Multiclavula mucida | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Basidiomycota |
Class: | Agaricomycetes |
Order: | Cantharellales |
Family: | Hydnaceae |
Genus: | Multiclavula |
Species: | M. mucida |
Binomial name | |
Multiclavula mucida (Pers.) R.H.Petersen (1967) | |
Synonyms [1] | |
List
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Multiclavula mucida is a globally distributed species of basidiolichen belonging to the family Hydnaceae. [5] Since its initial classification by Christiaan Hendrik Persoon in 1797, the species has been described under various synonyms and associated with multiple genera. Persoon described its fruiting bodies as gregarious (i.e. growing in groups but separate at the base), varying from simple to branching structures, predominantly whitish in colour with yellow to brownish tips. He also noted its frequent appearance in autumn on decaying, moist wood, often covered with a greenish crust necessary for its growth.
The cosmopolitan distribution of Multiclavula mucida and its variability in appearance have led to a complex nomenclature and historical confusion with other species and genera. Though widespread, it is considered rare in Europe and faces threats in many countries. It can be distinguished from similar species such as Multiclavula petricola , M. vernalis , M. coronilla , and Clavicorona taxophila by differences in basidiospore size, growth substrate, fruiting body structure, and microscopic features, despite similarities in colour and general form.
The fungus was first formally described by Christiaan Hendrik Persoon in 1797, as Clavaria mucida. Persoon characterised the fungus as gregarious, small, and either simple or branching. He noted its autumnal appearance on decaying, moist tree trunks covered with a crust or green powder, sometimes extending over a wide area. He described each specimen of the species as small and slender, occurring in both simple and more frequently in branched forms. The initial growth stage is marked by a thickened, wrinkled, and slightly curved apex. In branched specimens, the structure divides into thin, diverging, sharp branches originating from the middle or base. Persoon described the flesh as fleshy but not very tenacious, primarily whitish with the tip of the fungus being more intensely ochre in simpler forms, growing on a thin, whitish substrate or tomentum. He questioned the unique nature of the greenish crust common to this species, noting that it seemed necessary for its growth but disappeared when dried. Additionally, Persoon considered whether what he considered related species, Ramaria ornithopodoides and Clavaria albida, due to their significant size and branching nature, should be classified together with Clavaria mucida or recognised as distinct species. [6]
The fungus has acquired several synonyms as a result of having been transferred to different genera in its taxonomic history. After its initial placement in Clavaria , it has been proposed for transfer to Typhula (by Elias Fries in 1827), Calocera (Richard Wettstein, 1886), Pistillaria (Julien Noël Costantin and Léon Marie Dufour, 1891), Lentaria (E. J. H. Corner, 1950), and Stichoclavaria (Eckehard Paechnatz, 1987). [1] American mycologist Ron Petersen, in a 1967 publication transferred the taxon to the newly circumscribed genus Multiclavula in 1967. Before Corner's 1950 monograph on clavarioid fungi, [2] little attention was given to hyphal structure and morphology in their taxonomy. Despite Corner's extensive literature compilation, he couldn't inspect many type specimens, leading to reliance on past reports for species reclassification. During Petersen's study of Clavulinopsis and comparing it to species in Lentaria's subgenus Lentariopsis as defined by Corner, it became clear that several species across both genera could be congeneric. Corner himself was uncertain about the internal relationships within Lentaria, particularly about the evolutionary connections between its species and those in the Ramaria stricta series. This ambiguity led Petersen to reevaluate the genera housing these fungi. [7]
The reclassification of Multiclavula mucida was influenced by comparative analyses with other genera of lichen-forming fungi. While Clavulinopsis species are mostly terrestrial, characterised by white, thin- to slightly thick-walled spores and monomitic hyphae (a type of fungal tissue composed of uniformly structured hyphae), M. mucida has distinctive traits. Similarly, Lentaria, particularly exemplified by L. surculus , contrasts with M. mucida due to its predominantly lignicolous nature and thicker-walled hyphae. Despite these differences, the species within Lentaria have been subdivided into two distinct groups, neither of which aligns with Corner's subgenera, which indicated the necessity for a new genus classification. [7]
This reevaluation, spurred by inconsistencies and the unique characteristics of the C. mucida group, led to significant taxonomic revisions. Historical classifications that placed C. mucida within various genera were reexamined. Petersen's review of relevant type specimens, particularly those related to Lentariopsis, concluded that the subgenus should be amalgamated with Lentaria. This consolidation and the unique aspects of C. mucida facilitated the establishment of Multiclavula as a distinct genus, addressing both phylogenetic and morphological disparities. [7]
The species epithet mucida is from the Latin word mucidum, meaning "mouldy" or "slimy"; the genus name Multiclavula means "many clubs". [8] In North American literature, this fungus is known by several vernacular names, including "scum lover", [9] "common club mushroom lichen", [10] "frail bones", [8] and "club lichen", [11] the last of which is also used to refer to the entire genus. [12]
In the thallus of M. mucida, the algae are not merely scattered. Instead, they are organised into small, closely situated globules embedded in a gelatinous matrix on moist, rotting wood. [13] This Botrydina-type thallus structure, common in basidiolichens, serves multiple functions. It protects the algae from UV light and desiccation, while ensuring close contact through a hyphal network that provides a continuous water and minerals supply. [14] The fruiting bodies (basidiocarps) are typically simple to lobed and can reach up to 2 cm (0.8 in) in height. They are waxy-tough but can become slimy, transitioning from white or creamy colour to grey or pale tan as they mature. These structures often dry to show a pure white tip. [7] The fruiting bodies' stickiness often causes debris to adhere to their surfaces. The algal partners are encapsulated in tiny pockets of fungal tissue, yet they retain a loose, unstructured form, manifesting as a green granular crust where the fungus produces its fruiting bodies. Despite their mutualistic relationship, both the algae and the fungus maintain their individual morphology. [15] The relationship between coccomyxoid green algae and M. mucida was documented by Geitler in 1955, noting that the algae are larger within the lichen than when free-living. [16] He suggested in 1956 that this size difference could be due to a decrease in algal division frequency induced by the fungal partner, the mycobiont . [17] Using electron microscopy, Hiroshi Masumoto identified the photobiont partner of M. mucida as Elliptochloris subsphaerica . [18]
The fungus's internal structure features thin to slightly thick-walled hyphae. These threads are generally parallel, interlocking, and often merged, showing many branching and connecting points (anastomoses). These hyphae also feature clamp connections, which are small bridge-like structures connecting the cells, and can be up to 200 μm long. [7]
Below the surface layer, thinner, more twisted hyphae form the basidia as side branches. Basidia are typically short and narrow, with four to six outgrowths (sterigmata) from which spores are released. They measure 15–25 by 4–6 μm. The spores themselves are ovoid to ellipsoid, smooth, and can contain one or two oil droplets (guttulate), appearing white in spore prints and measuring 4.5–7.7 by 1.8–3.2 μm. [7] M. mucida does not react with any of the standard chemical spot tests and is not known to contain any secondary metabolites (lichen products). [10]
David Arora, in his popular field guide Mushrooms Demystified , describes the edibility of these tiny and temporary fruit bodies as "utterly inconsequential". [9]
Multiclavula mucida shares similarities with the Japanese species Multiclavula petricola . They resemble each other in the colour and general structure of their fruiting bodies, which are typically white and primarily unbranched. However, the two species differ significantly in the size of their basidiospores: those of M. mucida measure 4.5–7.7 μm in length, which are shorter than the 7.3–9.5 μm-basidiospores of M. petricola. Additionally, while M. mucida typically grows on decaying wood, M. petricola prefers rocky substrates. [19] Multiclavula vernalis , while somewhat similar in appearance to M. mucida, is distinguished by its orange colour, its preference for growing on soil, and the presence of cystidia. [9] Multiclavula coronilla , which grows on similar substrates, bears resemblance but can be differentiated by its shorter and simpler (unbranched) fruiting bodies, which are up to 1 cm tall, and its by its basidia, which have 6 to 8 sterigmata. Differentiating these two species typically requires microscopic examination. [10]
Another species resembling M. mucida is Clavicorona taxophila . This small, whitish species, characterised by a broadened, flattened tip, typically grows on twigs, needles, and other types of debris, rather than amidst scum on rotting wood. [9] In the Neotropics, M. mucida closely resembles Sulzbacheromyces leucodontium in appearance. However, their ecological preferences differ; M. mucida mainly grows on decaying logs, while S. leucodontium typically inhabits terrestrial environments in lowland and sub-lowland forests. Additionally, Multiclavula species are more commonly found in mountainous regions. [20]
This species is commonly found on decomposing wood, often in the presence of algae. [7] It thrives on a wide variety of substrates, including bamboo, [21] beech, [22] cedar, [23] poplar, and oak. [24] In Central Europe, coniferous wood is a particularly favoured substrate. [13] M. mucida is widely distributed, found in Asia, Europe, North and South America, and possibly Australia, [7] [10] with its North American range extending as far north as Alaska. [25] The discovery of M. mucida at unusually low elevations, specifically around 130 to 135 m (427 to 443 ft) above sea level in the Bienwald area of Rhineland-Palatinate, has prompted a reevaluation of its distribution and naming inconsistencies across different scientific literatures. Unlike other members of its genus, which prefer sandy soils, this species predominantly relies on rotting wood in moist environments. [26] Lothar Geitler first identified this species as an extratropical lichen in 1955, [16] challenging the then-prevailing view that basidiolichens were exclusively tropical. [13]
Previously considered native to Northern Europe and the mountains of Central Europe, M. mucida has been increasingly observed across most European countries and globally. This expansion in its geographical range could be attributed to the establishment and maintenance of natural forest reserves, environments that foster the decay of thick logs and provide suitable habitats for the fungus. In addition to its growing presence in traditional locales, the species has been documented in regions as far-reaching as Australia, including Tasmania, and New Zealand. [27]
Historically, however, the species has faced declines due to intensive forestry and environmental pollution, such as soil acidification from sulfur dioxide emissions, which damage its natural habitats. [26]
Multiclavula mucida has been categorised differently in mycological and lichenological circles, leading to its varying status in conservation lists. In Germany, for example, it has been listed under different categories in the Red List of endangered plants and macrofungi, reflecting a disparity in its assessment between being recognised as a fungus and as a lichen. [26] In Finland's Regional Red List, it is classified as endangered, with a population size of less than 2,500 individuals and a continual decline. [28] In Lithuania, it is considered very rare, being limited to only 1–3 known localities, [29] and in Estonia, it is categorised as vulnerable. [30] In the latter country, it is considered an indicator of old-growth forest. These primeval forests, where the lichen is found, are increasingly rare in Europe and are mainly preserved in nature reserves. [31] Because of its small size and seasonal appearance, M. mucida has been described as inconspicuous and easy to overlook. [32] [33]
A lichen is a hybrid colony of algae or cyanobacteria living symbiotically among filaments of multiple fungi species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualistic relationship. Lichens are the lifeform that first brought the term symbiosis under biological context.
Basidiolichens are lichenized members of the Basidiomycota that form a much smaller group of lichens than the far more common ascolichens in the Ascomycota. In arctic, alpine, and temperate forests, the most common basidiolichens are in the agaric genus Lichenomphalia and the clavarioid genus Multiclavula. Several lichenized genera occur in tropical regions, the most common being the foliose Dictyonema. Previously basidiolichens had been classified in their own subclass, Basidiolichenes. Molecular based phylogeny does not support classification of the genera together.
Clavaria is a genus of fungi in the family Clavariaceae. Species of Clavaria produce basidiocarps that are either cylindrical to club-shaped or branched and coral-like. They are often grouped with similar-looking species from other genera, when they are collectively known as the clavarioid fungi. All Clavaria species are terrestrial and most are believed to be saprotrophic. In Europe, they are typical of old, mossy, unimproved grassland. In North America and elsewhere, they are more commonly found in woodlands.
The clavarioid fungi are a group of fungi in the Basidiomycota typically having erect, simple or branched basidiocarps that are formed on the ground, on decaying vegetation, or on dead wood. They are colloquially called club fungi and coral fungi.
Clavaria fragilis, commonly known as fairy fingers, white worm coral, or white spindles, is a species of fungus in the family Clavariaceae. It is synonymous with Clavaria vermicularis. The fungus is the type species of the genus Clavaria and is a typical member of the clavarioid or club fungi. It produces tubular, unbranched, white basidiocarps that typically grow in clusters. The fruit bodies can reach dimensions of 15 cm (5.9 in) tall by 0.5 cm (0.2 in) thick. Clavaria fragilis is a saprobic species, growing in woodland litter or in old, unimproved grassland. It is widespread throughout temperate regions in the Northern Hemisphere, but has also been reported from Australia and South Africa. The fungus is edible, but insubstantial and flavorless. There are several other small white coral-like fungi with which C. fragilis may be confused.
Clavaria zollingeri, commonly known as the violet coral or the magenta coral, is a widely distributed species of fungus. It produces striking tubular, purple to pinkish-violet fruit bodies that grow up to 10 cm (3.9 in) tall and 7 cm (2.8 in) wide. The extreme tips of the fragile, slender branches are usually rounded and brownish. A typical member of the clavarioid or club fungi, Clavaria zollingeri is saprobic, and so derives nutrients by breaking down organic matter. The fruit bodies are typically found growing on the ground in woodland litter, or in grasslands. Variations in branching and color can often be used to distinguish C. zollingeri from similarly colored coral fungi such as Alloclavaria purpurea and Clavulina amethystina, although microscopy is required to reliably identify the latter species.
Clavariadelphus ligula, commonly known as the strap coral, is a species of fungus in the family Gomphaceae. It produces club-shaped fruit bodies with spongy flesh that grow in groups on the forest floor. It is found in Asia, Europe, and North America.
Multiclavula is a genus of basidiolichens in the family Hydnaceae. The widespread genus contains 14 species. The genus was circumscribed by the American mycologist Ron Petersen in 1967, with Multiclavula corynoides assigned as the type species.
Clavulinopsis amoena is a clavarioid fungus in the family Clavariaceae. It forms slender, cylindrical, golden-yellow fruiting bodies that grow on the ground among plant litter. It was originally described from Indonesia and appears to be distributed in temperate areas of the southern hemisphere.
Clavulinopsis helvola is a clavarioid fungus in the family Clavariaceae. In the UK, it has the recommended English name of yellow club. It forms slender, cylindrical, yellow fruiting bodies that grow on the ground in agriculturally unimproved grassland or in woodland litter. It was originally described from Europe.
Clavulinopsis fusiformis is a clavarioid fungus in the family Clavariaceae. In the UK, it has been given the recommended English name of golden spindles. In North America it has also been called spindle-shaped yellow coral or golden fairy spindle. Clavulinopsis fusiformis forms cylindrical, bright yellow fruit bodies that grow in dense clusters on the ground in agriculturally unimproved grassland or in woodland litter. It was originally described from England and is part of a species complex as yet unresolved.
Lepidostromatales is an order of fungi in the class Agaricomycetes. It is the only known order of basidiomycete fungi composed entirely of lichenized members. Morphologically, the fruiting bodies of all species are clavarioid. Six species are known, five of which were described within the span of 2007–2013. Due to its morphological similarity to the genus Multiclavula, its isolated phylogenetic position was not understood until quite recently. The photobionts that have been found in association with members of this group are not known to associate with any other types of lichenized fungi.
Multiclavula vernalis or the orange club-mushroom lichen is a species of clavarioid fungus in the Clavulinaceae family. It was originally named as a species of Clavaria in 1822 by Lewis David de Schweinitz. Ronald H. Petersen transferred it to Multiclavula in 1967.
Symbiosis in lichens is the mutually beneficial symbiotic relationship of green algae and/or blue-green algae (cyanobacteria) living among filaments of a fungus, forming lichen.
Hirticlavula is a fungal genus in the family Clavariaceae. It contains a single described species, Hirticlavula elegans. Formally described in 2014, the fungus has been collected from Norway and Denmark. H. elegans produces white fruit bodies up to 1.1 millimetres (0.043 in) in height. Each fruit body contains a fertile head atop a hairy stem. The fruit bodies grow directly from dead bark or wood, where they feed as saprotrophs. Both morphological and ecological details are distinctive when compared to other clavarioid fungi.
Clavulinopsis sulcata is a clavarioid fungus in the family Clavariaceae and is the type species of the genus Clavulinopsis. It forms very long, slender, cylindrical pinkish or orange fruiting bodies that grow on the ground among plant litter. A vernacular name that has been used for the species is flame fungus.
Multiclavula petricola is a species of saxicolous (rock-dwelling) basidiolichen in the family Hygrophoraceae. Found in Japan, it was formally described as a new species in 2020 by Hiroshi Masumoto and Yousuke Degawa. The type specimen was collected from Katashina at an altitude of 1,757 m (5,764 ft); here, in a shady coniferous forest, it was found growing on wet volcanic rock. The lichen makes tiny white fruitbodies and has a globular thallus. It is only known from the type locality. The photobiont partner of the lichen is the green algae that lives on the rock surface. The Latin species epithet petricola means "dweller on rocks". The Japanese name Iwa-no-shira-tsuno combines the Japanese words iwa-no with shira ("white") and tsuno ("horns").
Lichenomphalia velutina is a species of basidiolichen in the family Hygrophoraceae. It was first formally described in 1886 by the French mycologist Lucien Quélet, who classified it in the genus Omphalia. It was transferred to its current genus, Lichenomphalia, in 2002. Known primarily as having a European and North American distribution, the fungus was recorded from China for the first time in 2018. Chinese collections occur at elevations greater than 3,000 m (9,800 ft), while those in Europe tend to be at elevations under 1,500 m (4,900 ft). Lichenomphalia velutina also occurs in Greenland. It grows on soil and on moss.
Normandina pulchella, commonly known as the elf-ear lichen or blue heart, is a species of squamulose lichen in the family Verrucariaceae. This cosmopolitan species is widely distributed across both hemispheres, where it thrives in moist microhabitats. It favours moss-covered deciduous trees and rocks, often colonising over mosses and bryophytes. It occasionally grows on bare bark and on other lichens. Distinctive features of N. pulchella include its bluish-green squamules (scales) with sharply raised margins, non-reactivity to standard chemical spot tests, and growth in humid habitats. Initially, Nannochloris normandinae, a green alga, was thought to be its photobiont. However, recent studies have revised this understanding, now suggesting Diplosphaera as the algal partner.
Dictyocatenulata is a fungal genus of uncertain classification in the subclass Ostropomycetidae. It contains the single species Dictyocatenulata alba. Originally described in 1967 as a bark-dwelling fungus, it was later recognised as a lichen in 2004. The genus is characterised by its unique reproductive structures called synnemata, which are upright, stem-like formations that produce spores. D. alba has a widespread distribution, found in North and Central America, Asia, and Europe, typically growing on tree bark in humid forests. Recent molecular studies suggest that Dictyocatenulata may be closely related to the genus Thelenella, potentially representing an asexual stage of unknown Thelenella species. This lichen is distinguished by its thin, greenish thallus (body) and its spores, which are divided into many cells and arranged in chains, features that give the genus its name.