Cladoniaceae

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Cladoniaceae
Cladonia subulata is the type species of the type genus of the family Cladoniaceae
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Lecanoromycetes
Order: Lecanorales
Family: Cladoniaceae
Zenker (1827)[1]
Type genus
P.Browne
(1756)
Synonyms[2]

The Cladoniaceae are a

aridity
.

Many Cladoniaceae species are characterised by a

green algae from the class Trebouxiophyceae, usually the genus Asterochloris. Six Cladoniaceae species are included in the International Union for Conservation of Nature's Red List of Threatened Species
.

Systematics

Historical taxonomy

Cladoniaceae was formally introduced to science in 1827 by German naturalist

Baeomycetaceae, Icmadophilaceae, and Stereocaulaceae.[6] William Nylander included 53 Cladonia species worldwide in his 1860 work Synopsis lichenum.[7] When Edvard August Vainio published his three-volume monograph on the Cladoniaceae (Monographia Cladoniarum universalis, 1887,[8] 1894,[9] and 1897[10]), he included 134 species and subspecies. In his circumscription of the family, the genera Pycnothelia, Cladia, and Cladina were included in the genus Cladonia.[11] In more recent history, significant progress in the knowledge of the taxonomy and biogeography of the family can be attributed to several decades of devoted research by Teuvo Ahti.[12]

Cladoniaceae is now one of the largest families of lichen-forming fungi,[13] with about 560 species distributed amongst 18 genera. The type genus is Cladonia, circumscribed by 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."[14]

Etymology

As is standard practice in botanical nomenclature,[15] 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 meaning "quality or state of a thing".[16]

Synonymy

Several

Cetradonia linearis,[3] was folded into the Cladoniaceae in 2006.[22]

In 2018, Kraichak and colleagues used a technique called temporal banding to reorganize the

synonymised the families Squamarinaceae and Stereocaulaceae with the Cladoniaceae, resulting in a large increase in the number of genera and species.[23] The Squamarinaceae had already been included in the Cladoniaceae by previous authors.[13] 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. As Robert Lücking explained, "merging of the two families under the name Cladoniaceae is not possible without a conservation proposal because Cladoniaceae (Zenker, 1827) is antedated by Stereocaulaceae (Chevallier, 1826) by one year."[24] 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".[25]

Description

In Cladonia coccifera, the conidia are embedded in a scarlet-coloured slime on the margins of the scyphi (cups).

The

soredia are common.[25]

The ascomata are in the form of an

conidia are non-septate, usually filiform (thread-like), and hyaline.[27]

Chemistry

In the Cladoniaceae, over 70 different secondary metabolites (lichen products), primarily polyphenols, have been identified. 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 analysis becomes crucial for identifying the major lichen products in the specimens and verifying species identification.[29]

Photobionts

The symbiotic algal partner (

microbial community, which was found to be consistent amongst the different species, consisted largely of Alphaproteobacteria and Acidobacteriota.[33]

Development

Cladonia coniocraea (left) has subulate podetia, while the podetia of Cladonia fimbriata (middle) feature cup-shaped scyphi, commonly called "pixie cups", Cladonia cervicornis (right) has a more complex branching pattern.

The development of several Cladoniaceae genera have been studied in detail,[34][35][36][37] although the interpretation of results has sometimes been controversial.[28] 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.[38][39]

Cladoniaceae species begin development with the formation of a

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 very complex branching patterns.[29] 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.[40]

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.[29] The podetia are slow-growing, with an annual growth rate generally ranging from 1 to 15 mm.[41]

Branching in the Cladoniaceae occurs on the podetium due to the growth dynamics of fungal meristem tissue at its apex. Two primary branching patterns exist: one where branches emerge from late divisions of a large meristem that alters its shape, and another where branches come from small meristems that split early but maintain their shape. These meristem growth dynamics are crucial for interpreting phylogeny in Cladoniaceae mycobionts, with the trend towards smaller, early-splitting meristems seen as an evolutionary advancement.[42] In cladoniiform lichens, especially within the Cladoniaceae, a shift in meristem growth from isotropous (having uniform properties in all directions) to anisotropous (having properties that differ depending on the direction in which they are measured) leads to pronounced lateral elongation of the apical meristem. This change offers developmental flexibility, transitioning from a symmetrical growth to a more varied, asymmetrical growth, hinting at evolutionary processes within the Cladoniaceae. Despite this variability, such morphogenetic activities appear to be highly conserved even among species that are presumably distantly related.[43]

Genera

After more than a century of discovery and research, including recent advances in understanding revealed by

taxonomic authority
, year of publication, and the number of species:

Pilophorus acicularis
Pycnothelia papillaria

Myelorrhiza was transferred from the Cladoniaceae to the Ramalinaceae by Kistenich and colleagues in 2018.[59] Neophyllis, originally classified in the Cladoniaceae, was transferred to Sphaerophoraceae in 1999.[60]

Habitat and distribution

Cladoniaceae species have been recorded growing in many habitats and on a diversity of

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.[28]

In his 2000 monograph on the Cladoniaceae of the Neotropical realm, Ahti included 184 species in 4 genera, and showed that that South America is a hotspot of biodiversity for genus Clanodia.[12] 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.[62] 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.[63] A 2013 monograph of Northern European Cladoniaceae treated 100 species (95 Cladonia, 4 Pilophorus, and the monotypic genus Pycnothelia).[64] In the 2021 key to lichen species in Italy, 86 Cladoniaceae are included.[65] In Bulgaria, 55 species in two genera were reported in 2022.[66] In a study of the lichen biodiversity in Kazakhstan's Burabay National Park, the Cladoniaceae made up about 30 percent of the species diversity.[67]

A study on the distributional ecology of Cladina and Cladonia in western North America found that 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.[68]

Conservation

Cladonia perforata has been an endangered species on the IUCN redlist since 2003.

Each of the six Cladoniaceae species that have been assessed for the global

Cryptomeria japonica and Chamaecyparis obtusa.[74]

On the

critically endangered), Gymnoderma coccocarpum (endangered), and Gymnoderma insulare (endangered) are the representatives of the Cladoniaceae.[75]

Human interactions and uses

Some species in the Cladoniaceae have been involved in a profitable export business for decorative uses, with demand reaching thousands of kilograms in various markets. In Europe, Cladonia stellaris is used ornamentally in wreaths, floral decorations and architect's models. In the years 1970–75, 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.[76] In Sweden, Cladonia lichens were used historically as a partial insulation for storm windows.[77] 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.[78]

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Cited literature

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