The Cladoniaceae are a family of lichen-forming fungi in the order Lecanorales, comprising about 560 species distributed amongst 18 genera. This family is one of the largest among lichen-forming fungi and is globally distributed, from Arctic tundra to tropical rainforests, favouring humid environments while being intolerant of arid conditions. Molecular phylogenetics has significantly advanced the understanding of their complex taxonomic history, revealing intricate evolutionary relationships and leading to a refined classification. Notable members include reindeer moss and cup lichens of the genus Cladonia, which consist of about 500 species and forms a significant part of the diet for large mammals in taiga and tundra ecosystems.
A distinctive feature of many Cladoniaceae species is their dimorphic thallus: a scaly or crust-like primary form and a fruticose (shrub-like) secondary form known as a podetium or pseudopodetium . These lichens typically grow on soil, decaying wood, or tree trunks, with a few species found on rocks. They form symbiotic associations with green algae, usually from the genus Asterochloris. The family is known for its diverse secondary metabolites—over 70 have been identified—which play roles in species identification and ecological functions such as protection against UV radiation and herbivore deterrence. Genomic studies have uncovered significant variation in mitochondrial DNA among Cladoniaceae species, contributing to the understanding of their evolution and diversity.
Before the term 'lichen' was widely adopted, one of the earliest classification systems for these organisms was developed by the German botanist Johann Jacob Dillenius. In his 1741 system, Cladonia species were placed within the genus Coralloides.[5] In his Species Plantarum, Carl Linnaeus formally described several species that are now classified within the Cladoniaceae. Although he initially placed them in the eponymously named genus Lichen, a dozen of these species are recognised as belonging to the genus Cladonia.[6] Among these was Cladonia pyxidata, a representative of the "pixie cup" or "trumpet" lichens—so named for their distinctive shape,[7] as well as some well-known and widespread Cladonia species: C.coccifera, C.cornuta, C.deformis, C.digitata, C.fimbriata, C.gracilis, C.portentosa, C.rangiferina, C.stellaris, C.subulata, and C.uncialis.[7] These 12 species represent about 14% of the 80 Lichen species Linnaeus described in his seminal 1753 work.[6]
The pixie cup, Cladonia pyxidata, was once of the first lichens to be scientifically described.
The German naturalist Jonathan Carl Zenker formally introduced the Cladoniaceae to science in 1827, with his work appearing in a publication edited by Karl Goebel.[1] Zenker's initial concept of the family included genera such as Baeomyces, Icmadophila, and Stereocaulon, which have since been reclassified into separate families due to advances in taxonomic understanding.[8]William Nylander included 53 Cladonia species worldwide in his 1860 work Synopsis lichenum.[9] When Edvard August Vainio published his three-volume monograph on the Cladoniaceae (Monographia Cladoniarum universalis, 1887,[10] 1894,[11] and 1897[12]), he included 134 species and subspecies. In his circumscription of the family, the genera Pycnothelia, Cladia, and Cladina were included in the genus Cladonia.[13] In recent history, Teuvo Ahti's extensive research, including comprehensive monographs and revisions, has significantly advanced the understanding of Cladoniaceae taxonomy and biogeography.[14]
Cladoniaceae is now one of the largest families of lichen-forming fungi,[15] with about 560 species distributed amongst 18 genera. The type genus is Cladonia, circumscribed by the Irish physician and botanist Patrick Browne in 1756. He included eight species in his new genus. Of their occurrence, he wrote: "All these species are found in great abundance in the mountains of Liguanea: they grow mostly on the ground, among other sorts of moss, but a few ... species chiefly are found upon the decaying trunks of trees."[16]
Etymology and naming
As is standard practice in botanical nomenclature,[17] the name Cladoniaceae is based on the name of the type genus, Cladonia, with the ending -aceae indicating the rank of family. The genus name comprises the Greek word: κλάδος (klādos), meaning "branch", "bud", or "shoot"; and the Latin-ia, a suffix commonly used in Latin to form nouns, particularly in taxonomy to denote genera.[18]
It is well known that reindeer feed on lichens, which has led to the widespread but misleading name "reindeer moss".[19] This common name, along with "reindeer lichen" and "caribou lichen", is typically applied to the ground-dwelling, mat-forming species that were previously classified in the genus Cladina.[20]Cladonia species with cup-shaped structures borne at the tips of vertical stalks (podetia) are often known as "pixie cups". Examples include the "boreal pixie cup" (C.borealis),[21] the "finger pixie cup" (C.digitata),[22] and the "red-fruited pixie cup" (C.pleurota) [23] Additional names alluding to these characteristic structures include the "powdered funnel lichen" (C.cenotea),[24] and the "trumpet lichen" (C.fimbriata).[25] Some names reference the reddish hue of their fruiting structures, such as "British soldiers" (C.cristatella),[26] the "jester lichen" (C.leporina),[27] and the "lipstick powderhorn" (C.macilenta).[28]
Cladogram depicting phylogenetic relationships among selected Cladoniaceae taxa, based on Stenroos and colleagues' 2019 analysis.[29]
Molecular phylogenetics studies have significantly advanced the scientific understanding of relationships within the Cladoniaceae, particularly in the large and diverse genus Cladonia. Early studies in the early 2000s began to reveal the complexity of relationships within the family, challenging traditional morphology-based classifications. More recent comprehensive analyses have identified 13 major clades within Cladonia, providing a framework for understanding the genus's evolution and diversity. These clades, while generally well-supported by molecular data, often lack clear morphological synapomorphies (shared physical characteristics inherited from a common ancestor), highlighting the challenges in Cladonia taxonomy. Some clades, such as clade Erythrocarpae (characterised by red apothecia) and clade Ochroleucae (with pale ochraceous apothecia), do share distinctive features. However, most clades encompass a wide range of morphological and chemical variation.[29]
Within these clades, several species complexes have been identified, such as the Cladonia gracilis group and the Cladonia humilis group. These complexes often include morphologically similar species that are difficult to distinguish based on traditional taxonomic characters. Molecular studies have revealed that many of these complexes contain cryptic diversity, with genetically distinct lineages that are not easily recognisable morphologically.[29]
Focused molecular studies have further refined the understanding of specific groups within Cladoniaceae. For instance, detailed analyses of the Cladonia furcata complex have revealed high levels of homoplasy (the occurrence of similar traits due to convergent evolution rather than shared ancestry) in the morphological characters traditionally used for species delimitation. Similarly, studies on the Cladonia cariosa group and the Cladonia pyxidata group have uncovered previously unrecognised diversity and highlighted the need for integrative approaches combining molecular, morphological, and chemical data in species delimitation. These phylogenetic studies have also shed light on biogeographic patterns within Cladoniaceae. For example, some clades show distinct geographic distributions, such as a group of predominantly African species within clade Perviae, while others have more cosmopolitan distributions. Morphologically distinct genera like Carassea, Pycnothelia, and Metus form a sister clade to Cladonia. These genera share some morphological and chemical traits, such as dimorphic thalli and the production of atranorin. However, their geographic distributions vary, with Carassea being endemic to Brazil, while Pycnothelia has a bipolar distribution (i.e., found in the high latitudes of both hemispheres) and Metus is found primarily in Australasia.[29]
Genomics
Genomic studies have revealed significant variation in the size and structure of mitochondrial DNA in Cladoniaceae, uncovering significant variation in genome size and structure. Within the genus Cladonia, mitochondrial genomes span from approximately 45,000 to 66,000 base pairs, consistently containing respiratory genes and ribosomal RNA regions across all examined species. Crustose lichens tend to have smaller mitochondrial genomes compared to their fruticose and foliose counterparts. The genomes often harbour homing endonuclease genes, which may influence genomic evolution. Notably, substantial intraspecific variation has been observed, particularly in widespread species such as C.rangiferina and C.submitis. Unlike some other symbiotic organisms, Cladoniaceae do not exhibit mitochondrial genome reduction, suggesting complex evolutionary dynamics.[30]
In 2018, Ekaphan Kraichak and colleagues used a technique called temporal banding to reorganise the Lecanoromycetes, proposing a revised system of classification based on correlating taxonomic rank with geological (evolutionary) age. They synonymised the families Squamarinaceae and Stereocaulaceae with the Cladoniaceae, resulting in a large increase in the number of genera and species.[37] The Squamarinaceae had already been included in the Cladoniaceae by previous authors.[15] Although this reorganisation has been used in some later publications,[2] the folding of the Stereocaulaceae into the Cladoniaceae was not accepted in a recent analysis. Robert Lücking highlighted that merging the two families under the name Cladoniaceae is not permissible without a formal conservation proposal because Stereocaulaceae, established in 1826, predates Cladoniaceae, which was established in 1827. According to the rules of botanical nomenclature, the earliest validly published name has priority and must be used when two families are combined unless an exception is granted. This means that if the families were merged without a conservation proposal, the combined family would have to be named Stereocaulaceae due to its earlier establishment.[38] In a 2021 treatment of the British and Irish Cladoniaceae, the authors also keep these families separate, noting "both families are monophyletic and easily distinguishable on both morphological and molecular terms".[39]
Description
Cladonia coccifera has a two-part (dimorphic) growth structure
The t points to the leaf-like scales (squamules) of the ground-hugging primary thallus.
The upright secondary thallus, with stalks (podetia) topped by cup-shaped structures (apothecia). The red slime on the cup rims contains asexual reproductive spores (conidia).
The thallus (body) of Cladoniaceae lichens often have a unique structure known as cladoniiform growth, where two distinct forms develop simultaneously within the same organism.[40] The primary thallus , which can be short-lived ( evanescent ) or long-lasting, grows close to the surface and may appear crusty, leafy, or scaly. From this base, a secondary thallus typically grows upright, bearing the reproductive structures (ascomata).[41] This secondary growth can range from a few millimetres to over 25 centimetres (10 inches) in height.[39] Due to this dual structure, the overall appearance of these lichens can be either fruticose (bushy) or foliose (leafy). However, some species in this family do not develop either the primary thallus or the upright structures, showing variations within the group.[29] As for vegetative propagules, isidia occur rarely in this family, whereas soredia are common.[39]
The ascomata are in the form of an apothecium, and are biatorine , meaning they are of the lecideine type – light in colour and soft in consistency. They often have a reduced margin.[41] Their colour is typically dark brown (sometimes pale brown), red, ochraceous, or black.[29] The hamathecium (referring to all hyphae between the asci in the hymenium) consists of sparsely branched paraphyses, and is amyloid, indicating that stains blue to blue-black with iodine-based reagents. The asci (spore-bearing cells) are somewhat fissitunicate , meaning they have two layers that separate during ascus dehiscence. The ascus structure consists of an apical dome and a tube (both of which are amyloid), which is cylindrical to clavate (club-shaped). Ascospores number eight per ascus, and they are usually non-septate (lacking internal partitions), ellipsoid to more or less spherical in shape, hyaline (translucent), and non-amyloid.[41] Except for a few genera that produce septate ascospores (Calathaspis, Pycnothelia and Pilophorus), the hymenium does not generally have characters that are useful in taxonomy.[29] The conidiomata (asexual fruiting bodies) are in the form of pycnidia; the conidia (asexual spores) are non-septate, usually filiform (thread-like), and hyaline.[41]
Chemistry
In the Cladoniaceae, over 70 different secondary metabolites (lichen products), primarily polyphenols, have been identified. These substances crucial roles in species identification and have ecological functions such as protection against UV radiation and deterrence of herbivores. To observe the fluorescence of certain depsides and depsidones, ultraviolet light is directly applied to the specimens. Traditional colour spot tests with reagents like KOH can be ineffective at low concentrations. Consequently, thin-layer chromatography, a laboratory technique used to separate chemical compounds, is essential for detecting and identifying lichen substances that are present in low concentrations.[42]Chemotaxonomy is an important aspect in the identification of Gray's pixie cup (Cladonia grayi): when lit by a UV light, it produces a light blue fluorescence.[43]
Photobionts
The symbiotic algal partner (photobiont) of most Cladoniaceae taxa are unicellular green algae, usually in the genus Asterochloris, but occasionally in the genus Chlorella;[42] both of these genera are in the class Trebouxiophyceae. Eleven species of Asterochloris have found to be associated with genus Cladonia; the algal genus – one of the most common lichen symbionts – occurs in the thalli of more than 20 lichen genera.[44] The most common photobionts in this genus that associate with Cladonia are A.glomerata, A.italiana, and A.mediterranea, with some lineages showing dominance in one or several climatic regions.[45] In contrast, Myrmecia was shown to be the main photobiont for the Mediterranean species Cladonia subturgida.[46] The algal genus Trebouxia, a common lichen photobiont, has not been recorded associating with the Cladoniaceae.[29] Some Pilophorus species form tripartite associations, involving the fungus, green algae, and cyanobacteria within gall-like structures called cephalodia.[42] The cyanobacterial genera Nostoc and Stigonema are involved in these tripartite associations.[29] In a study of several Cladonia lichens collected from Southern Finland, the associated microbial community, which was found to be consistent amongst the different species, consisted largely of Alphaproteobacteria and Acidobacteriota.[47]
Development
The development of several Cladoniaceae genera have been studied in detail,[48][49][50][51] although the interpretation of results has sometimes been controversial.[29] For example, two 1970 studies by Hans Jahns explored the development of fruiting bodies in Cladonia, significant for understanding the taxonomy and phylogeny within the Cladoniaceae. This work revealed two distinct ontogenetic types based on the formation of generative tissue and its role in developing the characteristic podetium of Cladonia species. This generative tissue, originating in the thallus horizontalis and growing vertically, is crucial for forming the podetium. The study identified variability within species, showing that different species can have more than one ontogenetic type, challenging previous assumptions about the uniformity of development patterns within the genus. This demonstrated variability had implications for the taxonomy and phylogenetic relationships within Cladoniaceae, suggesting a complex evolution of reproductive structures that did not strictly align with previously conceived taxonomic groupings.[52][53]
Cladoniaceae species begin development with the formation of a prothallus – a fungal layer upon which an alga-containing thallus will develop. It comprises the hyphae from the germination of an ascospore. After the protothallus contacts the alga, lichenisation begins with the development of small squamules (scale-like thallus segments) that make up the primary thallus, which is squamulose (scaly) or crustose (crustose-like). The secondary thallus consists of vertical structures that are shrubby and hollow, although they can be solid in rare cases. If these structures are made of generative tissue, they are called podetia; when they are made of vegetative tissue, they are called pseudopodetia. The morphology of these structures determines to a large part the taxonomy of the Cladoniaceae, which can range from simple to complex branching patterns.[42]Cladonia minisaxicola, found in the mountains of Bahia (Brazil) is the only species in that large genus that is completely crustose and does not develop podetia.[54]
Cladonia coniocraea (left) has subulate podetia, C.fimbriata (middle) features cup-shaped scyphi, commonly called "pixie cups", and C.cervicornis (right) has a more complex branching pattern.
The tips of the podetia have a wide range of morphology in the Cladoniaceae. They can be straight, tapering from a wide base to a point (called subulate ), or flaring on cup-shaped scyphi . The scyphi are sometimes closed, or have a central perforation, forming structures called funnels.[42] The podetia are slow-growing, with an annual growth rate generally ranging from 1 to 15mm.[55]
Branching in the Cladoniaceae occurs on the podetium, driven by the growth patterns of fungal meristem tissue at its tip. There are two main branching patterns: in one, branches emerge later from a large meristem that changes shape, while in the other, smaller meristems split early but keep their shape. These growth patterns help scientists understand the evolutionary relationships within Cladoniaceae.[56] A shift from isotropous growth (uniform in all directions) to anisotropous growth (different in various directions) allows for more flexibility in development. This transition, from symmetrical to more irregular growth, may signal evolutionary adaptations. Despite these changes, the branching processes remain highly consistent even among species in the family that are not closely related.[57]
Genera
After more than a century of discovery and research, including recent advances in understanding revealed by molecular phylogenetics studies, the Cladoniaceae encompass 18 genera and more than 500 species.[15][39] In terms of species diversity, the Cladoniaceae stood as the tenth-largest lichen-forming fungal family by 2017.[15] This is a list of the genera contained within the Cladoniaceae, based on the Catalogue of Life; this includes taxa formerly classified in the Squamarinaceae, but does not include the Stereocaulaceae. Following the genus name is the taxonomic authority, year of publication, and the number of species:
Cladoniaceae species have been recorded growing in many habitats and on a diversity of substrates, including soil, tree trunks, and rotten wood.[29] In a few cases, Cladoniaceae can grow on rocks, such as Cladonia salmonea which grows on the rock faces of vertical cliffs, or Cladonia pyxidata, which can grow on thin soil on rocks.[74] Cladoniaceae species are generally absent from arid environments due to their preference for humid conditions. The range of their habitats includes boreal forests, bogs, temperate forests, the tundra of the Arctic and Antarctic, man-made habitats (e.g. roadsides), tropical highlands, and the sandy tropical lowlands of the Amazon rainforest.[29]
In his 2000 monograph on the Cladoniaceae of the Neotropical realm, Ahti included 184 species in 4 genera, and showed that South America is a hotspot of biodiversity for genus Clanodia.[14] Bioclimatic variables significantly influence the distribution of Cladoniaceae species richness in the Neotropics, particularly under conditions of low precipitation and temperature, and high climatic variability. Areas with stable climates and higher temperatures and precipitation tend to support greater species richness.[75] Twenty-six Cladoniaceae species (25 Cladonia and 1 Cladia) are known to occur in the Galápagos Islands. There, some species form mats on lava flows that have developed little soil.[76] A 2013 monograph of Northern European Cladoniaceae treated 100 species (95 Cladonia, 4 Pilophorus, and the monotypic genus Pycnothelia).[77] In the 2021 key to lichen species in Italy, 86 Cladoniaceae are included.[78] In Bulgaria, 55 species in two genera were reported in 2022.[79] In a study of the lichen biodiversity in Kazakhstan's Burabay National Park, the Cladoniaceae made up about 30 percent of the species diversity.[80]
In western North America, the Coast Mountains of British Columbia act as a key phytogeographic barrier. This results in distinct oceanic and continental taxa groupings on either side. The research also suggests that the southern boundaries of certain species may be determined more by historical rather than purely ecological factors, indicating possible range expansions. The highest diversity of Cladonia species is found in British Columbia between 52°N and 56°N, an area that was covered by glaciers until about 10,000 to 13,000 years ago. The Cladoniaceae biodiversity in this region represents the richest assemblage of the family in western North America. Species diversity declines sharply south of 52°N, with a loss of three to five taxa for each degree of latitude.[81]
The glacial history of the region has played a crucial role in shaping the current distribution of Cladoniaceae. During the Pleistocene, most species likely survived in areas south of the Cordilleran ice sheet, with some persisting in nunataks, arctic regions, or small coastal refugia. This glacial legacy is still evident in the family's current distribution patterns. Cladoniaceae show a preference for specific habitats, with greater floristic and chemical diversity observed in humid areas and lower forested elevations compared to arid regions and alpine zones. This suggests that many species in the family are adapted to environments with relatively short periods of desiccation. The post-glacial period has seen significant changes in the distribution of Cladoniaceae. Many species that likely existed in Washington, Oregon, and California during the Pleistocene are now absent from these areas. This change is thought to be a result of climate shifts since deglaciation, particularly an increase in summer moisture deficits. While most Cladoniaceae species have reached a stable distribution, some are still in flux. Species like Cladina stellaris and C.trassii appear to be continuing their southward expansion from northern glacial refugia.[81]
Cladonia appalachiensis (endangered, 2020) growing on high-elevation Anakeesta Knob rock, faces threats from visitor disruption and changes in cloud cover and humidity.[83] The species is particularly vulnerable due to its restricted range and specific habitat requirements.[29] The main threats to Cladonia perforata (endangered, 2003) include habitat loss, hurricanes, and improper fire management, with a single natural event potentially causing substantial subpopulation reduction.[84]
Cladonia submitis (endangered, 2020) is primarily threatened by habitat loss and degradation due to land development, particularly around metropolitan areas. Climate change also poses significant risks through altered fire regimes and sea level rise, affecting its pine barren and sand dune habitats.[85] The species' limited distribution and specific habitat requirements make it particularly susceptible to these threats.[29]Pilophorus fibula (endangered, 2020) is threatened by habitat loss, alteration of hydrological regimes, recreational damage, and declining water quality.[86] The species is found in a limited number of locations, making it vulnerable to local extinctions.[29]
Gymnoderma insulare (endangered, 2014), primarily found in old-growth forests in Japan and Taiwan, faces threats from natural hazards like typhoons and is affected by the decline of its tree hosts, Cryptomeria japonica and Chamaecyparis obtusa.[87] The species' dependence on specific host trees and old-growth forest conditions makes it particularly vulnerable to forest degradation and climate change impacts.[29]
Cladonia lichens, particularly the "reindeer lichens" such as C.stellaris, C.rangiferina, and C.arbuscula, are a critical winter food source for reindeer (caribou) in northern boreal and arctic regions.[89] In some areas, these lichens can constitute up to 80% of reindeer winter diet.[90] The average annual linear growth rate of these reindeer lichens is about 5mm per year, accounting for their slow recovery rate after grazing or disturbance.[91] Satellite-based studies have shown a significant decline in caribou lichen cover across large areas of Eastern Canada over the past three decades, likely due to factors such as climate change-induced shrub encroachment, increased wildfire frequency, and grazing pressure, which could have serious implications for caribou populations and ecosystem dynamics.[92] The abundance of lichen-rich forests has significant economic implications for reindeer husbandry, particularly for indigenous Sámi people in Fennoscandia.[93] However, lichen-dominated forests have declined significantly in recent decades due to factors such as intensive forestry practices, overgrazing, and fire suppression.[93] This decline has prompted efforts to restore lichen habitats, including experimental transplantation of lichen fragments to accelerate recovery after disturbances like forest fires.[94]
Five Cladoniaceae species are depicted in Haeckel's 1904 lichen lithograph.
Some Cladoniaceae species are exploited in a profitable export business for decorative uses,[95] with demand reaching thousands of kilograms. In Europe, Cladonia stellaris is used ornamentally in wreaths, floral decorations and architectural models. From 1970–1975, an average of nearly 3000 metric tonnes were exported each year from Finland, Norway and Sweden; most of these exports (about 80%) went to West Germany.[96] Fruticose Cladonia species, often dyed green and glycerol-treated for flexibility, are common in model train displays as miniature trees and shrubs.[97] The product commercially sold as "ball moss" or "Icelandic moss" in hobby and craft stores is often Cladonia stellaris and other similar species, often dyed in various colours.[98] In Sweden, Cladonia lichens were used historically as a partial insulation for storm windows.[99]
The complex net-like structures of the Australasian lichen Pulchrocladia retipora have been described as "of considerable beauty resembling lace or coral", and have been utilised in floral and architectural design.[100] This species' branches, characterised by its numerous small holes, exemplify nature's efficient use of latticework structures. This design, widely used in construction for structures such as transmission towers or bridges, allows the organism to maintain structural integrity while minimising the amount of biological material used in its construction.[101] The unique lichen architectures of five Cladoniaceae species are depicted in Ernst Haeckel's well-known and widely reproduced lichen-themed lithograph in his 1904 work Kunstformen der Nature (The art forms of nature). According to the lichenologists Robert Lücking and Toby Spribille, "the Cladonia growth form continues to be one of the most widely recognized lichen architectures, with their basal scales and erect, often trumpet-shaped podetia".[102]
Related Research Articles
Cladonia asahinae, the pixie cup lichen or Asahina's cup lichen, is a species of cup lichen in the family Cladoniaceae. C. asahinae occurs in Europe, North America, southern South America, and the Antarctic. It typically grows in high moisture environments in soil rich in humus or on dead wood.
Cladia is a genus of lichen-forming fungi in the family Cladoniaceae. Cladia species have a crustose or squamulose (scaly) primary thallus and a fruticose, secondary thallus, often referred to as pseudopodetium. The type species of the genus, Cladia aggregata, is widely distributed, occurring in South America, South Africa, Australasia and South-East Asia to southern Japan and India. Most of the other species are found in the Southern Hemisphere.
Cetradonia is a lichen genus in the family Cladoniaceae. A monotypic genus, Cetradonia contains the single species Cetradonia linearis. The genus was circumscribed in 2002 by Jiang-Chun Wei and Teuvo Ahti. The genus was once placed in the family Cetradoniaceae until that family was subsumed into the Cladoniaceae in 2006.
Carassea is a genus of lichenized fungi in the family Cladoniaceae. The genus was circumscribed in 2002 by Finnish lichenologist Soili Stenroos. A monotypic genus, Carassea contains the single species Carassea connexa. This species, originally named Cladonia connexa, was first documented by Edvard August Wainio, who collected specimens from Minas Gerais, Brazil, in 1885, and published a description of the species in the first of his three-volume Monographia Cladoniarum universalis. The Cladoniaceae genera most closely related to Carassea include Pycnothelia and Metus.
Pilophorus acicularis, commonly known as the nail lichen or the devil's matchstick lichen, is a species of matchstick lichen in the family Cladoniaceae.
Teuvo ("Ted") Tapio Ahti is a Finnish botanist and lichenologist. He has had a long career at the University of Helsinki that started in 1963, and then following his retirement in 1997, at the Botanical Museum of the Finnish Museum of Natural History. Known as a specialist of the lichen family Cladoniaceae, Ahti has published more than 280 scientific publications. A Festschrift was dedicated to him in 1994, and in 2000 he was awarded the prestigious Acharius Medal for lifetime contributions to lichenology.
Cladonia sobolescens, commonly known as the peg lichen, is a species of fruticose lichen. It is found in temperate eastern North America and East Asia.
Rexiella is a lichen genus in the family Cladoniaceae. The genus, originally circumscribed with the name Rexia by authors Soili Stenroos, Raquel Pino-Bodas, and Teuvo Ahti in 2018, was created to contain the species Cladonia sullivanii, first formally described in 1882 by Swiss botanist Johannes Müller Argoviensis. After publication of the new genus, it was discovered that the name was illegitimate, because an earlier homonym had been published; the generic name RexiaD.A.Casamatta, S.R.Gomez & J.R.Johansen had already been created in 2006 to contain the cyanobacterial species Rexia erecta. The new name Rexiella was therefore proposed in 2019. This name honours the Australian lichenologist Rex Filson, who published monographs on the genera Cladia and Heterodea.
Cladonia gracilis or the smooth cup lichen is a species of fruticose, cup lichen in the family Cladoniaceae. It was first described as a new species by Carl Linnaeus in his 1753 work Species Plantarum. German botanist Carl Ludwig Willdenow transferred it to the genus Cladonia in 1787. In North America, it is known colloquially as the "smooth Cladonia".
Pulchrocladia is a genus of fruticose lichens in the family Cladoniaceae. It has three species. The genus was circumscribed in 2018 by lichenologists Soili Stenroos, Raquel Pino-Bodas, Helge Thorsten Lumbsch, and Teuvo Ahti. The genus name refers to "the beautiful morphology of its species".
Cladonia alpina is a species of fruticose lichen that is in the family Cladoniaceae. It has been found in Japan, Norway, and the United Kingdom.
Cladonia mitis is a species of fruticose lichen in the family Cladoniaceae. It was formally described as a new species in 1918 by German lichenologist Heinrich Sandstede. It has previously been classified in genus Cladina before molecular phylogenetic studies showed this to be a part of Cladonia. Cladonia mitis is morphologically quite similar to Cladonia arbuscula, and some authors have considered it to be a variety or subspecies of the latter. They differ mainly in the production of secondary compounds: Cladonia mitis produces chemicals in the rangiformic acid complex, which C. arbuscula does not.
Pulchrocladia retipora, commonly known as the coral lichen, is a species of fruticose lichen in the family Cladoniaceae. Found predominantly in Australasia, its habitats range from the Australian Capital Territory to New Zealand's North and South Islands, and even the Pacific region of New Caledonia, where it grows in coastal and alpine heathlands. The lichen features coral-like branches and subbranches with numerous intricate, netlike perforations. It is known by multiple names, with some sources referring to it by its synonym Cladia retipora, or the common name lace lichen.
Pulchrocladia corallaizon is a species of lichen in the family Cladoniaceae. It was first formally described as Cladia corallaizon. The specific epithet corallaizon, modified from Greek, means "ever-living coral". In 2018, it was transferred to the newly circumscribed genus Pulchrocladia.
Pulchrocladia ferdinandii is a species of lichen in the family Cladoniaceae. It was first formally described as Cladonia ferdinandii by Swiss lichenologist Johannes Müller Argoviensis in 1882. The specific epithet honours German-Australian botanist Ferdinand von Mueller, who collected the type specimen near Esperance, Western Australia. Rex Filson transferred the taxon to Cladia in 1970. In 2018, it was transferred to the newly circumscribed genus Pulchrocladia.
Lobaria anomala, commonly known as the netted specklebelly, is a species of foliose lichen in the family Peltigeraceae. It is found in coastal western North America, where it grows on trees in humid environments. The lichen was first described as a new species in 1987 as a species of Pseudocyphellaria, though it had been mentioned in scientific papers before.
Cladonia longisquama is a species of fruticose lichen in the family Cladoniaceae. It occurs in the Seychelles, where it grows on moss-covered rocks.
Cladonia glacialis is a species of fruticose lichen in the family Cladoniaceae. Found in Iceland, it was described as a new species in 2009 by Hördur Kristinsson and Teuvo Ahti. The first author collected the type specimen in 1979 from Arnarfellsmúlar at an elevation of 600 m (2,000 ft). This type locality is on the southeast side of Múlajökull, which is an outlet glacier of the Hofsjökull ice cap. Cladonia glacialis is only known to occur at the type locality. It contains fumarprotocetraric acid as its major lichen product. The authors suggest that the species belongs to the "supergroup" Cladonia, and that it may be closely related to Cladonia phyllophora.
Cladonia krogiana is a species of fruticose lichen in the family Cladoniaceae, described to science in 2002. This lichen is characterised by its greyish-green to medium-brown squamulose (scaly) primary thallus, forming cushions up to 10 cm (4 in) wide, with distinctive chemical components including barbatic acid and the rare xanthone compound chlorovinetorin. Originally discovered in southeastern Norway, C. krogiana has since been found in the Czech Republic and New Brunswick, Canada, typically growing on sun-exposed siliceous rock in periodically wet, open areas near forests and water bodies.
↑ Hafellner, J. (1984). "Studien in Richtung einer natürlichen Gliederung der Sammelfamilien Lecanoracae und Lecideaceae" [Studies towards a natural classification of the collective families Lecanoracae and Lecideaceae]. Beihefte zur Nova Hedwigia. 79: 241–371.
1 2 Jørgensen, Per M. (1994). "Linnaean lichen names and their typification". Botanical Journal of the Linnean Society. 115 (4): 261–405. doi:10.1111/j.1095-8339.1994.tb01784.x.
↑ Wainio, Edvard August (1894). Monographia Cladoniarum universalis: II[Universal Monograph of Cladonias: II]. Acta Societatis pro Fauna et Flora Fennica. Vol.10. pp.1–499.
↑ Wainio, Edvard August (1897). Monographia Cladoniarum universalis: III[Universal Monograph of Cladonias: III]. Acta Societatis pro Fauna et Flora Fennica. Vol.14. pp.1–268.
1 2 Coppins, B. (2002). "Cladoniaceae. Teuvo Ahti. Flora Neotropica Monograph No. 78. New York: New York Botanical Garden Press. 2000. 362pp., 215 figures, 6 tables. ISBN 0 89327 431 3". Edinburgh Journal of Botany. 59 (3): 459–466. doi:10.1017/S0960428602230292.
1 2 3 4 Lücking, Robert; Hodkinson, Brendan P.; Leavitt, Steven D. (2017). "The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota–Approaching one thousand genera". The Bryologist. 119 (4): 371, 384–385. doi:10.1639/0007-2745-119.4.361.
↑ Hoffman, Jordan R.; Karol, Kenneth G.; Ohmura, Yoshihito; Pogoda, Cloe S.; Keepers, Kyle G.; McMullin, Richard T.; Lendemer, James C. (3 February 2023). "Mitochondrial genomes in the iconic reindeer lichens: Architecture, variation, and synteny across multiple evolutionary scales". Mycologia. 115 (2): 187–205. doi:10.1080/00275514.2022.2157665.
↑ Wedin, Mats; Döring, Heidi; Ekman, Stefan (2000). "Molecular phylogeny of the lichen families Cladoniaceae, Sphaerophoraceae, and Stereocaulaceae (Lecanorales, Ascomycotina)". The Lichenologist. 32 (2): 171–187. Bibcode:2000ThLic..32..171W. doi:10.1006/lich.1999.0236.
↑ Zhou, Qi-Ming; Wei, Jiang-Chun; Ahti, Teuvo; Stenroos, Soili; Högnabba, Filip (2006). "The systematic position of Gymnoderma and Cetradonia based on SSU rDNA sequences". Journal of the Hattori Botanical Laboratory. 100: 871–880.
↑ Kraichak, Ekaphan; Huang, Jen-Pan; Nelsen, Matthew; Leavitt, Steven D.; Lumbsch, H. Thorsten (2018). "A revised classification of orders and families in the two major subclasses of Lecanoromycetes (Ascomycota) based on a temporal approach". Botanical Journal of the Linnean Society. 188 (3): 233–249. doi:10.1093/botlinnean/boy060.
↑ Ahti, T. (1982). "The morphological interpretation of cladoniiform thalli in lichens". The Lichenologist. 14 (2): 105–113. doi:10.1017/s0024282982000255.
1 2 3 4 Jaklitsch, Walter; Baral, Hans-Otto; Lücking, Robert; Lumbsch, H. Thorsten (2016). Frey, Wolfgang (ed.). Syllabus of Plant Families: Adolf Engler's Syllabus der Pflanzenfamilien. Vol.1/2 (13ed.). Berlin Stuttgart: Gebr. Borntraeger Verlagsbuchhandlung, Borntraeger Science Publishers. p.121. ISBN978-3-443-01089-8. OCLC429208213.
↑ Škaloud, Pavel; Steinová, Jana; Řídká, Tereza; Vančurová, Lucie; Peksa, Ondřej; De Clerck, O. (2015). "Assembling the challenging puzzle of algal biodiversity: species delimitation within the genus Asterochloris (Trebouxiophyceae, Chlorophyta)". Journal of Phycology. 51 (3): 507–527. Bibcode:2015JPcgy..51..507S. doi:10.1111/jpy.12295. PMID26986666.
↑ Jahns, H.M.; Beltman, H.A. (2007). "Variations in the ontogeny of fruiting bodies in the genus Cladonia and their taxonomic and phylogenetic significance". The Lichenologist. 5 (5–6): 349–367. doi:10.1017/S0024282973000447.
↑ Hammer, Samuel (2018). "Primary tissue and the structure of the podetium in Cladonia". Mycologia. 87 (1): 46–53. doi:10.1080/00275514.1995.12026501.
↑ Grube, Martin; Hawksworth, David L. (2007). "Trouble with lichen: the re-evaluation and re-interpretation of thallus form and fruit body types in the molecular era". Mycological Research. 111 (9): 1116–1132. doi:10.1016/j.mycres.2007.04.008. PMID17698333.
↑ Jahns, H.M. (1970). Untersuchungen zur Entwicklungsgeschichte der Cladoniaceen unter besonderer Berücksichtigung des Podetien- Problems[Studies on the developmental history of the Cladoniaceae with special consideration of the podetium problem]. Nova Hedwigia (in German). Vol.20. pp.1–177.
↑ Jahns, H.M. (1970). "Induktion der Apothecienbildung bei Cladia aggregata (Sw.) Nyl" [Induction of apothecia formation in Cladia aggregata (Sw.) Nyl.]. Berichte der Deutschen Botanischen Gesellschaft (in German). 83: 33–40. doi:10.1111/j.1438-8677.1970.tb02301.x.
↑ Aptroot, André; da Silva Cáceres, Marcela Eugenia (2018). "New lichen species from Chapada Diamantina, Bahia, Brazil". The Bryologist. 121 (1): 67–79. doi:10.1639/0007-2745-121.1.067.
↑ Hammer, Samuel (2000). "Meristem growth dynamics and branching patterns in the Cladoniaceae". American Journal of Botany. 87 (1): 33–47. doi:10.2307/2656683. JSTOR2656683. PMID10636828.
↑ Lamb, I.M.; Weber, W.A. (1974). "Calathaspis, a new genus of lichens (Cladoniaceae) from New Guinea". Occasional Papers of the Farlow Herbarium of Cryptogamic Botany. 4: 1–12.
↑ Nylander, William (1870). Recognitio Monographica Ramalinarum[Monographic revision of the Ramalina lichens] (in French). Caen: Impr. de P. Le Blanc-Hardel. p.69.
↑ Galloway, D.J.; James, P.W. (1987). "Metus, a new austral lichen genus and notes on an Australian species of Pycnothelia". Notes from the Royal Botanical Garden Edinburgh. 44 (3): 561–579.
↑ Jørgensen, P.M.; Jahns, H.M. (1987). "Muhria, a remarkable new lichen genus from Scandinavia". Notes from the Royal Botanical Garden Edinburgh. 44: 581–599.
↑ Liu, Jian Kui; Hyde, Kevin D.; Jones, E. B. Gareth; Ariyawansa, Hiran A.; Bhat, Darbhe J.; Boonmee, Saranyaphat; etal. (2015). "Fungal diversity notes 1–110: taxonomic and phylogenetic contributions to fungal species". Fungal Diversity. 72 (1): 1–197. doi:10.1007/s13225-015-0324-y.
↑ Fries, Th. M. (1857). "Ad amimadversiones Cl. W. Nylanderi"[In response to the remarks of the distinguished W. Nylander]. Botaniska Notiser (in Latin). 1857: 167–174.
↑ Vainio, E.A. (1890). "Étude sur la classification naturelle et la morphologie des Lichens du Brésil. Pars prima" [Study on the natural classification and morphology of the Lichens of Brazil. Part one]. Acta Societatis Pro Fauna et Flora Fennica (in Latin). 7 (1): 7.
↑ Kistenich, Sonja; Timdal, Einar; Bendiksby, Mika; Ekman, Stefan (2018). "Molecular systematics and character evolution in the lichen family Ramalinaceae (Ascomycota: Lecanorales)". Taxon. 67 (5): 871–904. doi:10.12705/675.1. hdl:10852/67955.
↑ Wedin, Mats; Döring, Heidi (1999). "The phylogenetic relationship of the Sphaerophoraceae, Austropeltum and Neophyllis (lichenized Ascomycota) inferred by SSU rDNA sequences". Mycological Research. 103 (9): 1131–1137. doi:10.1017/S0953756298008223.
↑ Soto-Medina, Edier Alberto (2013). "Patrones de Riqueza de Especies de la Familia Cladoniaceae en el Neotrópico Patterns of Species Richness of Family Cladoniaceae in the Neotropics" [Patterns of species richness of family Cladoniaceae in the Neotropics]. Cryptogamie, Mycologie (in Spanish). 34 (2): 137–148. doi:10.7872/crym.v34.iss2.2013.137.
↑ Yánez, Alba; Ahti, Teuvo; Bungartz, Frank (2013). "The Family Cladoniaceae (Lecanorales) in the Galapagos Islands". Phytotaxa. 129 (1): 1–33. doi:10.11646/phytotaxa.129.1.1.
↑ Ahti, Teuvo; Stenroos, Soili; Moberg, Roland (2013). Nordic Lichen Flora. Cladoniaceae. Vol.5. Uppsala: Museum of Evolution, Uppsala University. ISBN978-91-85221-29-5.
↑ Burgaz, Ana Rosa; Márquez, Rodrigo; PinoBodas, Raquel (2022). "The Cladoniaceae (Lecanorales, Ascomycota) from Bulgaria". Herzogia. 35 (2): 510–540. doi:10.13158/heia.35.2.2022.510.
↑ Bukabayeva, Zhanilxan; Abiev, Sardarbek; Korolev, Alexander (2022). Species diversity of lichens on the territory of the Burabay National Park (Republic of Kazakhstan) (Report). pp.1–13. doi:10.21203/rs.3.rs-2195522/v1.
↑ Allen, J.; Lendemer, J.; McMullin, T. (11 August 2015). "Rock Gnome Lichen. Cetradonia linearis". The IUCN Red List of Threatened Species 2020. Retrieved 19 February 2021.
↑ Lendemer, J.; Allen, J.; McMullin, T. (6 October 2019). "Cladonia appalachiensis". The IUCN Red List of Threatened Species 2020. Retrieved 19 February 2021.
↑ Yahr, R. (30 April 2003). "Cladonia perforata". The IUCN Red List of Threatened Species 2003. Retrieved 19 February 2021.
↑ Ohmura, Y.; Nadyeina, O.; Scheidegger, C. (14 August 2014). "Gymnoderme insulare". The IUCN Red List of Threatened Species 2014. Retrieved 19 February 2021.
↑ Roturier, Samuel; Ollier, Sébastien; Nutti, Lars-Evert (2017). "Restoration of reindeer lichen pastures after forest fire in northern Sweden: Seven years of results". Ecological Engineering. 108: 143–151. doi:10.1016/j.ecoleng.2017.07.011.
↑ Hutchison, Melissa. "Pulchrocladia retipora". New Zealand Plant Conservation Network. Archived from the original on 19 April 2022. Retrieved 28 March 2022.
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