Bilimbia
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''Bilimbia'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
-forming
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
in the family
Ramalinaceae The Ramalinaceae are a family of lichen-forming fungi in the order Lecanorales. First proposed by Carl Adolph Agardh in 1821, the family now comprises 63 genera and about 750 species. Ramalinaceae lichens exhibit diverse growth forms, includin ...
. The genus forms
crustose Crustose is a Habit (biology), habit of some types of algae and lichens in which the organism grows tightly appressed to a substrate, forming a biological layer. ''Crustose'' adheres very closely to the Substrate (biology), substrates at all poin ...
(crust-like) lichens that appear as whitish-grey to grey patches on calcium-rich soils or
moss Mosses are small, non-vascular plant, non-vascular flowerless plants in the taxonomic phylum, division Bryophyta (, ) ''sensu stricto''. Bryophyta (''sensu lato'', Wilhelm Philippe Schimper, Schimp. 1879) may also refer to the parent group bryo ...
es growing over
alkaline In chemistry, an alkali (; from the Arabic word , ) is a basic salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as a base that dissolves in water. A solution of a soluble base has a pH greater than 7.0. The ...
surfaces. The genus is distinguished by its sessile
apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
(fruiting bodies) that range from light
ochre Ochre ( ; , ), iron ochre, or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colou ...
to black, eight-spored asci, and colourless
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s with multiple cross-walls.
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 ...
studies have confirmed that ''Bilimbia'' forms a well-supported
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
group within the Ramalinaceae, and as of 2025, it contains 24 accepted species.


Taxonomy

Before ''Bilimbia'' was
circumscribed In geometry, a circumscribed circle for a set of points is a circle passing through each of them. Such a circle is said to ''circumscribe'' the points or a polygon formed from them; such a polygon is said to be ''inscribed'' in the circle. * Circum ...
by
Giuseppe De Notaris Giuseppe De Notaris (18 April 1805 – 22 January 1877) was an Italian botanist generally known for his work with cryptogams native to Italy. Biography Born 18 April 1805, in Milan, he studied medicine at the University of Pavia, obtaining his ...
, the name had been used in botany. Reichenbach had attempted to use it for what is now known as the Bilimbi or Cucumber tree (''
Averrhoa bilimbi ''Averrhoa bilimbi'' (commonly known as bilimbi, cucumber tree, or tree sorrel) is a fruit-bearing tree of the genus ''Averrhoa'', family (biology), family Oxalidaceae. It is believed to be originally native to the Maluku Islands of Indonesia but ...
''), but this usage was never validly published under botanical nomenclature rules. The genus ''Bilimbia'' as defined by De Notaris in 1846 included two species, ''B. hexamera'' and ''B. tetramera''. It became widely adopted among lichenologists, particularly in the British Isles where it was used consistently from William Mudd's 1861 flora through Duncan's 1959 guidebook. However, its use declined in the early 1900s when
Alexander Zahlbruckner Alexander Zahlbruckner (31 May 1860, Svätý Jur – 1938, Vienna) was an Austrian- Hungarian botanist who specialized in the study of lichens. Johann Babtist Zahlbruckner, an earlier Austrian botanist, was his grandfather. From 1878 to 1883 ...
transferred the ''Bilimbia'' species into his broader concept of the genus ''
Bacidia ''Bacidia'' is a genus of lichen-forming fungi in the family Ramalinaceae. Taxonomy The genus was circumscribed by Giuseppe De Notaris in 1846. Description ''Bacidia'' is characterised by its crustose (crust-like) growth form. The main body ...
''. Several related genera were proposed during this period of taxonomic uncertainty.
Philipp Maximilian Opiz Philipp (Filip) Maximilian Opiz (5 June 1787 in Čáslav – 20 May 1858 in Prague) was a Czech-German forester and botanist. He made contributions to European botany during the early 19th century. Showing an early interest in botany from child ...
created ''Weitenwebera'' in 1857 specifically to replace ''Bilimbia'' , though this
replacement name In biological nomenclature, a ''nomen novum'' (Latin for "new name"), replacement name (or new replacement name, new substitute name, substitute name) is a replacement scientific name that is created when technical, nomenclatural reasons have mad ...
itself turned out to be a later
homonym In linguistics, homonyms are words which are either; '' homographs''—words that mean different things, but have the same spelling (regardless of pronunciation), or '' homophones''—words that mean different things, but have the same pronunciat ...
. ''Mycobilimbia'' was established in 1889 as a separate genus, notable for having no nomenclatural connections to De Notaris's original ''Bilimbia'' species. Another genus name, ''Probilimbia'', was briefly mentioned by
Edvard August Vainio Edvard August Vainio (born Edvard Lang; 5 August 185314 May 1929) was a Finnish lichenology, lichenologist. His early works on the lichens of Lapland (Finland), Lapland, his three-volume monograph on the lichen genus ''Cladonia'', and, in part ...
in 1899 but is considered
superfluous Superfluous means unnecessary or excessive. It may also refer to: *Superfluous precision, the use of calculated measurements beyond significant figures *''The Diary of a Superfluous Man'', an 1850 novella by Russian author Ivan Turgenev *Superfluo ...
. The taxonomy was later clarified when research showed that Reichenbach's use of ''Bilimbia'' did not constitute valid publication, meaning De Notaris's use remained valid. By 2001, four main European species were recognised: '' B. accedens'', '' B. lobulata'', '' B. microcarpa'', and '' B. sabuletorum'', with the type species ''B. hexamera'' now considered a synonym of ''B. sabuletorum''.
Molecular phylogenetic 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 ...
evidence suggests that the genus forms a well-supported
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
within the
Ramalinaceae The Ramalinaceae are a family of lichen-forming fungi in the order Lecanorales. First proposed by Carl Adolph Agardh in 1821, the family now comprises 63 genera and about 750 species. Ramalinaceae lichens exhibit diverse growth forms, includin ...
. De Notaris did not specify a
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
when he created the genus. Bruce Fink designated ''B. hexamera'' as the type in 1910. However, he used the now-obsolete ''American Code'', and the new nomenclatural rules allow for his typification to be superseded by any later typification, unless the typification has been reaffirmed in the interim. In 1952,
Rolf Santesson Rolf Santesson (1916–2013) was a Swedish lichenologist and university lecturer. He was awarded the Acharius Medal in 1992 for his lifetime contributions to lichenology. Early life and education Santesson was born in 1916 in Trollhättan, Sw ...
set ''B. hexamera'' as the type. In 1984,
Josef Hafellner Josef Hafellner (1951– ) is an Austrian mycologist and lichenologist. He was awarded the Acharius Medal in 2016 for his lifetime contributions to lichenology. Before his retirement, he was a professor at the Karl-Franzens-Universität in Graz ...
redesignated ''B. tetramera'' as the type, a decision that was followed by Timdal in 1991. However, ''B. tetramera'' is now known as ''Mycobilimbia tetramera''. If ''B. tetramera'' is assigned as the type of ''Bilimbia'', then ''Bilimbia'' becomes a synonym of, and the correct name for ''Mycobilimbia'', which is taxonomically unfeasible. For this reason, a proposal was published in 2020 to conserve the name ''Bilimbia'' with ''B. hexamera'' as a conserved type. The proposal was accepted by the Nomenclature Committee for Fungi in 2023.


Description

''Bilimbia'' is a genus of
crustose Crustose is a Habit (biology), habit of some types of algae and lichens in which the organism grows tightly appressed to a substrate, forming a biological layer. ''Crustose'' adheres very closely to the Substrate (biology), substrates at all poin ...
(crust-forming) lichens. These organisms form thin, spreading patches that range in colour from whitish grey to grey. Unlike some lichens, they do not produce
soredia Soredia are common reproduction, reproductive structures of lichens. Lichens asexual reproduction, reproduce asexually by employing simple fragmentation and production of soredia and isidia. Soredia are powdery propagules composed of fungus, fung ...
(powdery reproductive structures) and lack a (a preliminary growth stage). The main body (
thallus Thallus (: 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. A thallus usually names the entir ...
) of the lichen contains green algal cells as its partner. The fungal reproductive structures (
ascomata An ascocarp, or ascoma (: ascomata), is the fruiting body (sporocarp (fungi), sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded ascus, asci, each of which typically contains four to ...
) appear as small, rounded (
apothecia An ascocarp, or ascoma (: ascomata), 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. As ...
) that sit directly on the surface. These apothecia are typically convex and lack a distinct rim, though young specimens may have flat tops with a shallow margin. They range in colour from light
ochre Ochre ( ; , ), iron ochre, or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colou ...
to dark brown or black, and have an undusted () appearance. When viewed under a microscope, the apothecia show several distinct layers. The outer protective layer () curves backward and contains densely packed, radiating fungal threads (
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 o ...
e) held together by a firm gel. This layer often contains dark brown or pinkish-brown pigments that intensify when treated with
potassium hydroxide Potassium hydroxide is an inorganic compound with the formula K OH, and is commonly called caustic potash. Along with sodium hydroxide (NaOH), KOH is a prototypical strong base. It has many industrial and niche applications, most of which utili ...
solution ( K) and turn purple with
nitric acid Nitric acid is an inorganic compound with the formula . It is a highly corrosive mineral acid. The compound is colorless, but samples tend to acquire a yellow cast over time due to decomposition into nitrogen oxide, oxides of nitrogen. Most com ...
(N). The spore-producing layer (
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in oth ...
) is 50–90
micrometre The micrometre (English in the Commonwealth of Nations, Commonwealth English as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American English), also commonly known by the non-SI term micron, is a uni ...
s (μm) tall, occasionally reaching 110 μm, and usually lacks a well-defined top layer. The reproductive cells ( asci) each produce eight
spore In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s and have a distinctive structure that shows complex staining patterns when treated with
iodine Iodine is a chemical element; it has symbol I and atomic number 53. The heaviest of the stable halogens, it exists at standard conditions as a semi-lustrous, non-metallic solid that melts to form a deep violet liquid at , and boils to a vi ...
and potassium hydroxide solution. The spores themselves are colourless and elongated, with 0 to 7 (rarely up to 9) cross-walls (
septa SEPTA, the Southeastern Pennsylvania Transportation Authority, is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly four million people througho ...
), and often have a rough outer coating. The genus may contain a
secondary metabolite Secondary metabolites, also called ''specialised metabolites'', ''secondary products'', or ''natural products'', are organic compounds produced by any lifeform, e.g. bacteria, archaea, fungi, animals, or plants, which are not directly involved ...
, called
zeorin Zeorin is a triterpene with the molecular formula C30H52O2 which occurs in many lichens A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, al ...
, though many species lack any detectable secondary compounds. Various olive-green and brown pigments occur throughout the fungal tissues, though these remain chemically uncharacterised.


Habitat and distribution

''Bilimbia'' species are typically found growing on calcium-rich soils or on mosses that grow over alkaline surfaces. This preference for basic (non-acidic) environments and their association with moss communities is characteristic of the genus.


Species

,
Species Fungorum ''Index Fungorum'' is an international project to index all formal names (Binomial nomenclature, scientific names) in the fungus Kingdom (biology), kingdom. As of 2015, the project is based at the Royal Botanic Gardens, Kew, one of three partn ...
(in the
Catalogue of Life The Catalogue of Life (CoL) is an online database that provides an index of known species of animals, plants, fungi, and microorganisms. It was created in 2001 as a partnership between the global Species 2000 and the American Integrated Taxono ...
) accept 24 species of ''Bilimbia'': * '' Bilimbia caloosensis'' * '' Bilimbia caudata'' * '' Bilimbia cinereoglauca'' * '' Bilimbia corcovadensis'' * '' Bilimbia declinis'' * '' Bilimbia gallica'' * '' Bilimbia granosa'' * '' Bilimbia fiumensis'' * '' Bilimbia jeanjeanii'' * '' Bilimbia lobulata'' * '' Bilimbia microcarpa'' * '' Bilimbia minima'' * '' Bilimbia myriocarpa'' * '' Bilimbia myrtillicola'' * '' Bilimbia novohebridica'' * '' Bilimbia pulchra'' * '' Bilimbia ravenelii'' * '' Bilimbia rubidofusca'' * '' Bilimbia rubricosa'' * '' Bilimbia sabuletorum'' * '' Bilimbia sabulosa'' * '' Bilimbia salevensis'' * '' Bilimbia sibiriensis'' * '' Bilimbia suballinita'' * '' Bilimbia sublecanorina'' * '' Bilimbia tetramera''


References

{{Taxonbar , from1=Q10429579 , from2=Q10591066 , from3=Q105496976 Ramalinaceae Lichen genera Lecanorales genera Taxa described in 1846 Taxa named by Giuseppe De Notaris