{{Short description|Family of fungi}} {{Use dmy dates|date=October 2024}} {{Use British English|date=October 2024}} {{bots|deny=Citation bot}} {{Automatic taxobox | image = Saare-rihmsamblik.JPG | image_caption = ''Ramalina fraxinea''—type species of the type genus of the Ramalinaceae | taxon = Ramalinaceae | authority = C.Agardh (1821) | type_genus = ''Ramalina'' | type_genus_authority = Ach. (1809) | subdivision_ranks = Genera | subdivision = | synonyms_ref = <ref name="Frey 2016"/> | synonyms = {{Collapsible list|bullets=on | Bacidiaceae {{au|Walt.Watson (1929)}} | Biatoraceae {{au|A.Massal. ex Stizenb. (1862)}} | Catinariaceae {{au|Hale ex Hafellner (1984)}} | Crocyniaceae {{au|M.Choisy ex Hafellner (1984)}} | Lecaniaceae {{au|Walt.Watson (1929)}} | Megalariaceae {{au|Hafellner (1984)}} | Phyllopsoraceae {{au|Zahlbr. (1905)}} }} }}
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 71 genera and more than 1200 species. Ramalinaceae lichens exhibit diverse growth forms, including crustose, fruticose, squamulose, leprose, and byssoid thalli, and form symbiotic relationships primarily with green algae of the genus ''Trebouxia''. The family is characterised by pale-coloured thalli, apothecia (fruiting bodies) that are typically pale but may darken with age, and ascospores that vary in shape and septation.
Members of the Ramalinaceae are found in a wide range of habitats worldwide, from coastal fog deserts to boreal, temperate, and tropical forests. Some genera, such as ''Namibialina'', ''Vermilacinia'', and ''Niebla'', are endemic to specific coastal desert regions, whilst others like ''Ramalina'' have an almost worldwide distribution. Several species within the family face conservation challenges due to their limited distributions and specific environmental threats, with some being listed as vulnerable or critically endangered on the IUCN Red List.
==Systematics== ===Taxonomy=== The family was proposed by the Swedish botanist Carl Adolph Agardh in 1821.<ref name="Agardh 1821"/> While Agardh initially classified it as an "ordo" (order), he used it in a way that suggested a family ranking, referring to it as "Ramalineae".<ref name="Blanchon et al. 1996"/> According to the nomenclatural authority Index Fungorum, while Agardh initially classified it as an "ordo" (order), he used it in a way that suggested a family ranking, referring to it as "Ramalineae". The first explicit use of Ramalinaceae as a family name came from Antoine Laurent Apollinaire Fée in 1824, but this was not considered valid under Article 32.1(b) of the nomenclature rules, meaning it was not accompanied by a description or {{lichengloss|diagnosis}} or a reference to a previously published description or diagnosis. The first correctly spelled use of the family name Ramalinaceae in accordance with Article 18.4 (i.e., with the ending -''aceae'') is attributed to Watson in a 1929 publication.<ref name="Watson 1929"/><ref name="IF: Ramalinaceae"/>
Early taxonomists proposed various classification schemes for the family. William Nylander (1870) subdivided it based on internal thallus structures,<ref name="Nylander 1870"/> while Edvard Vainio (1890) emphasized the anatomical structure of the cortex and established sections including ''Fistularia'' and ''Myelopoea''.<ref name="Vainio 1920"/> Later, Gustaf Einar Du Rietz (1926) treated the old genera ''Desmazieria'' and ''Euramalina'' as subgenera under ''Ramalina'', further dividing them into sections and subsections based on thallus anatomy.<ref name="Du Rietz 1926"/> However, none of these classification schemes gained full acceptance among modern-day taxonomists.<ref name="Blanchon et al. 1996"/>
In 2018, Sonja Kistenich and colleagues published a large-scale molecular phylogenetic analysis of the family. The study demonstrated five well‐supported clades in the Ramalinaceae; they are named after the largest genera within them, viz. the ''Bacidia''-, ''Biatora''‐, ''Ramalina''‐, ''Rolfidium''‐, and ''Toninia''‐groups. The genera ''Bacidia'', ''Phyllopsora'', ''Physcidia'' and ''Toninia'' were found to be polyphyletic and split into segregates. The study also traced the character evolution of the morphological and ecological nature of the Ramalinaceae ancestor. The ancestor probably arose from moist, temperate forests growing on the bark of trees with a crustose growth form and reproduced mainly by forming apothecia and long, multi-septate spores.<ref name="Kistenich et al. 2018"/>
A 2020 study by Richard Spjut and colleagues provided further insights into the taxonomy of Ramalinaceae, particularly focusing on the fruticose genera. The research revealed that the fruticose genera within Ramalinaceae are not monophyletic (derived from a single ancestor) but form two distinct lineages: 1) ''Ramalina'' + ''Namibialina'' and 2) ''Vermilacinia'' + ''Niebla''. These lineages are nested within accessions of the crustose genus ''Cliostomum''.<ref name="Spjut et al. 2020"/>
The divergence between these two main lineages occurred approximately 48 million years ago. ''Ramalina'' began to spread worldwide around 43 million years ago, whilst its sister genus ''Namibialina'', newly described in this study, radiated later (about 19–20 million years ago) in the coastal deserts of southwest Africa. ''Vermilacinia'' and ''Niebla'', which diverged around 30 million years ago, are primarily found in coastal deserts of the New World.<ref name="Spjut et al. 2020"/>
The study highlighted challenges in delimiting species boundaries within ''Niebla'' and ''Vermilacinia'', indicating that more data are required for a comprehensive understanding. Notably, the taxonomy proposed by Spjut (1996) for ''Niebla'' was not fully corroborated by molecular data, whereas that for saxicolous ''Vermilacinia'' received substantial support. The research also led to the description of new ''Vermilacinia'' species.<ref name="Spjut et al. 2020"/>
Regarding the genus ''Ramalina'', the study examined 50 identified species out of an estimated total of 230. Species lacking secondary metabolites (except usnic acid) were resolved at the base of the phylogenetic tree but did not form a monophyletic group. Some clades corresponded to the production of specific secondary metabolites, although these were not always autapomorphies. The study also resulted in the recognition or resurrection of several ''Ramalina'' species, including ''R. krogiae'' and ''R. lusitanica''.<ref name="Spjut et al. 2020"/>
===Synonymy=== Some genera now classified in the Ramalinaceae were considered by past authors to be distinctive enough to warrant inclusion in their own family. These historical family names are considered synonymous with Ramalinaceae:<ref name="Frey 2016"/>
* Bacidiaceae {{au|Walt.Watson (1929)}} * Biatoraceae {{au|A.Massal. ex Stizenb. (1862)}} * Catinariaceae {{au|Hale ex Hafellner (1984)}} * Crocyniaceae {{au|M.Choisy ex Hafellner (1984)}} * Lecaniaceae {{au|Walt.Watson (1929)}} * Megalariaceae {{au|Hafellner (1984)}} * Phyllopsoraceae {{au|Zahlbr. (1905)}}
===Etymology===
As is standard practice in botanical nomenclature,<ref name="Hawksworth 1974"/> the name Ramalinaceae is based on the name of the type genus, ''Ramalina'', with the ending {{nowrap|''-aceae''}} indicating the rank of family. The genus name, assigned by the mycologist Erik Acharius in 1809,<ref name="Acharius 1809"/> comprises the Latin word {{lang|la|ramus}} meaning {{gloss|branch}} and {{lang|la|ramalis}}, meaning {{gloss|pertaining to a branch or having branches}}, and -''ina'', a suffix that denotes similarity. It refers to the typically fruticose, highly branched thalli characteristic of many ''Ramalina'' species.<ref name="Ulloa & Aguirre-Acosta 2020"/>
==Description==
The Ramalinaceae consists of lichen-forming fungi with a highly varied appearance. The thallus, which is the body of the lichen, can take different forms such as crusty (crustose), bushy (fruticose), scale-like (squamulose), or even granular (leprose) or cottony (byssoid). A few species within this family also grow on other lichens (lichenicolous). The colouration of these lichens tends to be pale, and some species may develop small reproductive structures called isidia or soralia that help with vegetative reproduction. The lichen's {{lichengloss|photobiont}} partner, which is the photosynthetic organism living within the fungus, is of the {{lichengloss|chlorococcoid}} type, meaning it consists of green algae that are spherical or slightly elongated.<ref name="Cannon et al. 2023"/>
Ramalinaceae lichens reproduce [sexually via apothecia (fruiting bodies), which are usually pale and may appear black with age. These apothecia may sit directly on the thallus surface or occasionally have short stalks. The edge of the apothecium (the margin) typically lacks a thallus-like covering, but a structural layer called the {{lichengloss|exciple}} is often present, though it may disappear over time as the lichen matures. The {{lichengloss|disc}} of the apothecium can range from flat to strongly convex.<ref name="Cannon et al. 2023"/>
Within the apothecia, there are microscopic structures called paraphyses, which are unbranched or branched filaments that surround the spore-producing cells. The tips of these paraphyses are often swollen. The asci (spore-producing cells) are typically cylindrical to club-shaped. These asci belong to either the ''Bacidia'' or ''Biatora'' types, which are distinguished by specific staining patterns when exposed to iodine (K/I+), showing a dark blue reaction. Each ascus usually contains eight spores. The {{lichengloss|ascospores}}, which are the fungal reproductive units, can vary widely in shape, from broadly elliptical to thread-like. They may be divided by one or more walls (septate) or lack divisions entirely (aseptate), and they are colourless without a surrounding layer.<ref name="Cannon et al. 2023"/>
Members of this family may produce asexual reproductive structures known as pycnidia, which are embedded in the lichen tissue or sit on the surface. These structures release conidia (asexual spores), which also come in various shapes, sometimes septate.<ref name="Cannon et al. 2023"/>
==Photobiont== Species in the Ramalinaceae form symbiotic relationships with photobionts, primarily green algae from the genus ''Trebouxia''. A 2024 study focusing on the ''Ramalina farinacea'' group identified two main photobiont species: ''Trebouxia jamesii'' and ''T. lynnae''.<ref name="Moya et al. 2024"/> These photobionts show distinct geographical distributions and ecological preferences: ''T. jamesii'' predominates in continental Europe and Mediterranean islands, favouring inland areas, whereas ''T. lynnae'' is more common in Macaronesian archipelagos and coastal regions and is better adapted to warmer, more humid climates.<ref name="Moya et al. 2024"/> Specificity varies within the family, as ''Ramalina farinacea'' can associate with both ''T. jamesii'' and ''T. lynnae'', whereas the Canarian endemic ''R. alisiosae'' shows a strict association with ''T. lynnae''. This flexibility likely contributes to the ecological adaptability of the group.<ref name="Moya et al. 2024"/> Both ''T. jamesii'' and ''T. lynnae'' also partner with lichens from other genera and families beyond Ramalinaceae, underlining their broader ecological role.<ref name="Moya et al. 2024"/>
A 2022 study on the ''Ramalina decipiens'' group found that about 50% of thalli contained multiple algal species, though typically one was dominant, accounting for about 94% of algal cells in mixed thalli.<ref name="Blázquez et al. 2022"/> Island of origin and macroclimate exerted greater influence on photobiont community structure than the particular ''Ramalina'' species, consistent with local photobiont adaptation.<ref name="Blázquez et al. 2022"/> Multiple genetic variants (haplotypes) of ''T. jamesii'' occur, some widespread across Europe and others confined to specific regions, a diversity that may enhance the symbiosis' adaptive potential.<ref name="Moya et al. 2024"/> ''T. jamesii'' tended to favour continental areas, whereas ''T. lynnae'' preferred coastal regions and islands, patterns that may reflect differences in temperature tolerance and salinity adaptation.<ref name="Blázquez et al. 2022"/> The capacity of some Ramalinaceae to switch or adjust photobiont partnerships may facilitate colonisation and persistence across varied habitats. For example, ''Ramalina maderensis'' associates with different photobionts in different parts of its range.<ref name="Blázquez et al. 2022"/> However, a study of the ''R. decipiens'' group found no evidence for trophic niche differentiation among ''Ramalina'' species, suggesting that photobiont association is not a primary driver of speciation in that group.<ref name="Blázquez et al. 2022"/>
==Habitat and distribution==
The Ramalinaceae family exhibits a diverse range of habitats and distributions, with several genera showing highly specialised ecological niches. Three fruticose genera—''Namibialina'', ''Vermilacinia'', and ''Niebla''—are endemic to coastal fog deserts. ''Namibialina'' is found in southwestern Africa, whilst ''Vermilacinia'' occurs along the Pacific coasts of South and North America. ''Niebla'' is restricted to North America. In contrast, ''Ramalina'' has a subcosmopolitan distribution, colonising a wide range of habitats from saxicolous sea-shores to trunks and branches in boreal, temperate, and tropical forests. Many species within these genera are found on coastal rocks in fog deserts, while others are epiphytic. Notably, there is evidence of micro-endemism, particularly in ''Niebla'' and saxicolous ''Vermilacinia'' species, with many taxa having very restricted geographical ranges. The distribution and speciation of these genera have been significantly influenced by fog conditions and climate changes since the Miocene. Some species exhibit disjunct distributions, such as ''Vermilacinia zebrina'', which is found in both North America and Namibia.<ref name="Spjut et al. 2020"/>
==Genera==
In a comprehensive molecular phylogeny of the family in which 6 existing genera were reduced to synonymy, Kistenich and colleagues accepted 39 genera in the Ramalinaceae.<ref name="Kistenich et al. 2018"/> Several genera have since been added, some newly proposed and some resurrected from previously disused names. {{As of|2025|November}}, Species Fungorum (in the Catalogue of Life), accepts more than 1200 species distributed amongst 71 genera in the Ramalinaceae.<ref name="CoL_FJB"/> [[File:Crocynia pyxinoides - Flickr - pellaea.jpg|thumb|right|''Crocynia pyxinoides'']] [[File:Powdery Fog Lichen (982924669).jpg|thumb|right|''Niebla cephalota'']] [[File:Toninia sedifolia 090308.jpg|thumb|right|''Toninia sedifolia'']] [[File:Ramalina darwiniana - Flickr - pellaea (1).jpg|thumb|right|''Ramalina darwiniana'']] *''Aciculopsora'' {{au|Aptroot & Trest (2006)}}<ref name="Aptroot et al. 2006"/> – 3 spp. *''Appressodiscus'' {{au|Aptroot & L.A.Santos (2023)}}<ref name="Aptroot et al. 2023"/> *''Auriculora'' {{au|Kalb (1988)}}<ref name="Kalb 1988"/> – 1 sp. *''Bacidia'' {{au|De Not. (1846)}}<ref name="De Notaris 1846"/> – 230 spp. *''Bacidiopsora'' {{au|Kalb (1988)}} – 6 spp.{{#tag:ref|Molecular phylogenetic analysis has indicated that the type species of ''Bacidiopsora'' is nested in ''Bacidia''.<ref name="Kistenich et al. 2018"/>|group=note}} *''Bacidina'' {{au|Vězda (1991)}}<ref name="Vězda 1990"/> – 12 spp. *''Badimia'' {{au|Vězda (1986)}}<ref name="Vězda 1986"/> – 20 spp. *''Bellicidia'' {{au|Kistenich, Timdal, Bendiksby & Ekman (2018)}}<ref name="Kistenich et al. 2018"/> – 1 sp. *''Biatora'' {{au|Fr. (1819)}}<ref name="Kondratyuk et al. 2019"/> – 42 spp. *''Bibbya'' {{au|J.H.Willis (1956)}}<ref name="Willis 1956"/> – 10 spp. *''Bilimbia'' {{au|De Not. (1846)}}<ref name="De Notaris 1846"/> – 6 spp. *''Catinaria'' {{au|Vain. (1922)}}<ref name="Vainio 1922"/> – 6 spp. *''Cenozosia'' {{au|A.Massal. (1845)}}<ref name="Massalongo 1854"/> – 1 sp. *''Cliomegalaria'' {{au|van den Boom & Alvarado (2019)}}<ref name="van den Boom & Alvarado 2019"/> – 1 sp. *''Cliostomum'' {{au|Fr. (1825)}}<ref name="Fries 1825"/> – 25 spp. *''Compsocladium'' {{au|I.M.Lamb (1956)}}<ref name="Lamb 1956"/> – 2 spp. *''Coppinsidea'' {{au|S.Y.Kondr., Farkas & Lőkös (2019)}}<ref name="Kondratyuk et al. 2019"/> – 2 spp. *''Crocynia'' {{au|(Ach.) A.Massal. (1860)}}<ref name="Massalongo 1860"/> – 5 spp.{{#tag:ref|''Crocynia'' is nested within ''Phyllopsora''; a proposal has been made to conserve the name ''Phyllopsora'' against ''Crocynia''.<ref name="Kistinech et al. 2019"/>|group=note}} *''Crustospathula'' {{au|Aptroot (1998)}}<ref name="Aptroot 1998"/> – 5 spp. *''Echidnocymbium'' {{au|Brusse (1987)}}<ref name="Brusse 1987"/> – 1 spp. *''Eschatogonia'' {{au|Trevis. (1853)}}<ref name="Trevisan de Saint-Léon 1853"/> – 7 spp. *''Heppsora'' {{au|D.D.Awasthi & K.Singh (1977)}}<ref name="Awasthi & Singh 1977"/> – 1 sp. *''Herteliana'' {{au|P.James (1980)}}<ref name="Hawksworth et al. 1980"/> – 4 spp.{{#tag:ref|''Herteliana'' is included in the Cladoniaceae by some recent authors.<ref name="Wijayawardene et al. 2022"/>|group=note}} *''Ivanpisutia'' {{au|S.Y.Kondr., Lőkös & Hur (2015)}}<ref name="Kondratyuk et al. 2015"/> (resurrected) – 2 spp. *''Jarmania'' {{au|Kantvilas (1996)}}<ref name="Kantvilas 1996"/> – 2 spp. *''Kiliasia'' {{au|Hafellner (1984)}}<ref name="Hallefner 1984"/> – 9 spp. *''Krogia'' {{au|Timdal (2002)}}<ref name="Timdal 2002"/> – 7 spp. *''Lecania'' {{au|A.Massal. (1853)}}<ref name="Massalongo 1853"/> – 50 spp. *''Lecaniella'' {{au|Jatta (1889)}}<ref name="Jatta 1889"/> (resurrected) – 2 spp. *''Lithocalla'' {{au|Orange (2020)}}<ref name="Orange 2020"/> – 2 spp. *''Lopezaria'' {{au|Kalb & Hafellner (1990)}}<ref name="Kalb 1990"/> – 2 spp.{{#tag:ref|Kistenich and colleagues (2018) suggest that ''Lopezaria'' is a synonym of ''Megalaria''.<ref name="Kistenich et al. 2018"/>|group=note}} *''Lueckingia'' {{au|Aptroot & Umaña (2006)}}<ref name="Aptroot et al. 2006"/> – 1 sp. *''Megalaria'' {{au|Hafellner (1984)}}<ref name="Hallefner 1984"/> – 46 spp. *''Mycobilimbia'' {{au|Rehm (1980)}}<ref name="Rehm 1889"/> – 5 spp. *''Myelorrhiza'' {{au|Verdon & Elix (1986)}}<ref name="Verdon & Elix 1986"/> – 2 spp. *''Myrionora'' {{au|R.C.Harris (1988)}}<ref name="Harris et al. 1988"/> (resurrected) – 5 spp. *''Namibialina'' {{au|Spjut & Sérus. (2020)}}<ref name="Spjut et al. 2020"/> – 1 sp.<ref name="Spjut et al. 2020"/> *''Niebla'' {{au|Rundel & Bowler (1978)}}<ref name="Rundel & Bowler 1978"/> – 23 spp. *''Parallopsora'' {{au|Kistenich, Timdal & Bendiksby (2018)}}<ref name="Kistenich et al. 2018"/> – 3 spp. *''Phyllopsora'' {{au|Müll.Arg. (1894)}}<ref name="Müller 1894"/> – 75 spp. *''Physcidia'' {{au|Tuck. (1862)}}<ref name="Tuckerman 1862"/> – 10 spp. *''Pseudohepatica'' {{au|P.M.Jørg. (1993)}}<ref name="Jørgensen 1993"/> – 2 spp. *''Pseudolepraria'' {{au|Kukwa, Jabłońska, Kosecka & Guzow-Krzem. (2023)}}<ref name="Kukwa et al. 2023"/> – 1 sp. *''Ramalina'' {{au|Ach. (1809)}} – 230 spp. *''Ramalinopsis'' {{au|(Zahlbr.) Follmann & Huneck (1969)}}<ref name="Follmann & Huneck 1969"/> – 1 sp.{{#tag:ref|Kistenich and colleagues (2018) suggest that ''Ramalinopsis'' can be reduced into synonymy with ''Ramalina''.<ref name="Kistenich et al. 2018"/>|group=note}} *''Rolfidium'' {{au|Moberg (1986)}}<ref name="Moberg 1986"/> – 3 spp. *''Schadonia'' {{au|Körb. (1859)}}<ref name="Körber 1859"/> – 4 spp.{{#tag:ref|The placement of ''Schadonia'' within Ramalinaceae is uncertain, with recent studies suggesting it may belong in Ectolechiaceae or warrant its own family, Schadoniaceae, pending further phylogenetic research.<ref name="Kistenich et al. 2018"/><ref name="Cannon et al. 2024"/>|group=note}} *''Scutula'' {{au|Tul. (1852)}}<ref name="Tulasne 1852"/> – 43 spp. *''Stirtoniella'' {{au|D.J.Galloway, Hafellner & Elix (2005)}}<ref name="Galloway 2005"/> – 1 sp. *''Tamasia'' {{au|Farkas (2023)}}<ref name="Farkas 2023"/> – 1 sp. *''Tasmidella'' {{au|Kantvilas, Hafellner & Elix (1999)}}<ref name="Kantvilas 1999"/> – 1 sp. *''Thalloidima'' {{au|A.Massal. (1852)}}<ref name="Massalongo 1852"/> – 17 spp. *''Thamnolecania'' {{au|(Vain.) Gyeln. (1933)}}<ref name="Gyelnik 1933"/> – 1 sp. *''Tibellia'' {{au|Vězda & Hafellner (1992)}}<ref name="Hallefner & Vězda 1992"/> – 1 sp. *''Toninia'' {{au|A.Massal. (1852)}}<ref name="Massalongo 1852"/> – 85 spp. *''Toniniopsis'' {{au|Frey (1926)}}<ref name="Frey 1926"/> – 7 spp. *''Tylocliostomum'' {{au|van den Boom & Magain (2020)}}<ref name="van den Boom & Magain 2020"/> – 1 sp. *''Vandenboomia'' {{au|S.Y.Kondr. (2019)}}<ref name="Kondratyuk et al. 2019"/> – 2 spp. *''Vermilacinia'' {{au|Spjut & Hale (1995)}}{{#tag:ref|Index Fungorum places ''Vermilacinia'' into synonymy with ''Ramalina''.<ref name="IF: Vermiciliana"/>|group=note}} *''Waynea'' {{au|Moberg (1990)}}<ref name="Moberg 1990"/> – 7 spp. *''Woessia'' {{au|D.Hawksw. & Poelt (1986)}}<ref name="Hawksworth & Poelt 1986"/> – 8 spp. *''Wolseleyidea'' {{au|S.Y.Kondr., Farkas & Lőkös (2019)}}<ref name="Kondratyuk et al. 2019"/> – 6 spp.
==Conservation== [[File:Ramalina menziesii 162611469.jpg|thumb|right|''Ramalina menziesii'', the state lichen of California, is assessed as Least Concern by the IUCN.]] The conservation status of 12 Ramalinaceae species has been assessed for the global IUCN Red List. Several species in the family are facing conservation challenges due to their limited distributions and specific environmental threats.
===Madeira archipelago endemics===
In the Madeira archipelago, particularly on Porto Santo Island, multiple rock-dwelling ''Ramalina'' species are assessed as Vulnerable. ''R. portosantana'',<ref name="IUCN: Ramalina portosantana"/> ''R. erosa'',<ref name="IUCN: Ramalina erosa"/> ''R. timdaliana'',<ref name="IUCN: Ramalina timdaliana"/> ''R. confertula'',<ref name="IUCN: Ramalina confertula"/> and ''R. jamesii'' each have restricted populations, ranging from 500 to 1,000 individuals across one to four locations. These species are primarily threatened by trampling and wildfires, which could swiftly devastate their entire populations.<ref name="IUCN: Ramalina jamesii"/> Additionally, ''R. nematodes'', although more abundant with twelve locations, is considered Near Threatened due to its very restricted area of occupancy and the potential impacts of the same threats.<ref name="IUCN: Ramalina nematodes"/>
===Galapagos endemics===
In the Galapagos Islands, ''Ramalina fragilis'' is the rarest among the four endemic ''Ramalina'' species. Unlike its congeners that thrive on shrubs and trees, ''R. fragilis'' is adapted to rocky substrates. It faces unique threats such as flash floods during El Niño events, erosion, and rising sea levels, which are expected to reduce its population size, area of occupancy, and habitat quality by up to 40% over the next 45 years.<ref name="IUCN: Ramalina fragilis"/>
===California endemics===
''Niebla ramosissima'', found solely on San Nicolas Island in Mediterranean California, is also listed as Vulnerable. Its limited distribution, confined to a single location with an area of occupancy up to 32 km<sup>2</sup>, makes it susceptible to habitat transformation caused by invasive species and climate change, which could alter its natural sea-side low-shrub vegetation.<ref name="IUCN: Niebla ramosissima"/>
===Other species===
''Lecania vermispora'' is known from three locations with a total area of occupancy of 8 km². This species faces threats from livestock grazing and climatic changes, which could lead to its decline and potential extirpation. Consequently, it is listed as Vulnerable under criterion D2.<ref name="IUCN: Lecania vermispora"/>
''Bacidia proposita'' is categorised as Critically Endangered. It is known from only one locality within the municipality of Honda, with an area of occupancy of 4 km². This site has experienced significant habitat loss due to logging and the expansion of urban, industrial, and agricultural frontiers. Despite recent surveys in similar habitats, the species has not been rediscovered.<ref name="IUCN: Bacidia proposita"/>
Conversely, ''Bacidia schweinitzii'' and ''Ramalina menziesii'' are assessed as Least Concern. ''Bacidia schweinitzii'' is widespread across eastern North America with scattered occurrences in eastern Asia, while ''Ramalina menziesii'' is common and locally abundant along the coastal regions of western North America. Neither species currently faces significant threats that would jeopardise their populations.<ref name="IUCN: Bacidia schweinitzii"/><ref name="IUCN: Ramalina menziesii"/>
==Notes== {{Reflist|group=note}}
==References== {{Reflist|colwidth=30em|refs=
<ref name="Acharius 1809">{{cite book |last=Acharius |first=E. |title=Lichenographia Universalis |year=1809 |pages=122, 598 |language=la}}</ref>
<ref name="Agardh 1821">{{cite book |last=Agardh |first=Carl Adolf |year=1821 |title=Aphorismi botanici |volume=v.1-16 |page=93 |publisher=Literis Berlingianis |location=Lundin |url=https://www.biodiversitylibrary.org/page/31116413 |doi=10.5962/bhl.title.44857}}</ref>
<ref name="Aptroot 1998">{{cite journal |last1=Aptroot |first1=André |year=1998 |title=New lichens and lichen records from Papua New Guinea, with the description of ''Crustospathula'', a new genus in the Bacidiaceae |journal=Bryophyte Diversity and Evolution|volume=14 |pages=25–35 |doi=10.11646/bde.14.1.6 |url=http://publikationen.ub.uni-frankfurt.de/files/30355/aptroot_1998_lichen.pdf}}</ref>
<ref name="Aptroot et al. 2006">{{cite journal |author-link1=André Aptroot |last1=Aptroot |first1=A. |last2=Umaña |first2=L. |last3=Chaves |first3=J.L. |last4=Trest |first4=M.T. |year=2006 |title=A first assessment of the Ticolichen biodiversity inventory in Costa Rica: three new squamulose genera (Lecanorales: Ramalinaceae and Pilocarpaceae) |journal=Journal of the Hattori Botanical Laboratory |volume=100 |pages=617–624 |doi=10.11646/phytotaxa.55.1.1 |url=https://www.researchgate.net/publication/286254232}}</ref>
<ref name="Aptroot et al. 2023">{{cite journal |last1=Aptroot |first1=André |last2=dos Santos | first2=Lidiane Alves |last3=Fraga Junior |first3=Carlos Augusto Vidigal |last4=da Silva Cáceres |first4=Marcela Eugenia |title=''Ramalea'' and the new genus ''Appressodiscus'' belong in the Ramalinaceae |journal=The Bryologist |volume=126 |issue=3 |year=2023 |doi=10.1639/0007-2745-126.3.360 |pages=360–366}}</ref>
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<ref name="IUCN: Ramalina erosa">{{cite iucn |last1=Sparrius |first1=L. |last2=Aptroot |first2=A. |last3=Sipman |first3=H. |last4=Pérez-Vargas |first4=I. |last5=Matos |first5=P. |last6=Gerlach |first6=A. |last7=Vervoort |first7=M. |date=2 February 2023 |title=Prime Strap Lichen. ''Ramalina erosa'' |url=https://www.iucnredlist.org/species/70382699/213347134 |access-date=8 October 2024}}</ref>
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<ref name="IUCN: Ramalina timdaliana">{{cite iucn |last=Sparrius |first1=L. |last2=Aptroot |first2=A. |last3=Sipman |first3=H. |last4=Pérez-Vargas |first4=I. |last5=Matos |first5=P. |last6=Gerlach |first6=A. |last7=Vervoort |first7=M. |year=2024 |title=Timdal's Strap Lichen. ''Ramalina timdaliana'' |url=https://www.iucnredlist.org/species/70384543/213347329 |access-date=8 October 2024}}</ref>
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}}
{{Taxonbar |from1=Q3066158 |from2=Q60974821 |from3=Q33137783 |from4=Q28812}}
Category:Ramalinaceae Category:Ascomycota families Category:Lichen families Category:Taxa described in 1821 Category:Taxa named by Carl Adolph Agardh