraceae sp ...
. They are commonly known as "lung wort" or "lungmoss" as their physical shape somewhat resembles a lung, and their ecological niche is similar to that of moss. ''Lobaria'' are unusual in that they have a three-part
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
, containing a
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
, and an
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 ...
(as other lichens do), but also a
cyanobacterium Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blue ...
that fixes nitrogen.


Taxonomy

''Lobaria'' was originally described as a
section Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sign ...
of the eponymous genus ''Lichen'' by German naturalist
Johann Christian Daniel von Schreber Johann Christian Daniel von Schreber (17 January 1739 in Weißensee, Thuringia – 10 December 1810 in Erlangen), often styled J.C.D. von Schreber, was a German naturalist. Career He was appointed professor of'' materia medica'' at the Univers ...
in 1786. It was proposed as a genus by
Georg Franz Hoffmann Georg Franz Hoffmann was a German botanist and lichenologist. He was born on 13 January 1760 in Marktbreit, Germany, and died on 17 March 1826 in Moscow, Russia. Professional career After graduating from the University of Erlangen in 1786, he w ...
in 1796. The establishment of ''Lobaria'' remained uncertain until
Edvard Vainio Edvard August Vainio (born Edvard Lang; 5 August 185314 May 1929) was a Finnish lichenologist. His early works on the lichens of Lapland, his three-volume monograph on the lichen genus ''Cladonia'', and, in particular, his study of the cla ...
also described it. He divided the genus into two
sections Section, Sectioning or Sectioned may refer to: Arts, entertainment and media * Section (music), a complete, but not independent, musical idea * Section (typography), a subdivision, especially of a chapter, in books and documents ** Section sig ...
based on different morphologies of the mature spore: ''Lobaria'' and ''Ricasolia''. In 2013, the concept of family Lobariaceae was revised with the help of
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, and, in addition to the creation of several new genera, '' Ricasolia'' was promoted to generic status. The family Lobariaceae was synonymized with the
Peltigeraceae The Peltigeraceae are a family of lichens in the order Peltigerales. The Peltigeraceae, which contains 15 genera and about 600 species, has recently (2018) been emended to include the families Lobariaceae and Nephromataceae. Many Peltigeraceae sp ...
in 2018.


Ecology

Lichenicolous fungi A lichenicolous fungus is a parasitic fungus that only lives on lichen as the host. A lichenicolous fungus is not the same as the fungus that is the component of the lichen, which is known as a lichenized fungus. They are most commonly specific to ...
that have been found growing on ''Lobaria'' species include ''
Stigmidium lobariae ''Stigmidium'' is a genus of lichenicolous (lichen-eating) fungi in the family Mycosphaerellaceae. The genus was circumscribed by Italian botanist Vittore Benedetto Antonio Trevisan de Saint-Léon in 1860, with '' Stigmidium schaereri'' assigned ...
'', ''
Calycina alstrupii ''Calycina'' is a genus of fungi within the family Pezizellaceae. The genus contains about 45 species. References External links *Calycina' at Index Fungorum ''Index Fungorum'' is an international project to index all formal names ( scien ...
'', and ''
Abrothallus halei ''Abrothallus halei'' is a species of lichenicolous fungus in the family Abrothallaceae. It was formally described as a new species in 2010 by lichenologists Sergio Pérez-Ortega, Ave Suija, David Leslie Hawksworth, and Rolf Santesson. The t ...
''. A ''Lobaria''-associated
actinobacterium The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to so ...
, ''
Subtercola lobariae ''Subtercola lobariae'' is a Gram-positive and aerobic bacterium from the genus '' Subtercola'' which has been isolated from the lichen '' Lobaria retigera'' from the Jiaozi Snow Mountain in China China, officially the People's Republic ...
'', was isolated from ''L. retigera'' collected from the
Jiaozi Snow Mountain Jiaozi Snow Mountain () is a mountain located in northern central Yunnan, China. The mountain peak lies on the border Dongchuan District and Luquan County in Kunming city-level prefecture. The peak is the highest point in the Eastern Yungui Pl ...
in
Yunnan Province Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the C ...
, China. About a third of the bacteria found colonizing the thallus surface of ''Lobaria pulmonaria'' were found to belong to the
Rhizobiales The ''Hyphomicrobiales'' are an order of Gram-negative Alphaproteobacteria. The rhizobia, which fix nitrogen and are symbiotic with plant roots, appear in several different families. The four families ''Nitrobacteraceae'', ''Hyphomicrobiaceae'' ...
. This order of bacteria is well-known in their role as beneficial partners in plant-microbe interactions. Advantages conferred by the presence of the bacteria include
auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
and
vitamin A vitamin is an organic molecule (or a set of molecules closely related chemically, i.e. vitamers) that is an Nutrient#Essential nutrients, essential micronutrient that an organism needs in small quantities for the proper functioning of its ...
production,
nitrogen fixation Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmo ...
, and
stress Stress may refer to: Science and medicine * Stress (biology), an organism's response to a stressor such as an environmental condition * Stress (linguistics), relative emphasis or prominence given to a syllable in a word, or to a word in a phrase ...
protection. Although the bacteria were most prevalent on the thallus surface, they were shown to be able to penetrate into the interhyphal gelatinous matrix of the upper lichen cortical layer. Occasionally, some bacteria colonize the interior of the fungal hyphae. Hydration traits determine much of a lichen's distribution pattern along a climatic gradient. A study demonstrated that ''Lobaria amplissima'' thalli with external cephalodia need more rain than thalli without, consistent with reports of decreasing frequency of external cephalodia from wet to drier climates. A study using
ecological niche modelling Species distribution modelling (SDM), also known as environmental (or ecological) niche modelling (ENM), habitat modelling, predictive habitat distribution modelling, and range mapping uses computer algorithms to predict the distribution of a spec ...
of occurrence data of three ''Lobaria'' species found in Italy predicts that
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
will impact their distribution range across the country and that there is a high extinction risk resulting from reduction of their range.


Evolutionary history

A fossil impression found in a 12–24  Myr-old
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
deposit from Trinity County in
northern California Northern California (colloquially known as NorCal) is a geographic and cultural region that generally comprises the northern portion of the U.S. state of California. Spanning the state's northernmost 48 counties, its main population centers incl ...
has strong similarities to extant species of ''Lobaria'', particularly ''L. pulmonaria'' and other species with reticulated edges, including '' L. anomala'' and '' L. retigera''. Using a
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleoti ...
-calibrated
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
to obtain a time estimate for ''Lobaria'' yielded a stem age (the time that that
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
descended from a
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
with its
sister clade In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
) of nearly 30 Mya. The evidence suggests that the
paleoclimate Paleoclimatology (British spelling, palaeoclimatology) is the study of climates for which direct measurements were not taken. As instrumental records only span a tiny part of Earth's history, the reconstruction of ancient climate is important to ...
and the closing or opening of the Bering Strait played a significant role in determining the distribution of most ''Lobaria'' species.


Species

*'' Lobaria anomala'' *'' Lobaria anthraspis'' *'' Lobaria discolor'' *''
Lobaria endochroma ''Lobaria'' is a genus of foliose lichens, formerly classified in the family (biology), family Lobariaceae, but now placed in the Peltigeraceae. They are commonly known as "lung wort" or "lungmoss" as their physical shape somewhat resembles a lu ...
'' – New Guinea *'' Lobaria hartmannii'' *'' Lobaria hengduanensis'' – China *''
Lobaria hertelii ''Lobaria hertelii'' is a species of foliose lichen in the family Peltigeraceae. Found in New Guinea, it was formally described as a new species in 2004 by Dutch lichenologist Harrie Sipman Henricus (Harrie) Johannes Maria Sipman (born 1945) ...
'' – New Guinea *'' Lobaria himalayensis'' – India *'' Lobaria irrugulosa'' – China *''
Lobaria isidiophora ''Lobaria'' is a genus of foliose lichens, formerly classified in the family (biology), family Lobariaceae, but now placed in the Peltigeraceae. They are commonly known as "lung wort" or "lungmoss" as their physical shape somewhat resembles a lu ...
'' – Asia *'' Lobaria latilobulata'' – China *'' Lobaria macaronesica''
Macaronesia Macaronesia (Portuguese: ''Macaronésia,'' Spanish: ''Macaronesia'') is a collection of four volcanic archipelagos in the North Atlantic, off the coasts of Africa and Europe. Each archipelago is made up of a number of List of islands in the At ...
*''
Lobaria oregana ''Lobaria oregana,'' also known as lettuce lichen and as Oregon lungwort, is a species of foliose lichen occurring in North American old-growth forests, such as the Hoh Rainforest in Washington state. Taking its common name from its lettuce-lik ...
'' *'' Lobaria orientalis'' – Asia *'' Lobaria plurimiseptata'' *'' Lobaria pseudoretigera'' – New Guinea *''
Lobaria pulmonaria ''Lobaria pulmonaria'' is a large epiphytic lichen consisting of an ascomycete fungus and a green algal partner living together in a symbiotic relationship with a cyanobacterium—a symbiosis involving members of three kingdoms of organisms. ...
'' *'' Lobaria retigera'' *'' Lobaria rhaphispora'' *'' Lobaria spathulata'' Several species formerly classified in ''Lobaria'' have been transferred to other genera in view of modern molecular phylogenetic studies. Examples include ''
Lobaria quercizans ''Lobaria quercizans'' or ''Ricasolia quercizans,'' commonly known as the smooth lungwort, is a macrolichen in the family Peltigeraceae. It forms large, smooth, grey (pale green when wet) growths that often have abundant red-brown apothecia. D ...
'' and '' Lobaria amplissima'' (now in '' Ricasolia''), and ''
Lobaria scrobiculata ''Lobaria scrobiculata'', commonly known as the textured lungwort, is a large foliose, epiphytic lichen in the family Peltigeraceae. Description The thallus of ''L. scrobiculata'' has broad, concave and rounded lobes, rather wider than in ''Lo ...
'' (now in '' Lobarina'').


Chemistry

Retigeranic acid is a sesterterpene compound isolated from ''Lobaria retigera''. The
ethyl acetate Ethyl acetate ( systematically ethyl ethanoate, commonly abbreviated EtOAc, ETAC or EA) is the organic compound with the formula , simplified to . This colorless liquid has a characteristic sweet smell (similar to pear drops) and is used in glues ...
extract of ''Lobaria orientalis'' collected in
Central Vietnam Central Vietnam ( vi, Trung Bộ or ), also known as Middle Vietnam or The Middle, formerly known as by South Vietnam, and Annam under French Indochina, is one of the three geographical regions within Vietnam. The name Trung Bộ was used by ...
led to the isolation of new β-
orcinol Orcinol is an organic compound with the formula CH3C6H3(OH)2. It occurs in many species of lichens including ''Roccella tinctoria'' and ''Lecanora''. Orcinol has been detected in the "toxic glue" of the ant species ''Camponotus saundersi''. It is ...
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, lobarientalones A and B, and several
diphenyl ether Diphenyl ether is the organic compound with the formula ( C6 H5)2 O. It is a colorless solid. This, the simplest diaryl ether, has a variety of niche applications. Synthesis and reactions Diphenyl ether and many of its properties were first rep ...
s, lobariethers A–E.


Uses

Three species of ''Lobaria'' are used as food by ethnic peoples in
Yunnan Province Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the C ...
(China): '' L. isidiophora'', '' L. kurokawae'', and '' L. yoshimurae''.


References

{{Taxonbar, from=Q3501273 Lichen genera Peltigerales genera Taxa described in 1791 Taxa named by Johann Christian Daniel von Schreber