Cladonia rei | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Fungi |
Division: | Ascomycota |
Class: | Lecanoromycetes |
Order: | Lecanorales |
Family: | Cladoniaceae |
Genus: | Cladonia |
Species: | C. rei |
Binomial name | |
Cladonia rei Schaer. (1823) | |
Synonyms [1] | |
List
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Cladonia rei, commonly known as the wand lichen, is a species of ground-dwelling, fruticose lichen in the family Cladoniaceae. It is a widely distributed species, having been reported from Africa, Asia, Australasia, Europe, and North America. It is identified by its slightly dirty-colored, rough-surfaced, slender podetia that grow up to 9 cm (3.5 in) tall. Diagnostic characters of the lichen include the continuously sorediate, green-and-brown-mottled, podetia that taper upward to a point, while chemically, it contains homosekikaic and sekikaic acids. Its reduced capacity to bioaccumulate toxic heavy metals from its surroundings, as well as its ability to switch photobiont partners, allows the lichen to colonize and survive highly polluted habitats. There are several other Cladonia species that are somewhat similar in appearance, but can be distinguished either by subtle differences in morphology, or by the secondary chemicals they contain.
The species was first described scientifically in 1823 by Swiss lichenologist Ludwig Emanuel Schaerer. It is named after Professor Re from Turin, who had sent the specimens to Schaerer from Italy. In his original description, Schaerer noted the new lichen's similarity to Cladonia gracilis . [2] Schaerer's type specimen, kept at the herbarium of the Conservatoire et Jardin botaniques de la Ville de Genève, contains both homosekikaic and fumarprotocetraric acids. [3] In the genus Cladonia, C. rei is a member of clade Cladonia, subclade Graciles. [4]
In North America Cladonia rei is commonly known as the wand lichen, [5] or the sordid powderhorn. [6]
In an 1850 publication, Schaerer proposed that the taxon was better classified as a variety of Cladonia cornuta . [7] In 1894, Finnish lichenologist and Cladonia specialist Edvard August Vainio proposed it as a form of Cladonia fimbriata . [8] The two proposed names are now considered homotypic or nomenclatural synonyms of Cladonia rei. [9]
In 1976, Haavard Østhagen proposed that Cladonia nemoxyna, a taxon originally described by Erik Acharius in 1803, should be considered a synonym of Cladonia rei. [3] This nomenclatural possibility had been suggested already a decade earlier by Juha Suominen and Teuvo Ahti when they typified the name in a 1966 publication. [10] This name, and several other with the varietal or form epithet nemoxyna are heterotypic (taxonomic) synonyms of Cladonia rei. [1]
Cladonia rei is morphologically similar to Cladonia subulata , and the two are difficult to distinguish. In 1993, Paus and colleagues studied all of the morphological characteristics that were used to distinguish between the two species, and concluded that there were none that could sufficiently distinguish between the two, but they could still be considered distinct species due to their different habitat preferences. [11] Leo Spier and André Aptroot conducted a chemical and morphological study on about 350 mostly West European specimens of Cladonia subulata and C. rei. They found about a third of the specimens that had been identified based on inferences about their habitat had been misidentified. They also reported that the long-wave UV test—the standard method for distinguishing between the two species in the field—was unreliable for detecting homosekikaic acid. As a result, many of the previous records of these taxa should also be considered unreliable. They concluded that there was "no indication of any correlation between chemistry and geography, morphology or substratum", and that C. rei should be considered a chemotype of Cladonia subulata, rather than a distinct species. They proposed that C. rei be placed in synonymy with Cladonia subulata, because the latter, as the older of the two names, has priority. [12]
Some studies published soon challenged Spier and Aptroot's proposal. Authors of a study of this taxon in Poland argued that the species should remain distinct, largely because "Their conclusions were based on material mostly from the Netherlands, with rather scanty representatives from other countries". After analysing almost 800 collections, they determined that 90% of the 228 specimens of C. rei had been misidentified, a figure they attributed to the original collectors not having corroborated their determinations with thin layer chromatography to accurately assess the chemistry. [13] In 2010, a molecular phylogenetics study, showed the subclade of C. subulata to be distinct from C. rei. The authors propose that the determining chemical feature for C. rei is the presence of homosekikaic acid. [14] Additional molecular support for separate species in another study also used scanning electron microscopy of the stereome surfaces that revealed distinct differences between the two species. [15]
Primary squamules are small and inconspicuous, often disappearing. [5] Podetia are gray-green to olive, sometimes browned, slender, coarsely or finely sorediate on the upper half. [5] They have dimensions in the range of 2–9 cm (13⁄16–3+9⁄16 in) tall by 1–2 mm (up to 3.5 mm) thick. [16] They have with narrow, symmetrical, or, more frequently lopsided cups, proliferating at the margins to resemble a star or magic wand. [5] Pale to dark brown apothecia are common on proliferations along the cup margins. [5] It is prudent to take heed of Spier and Aptroot's thoughts: "There is much individual variation between the specimens, which can vary from nearly unbranched to cup-shaped, from slender (less than 1 mm thick) to robust (reaching 5 mm) and from low (average height below 2 cm) to high (height over 5 cm, clearly higher than the surrounding Cladonia species). The distribution of these forms over the chemotypes is on the whole largely similar and differentiating characteristics could not be found." [12] Further, it is known that growth in contaminated sites can influence the growth and development of the Cladonia thalli, sometimes resulting in features that do not coincide with the normal taxonomic definition of the species. [17] This phenomenon has been documented with Cladonia rei. [18]
Chemical spot tests are undergone for identification. C.rei responds to P+ red for fumarprotocetraric acid, P− and UV+ white for homosekikaic acid. [5] PD reactions also turn C.rei yellow in low presences of fumarprotocetraric acid. [13] [19]
Cladonia rei invariably contains homosekikaic acid; sometimes it also has fumarprotocetraric acid. It also has smaller amounts of sekikaic acid, and well as trace amounts of the accessory substance 4'-O-methylnorhomosekikaic acid. The positive fluorescence of an ultraviolet test is traditionally used to detect the presence of homosekikaic acid. It can also be tested using the ferric chloride chemical spot test, which produces a violet-coloured spot if positive. [15] Other compounds that have been found in the lichen include antheraxanthin, astaxanthin, α-carotene, ε-carotene, β-cryptoxanthin, lutein, lutein epoxide, and violaxanthin. [20]
Cladonia subulata and Cladonia ochrochlora are similar species that can be distinguished by their secondary chemistry. Fumarprotocetraric acid is the only main lichen metabolite in C. subulata. Another lookalike, Cladonia glauca , contains squamatic acid, a substance that causes it to glow white when lit with a UV light. It also has a narrow longitudinal slit in most podetia. [4] Lots of details available. [11] Cladonia cenotea might be closely related. [15] Cladonia coniocraea and Cladonia cornuta tend to have larger primary squamules and do not normally proliferate at the cup margins; neither contains homosekikaic acid. [5]
Cladonia rei is a terricolous lichen that grows in soil, although more rarely it is found on rotting wood, [16] and there are few records of it growing on other substrates, including rock, plastic, [12] old straw, and mosses over rocks. [13] It prefers open areas with low humus content and a substrate with a more or less neutral soil pH. [15] often in dry and sunny places like grasslands, heaths and wastelands. [21] Another favoured habitat is limestone cliffs and limestone meadows, although the lichen is also found in disturbed areas, such as road embankments, gravel pits, or house ruins. [22] It is a pioneer species: able to quickly colonise bare ground, particularly in disturbed and anthropogenic habitats. [21] In some cases, it can grow on impoverished soil with high heavy metal content. In Krompachy (Slovakia), the lichen, along with the moss Ceratodon purpureus , was found to be very abundant in bare acid soil near copper smelters, where all vascular plants were severely damaged or have disappeared. [23] It is prevalent in temperate or somewhat arid regions, and does not occur in the Arctic or the Antarctic. [15] The lichen was used as a case study for a large-scale biodiversity monitoring program in Alberta, Canada. Historical records and collections suggested that the lichen was broadly distributed but rare in the province, but more recent material from surveys conducted by the Alberta Biodiversity Monitoring Institute showed that its distribution is entirely limited to the dry mixed grassland, northern fescue grassland and aspen parkland natural regions, where it is relatively common. [24]
It is a widespread species, with a range extending to Europe, Africa, Asia, Australasia, North America, [22] and South America. It is widely distributed in Europe, having been recorded in Austria, Belgium and Luxemburg, Czech Republic, Estonia, Lithuania, Norway, Poland, Slovakia, Spain, and Sweden. [13] More recently, it was reported from Armenia, [25] from the coastal sand-dunes of Portugal, [26] and, in 2021, from the Pyrénées-Orientales in the French Mediterranean region. [27] Its Australasian distribution includes both Australia [13] and New Zealand. [20] It was reported as new to India in 2002. [28] In 2006 it was recorded from Iran [29] and Turkey. [30]
In the United States, it has been considered by some authors to be widely distributed in the northwestern part of the country, but probably because of misidentification with the much more common C. ochlorochlora and C. verruculosa. Samuel Hammer suggests that the actual range in the region is limited to interior localities east of the Cascade Mountains, in Idaho and Washington. [31]
Because the lichen has often been recorded growing in extremely contaminated post-smelting slag dumps, its bioaccumulative abilities have been studied. The lichen has developed a restrained heavy-metal accumulation pattern that limits its exposure to these toxic compounds, and helps it to colonise these contaminated sites. [18] Efficient methods have been developed for extracting arsenic compounds from this species; in an arsenic-contaminated environment, Cladonia rei was shown to contain arsenous acid, arsenic acid, methylarsonic acid, dimethylarsinic acid, arsenobetaine, and trimethylarsine oxide. [32] Its high genetic variability within a single population is suggestive of great potential for colonising anthropogenic habitats, and underlies the role of this lichen as a pioneer in the early stages of natural regeneration of these contaminated sites. [33] Cladonia rei has a low bioaccumulation factor, meaning it is classified as a weak accumulator of heavy metals. Its erect and fruticose growth form help it survive in this polluted environment, due to the decrease in metal content along the vertical gradient of the thallus. [34] The level of cell membrane damage in the lichen is dependent on the heavy level concentrations of the polluted site. The amount of cell damage can be measured experimentally by determining relative electrical conductivity, and thus, this physiological parameter can serve as an early warning indicator for detection of elevated metal concentrations in soil. [35]
A phytosociological study of the pioneer cryptogamic communities associated with Cladonia rei in Polish post-smelting dumps showed the frequent co-occurrence of Diploschistes muscorum and Cladonia conista . In non-polluted sites, typical lichen associations include C. chlorophaea , C. fimbriata , C. pyxidata , Ceratodon purpureus , and Peltigera didactyla . [21] In northeastern North America, it is noted to be a common pioneer species in old field succession, along with Cladonia cristellata and Cladonia polycarpoides . [5] It is also a characteristic species of cryptogamic communities in dry grasslands of Saskatchewan, [36] and anthropogenic habitats in Germany. [37] Some cryptogamic communities, including those involving C. rei, are quite similar in Europe and North America, even though the vascular plant species composition often differs considerably. [21]
Epicladonia sandstedei is a lichenicolous fungus that parasitises Cladonia rei. [38]
In a study of the oribatid mite communities associated with three Cladonia species in a heavily contaminated habitat, Cladonia rei was shown to provide a good habitat for mites. Because of the great variation in the growth forms of its podetia, densely covered with propagules such as corticated granules, squamules and microsquamules as well as numerous non-corticated, farinose-to-granular soredia, the surface area/volume ratio of the lichen is higher compared to other studied Cladonia species. The rugged surface texture provides more microhabitats for both mites and the particles of organic matter that are their diet. This contrasts with Cladonia cariosa and Cladonia pyxidata , which have a less diverse morphology, and a lower diversity of oribatid inhabitants. [39]
The photobiont partners of Cladonia rei are to some extent dependent upon the environment. Although both Asterochloris and Trebouxia can associate with the fungus, "the presence of Trebouxia was directly related to anthropogenic sites with technogenic substrates, and the proportion of lichen specimens with these algae clearly depended on the level of heavy-metal soil pollution and the habitat type. Although Cladonia lichens were previously thought to be restricted to Asterochloris, they are able to start the relichenization process with Trebouxia under specific habitat conditions and to establish a stable association with these algae when colonization of disturbed sites takes place." [40]
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.
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.
Cladonia mongkolsukii is a species of fruticose lichen in the family Cladoniaceae. Described as new to science in 2011, it is found in lower-elevation montane scrub forests of northeast Thailand and in Sri Lanka. The specific epithet honors Pachara Mongolsuk, a Thai lichenologist.
Cladonia wainioi or the Wainio's cup lichen is a species of cup lichen found in boreal and arctic regions of the Russian Far East and northern North America.
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.
Cladonia subradiata is a widely distributed species of fruticose lichen in the family Cladoniaceae. It is found in Asia, Africa, Melanesia, Australia, New Zealand, and South, Central, and North America.
Cladonia arbuscula, also referred to as shrubby cup lichen or green reindeer lichen, is a species of cup lichen in the family Cladoniaceae.
Cladonia inflata is a rare species of terricolous (ground-dwelling) lichen in the family Cladoniaceae. Found in Bahia, Brazil, it was formally described as a new species in 2018 by lichenologists André Aptroot and Marcela Eugenia da Silva Cáceres. The type specimen was collected by the authors from Palmeiras, on the Mount of Pai Inácio, at an altitude between 1,050 and 1,140 m ; here the lichen was found growing on siliceous sandstone rock in a transitional forest. Cladonia inflata is only known to occur at the type locality, and is only known from the type specimen. At this location the lichen is conspicuous but not abundant, and forms extensive mats with many other Cladonia species, such as C. bahiana, C. clathrata, C. dissecta, C. divaricata, C. friabilis, C. furfuracea, C. metaminiata, C. miniata, C. obscurata, C. parvipes, C. pityrophylla, C. polyscypha, C. salmonea, C. secundana, and C. substellata. The lichen has a fruticose (bushy), mineral-grey thallus that consists of upright hollow podetia measuring about 4 to 7 cm high, atop a cushion up to 10 cm (4 in) in diameter. It contains the secondary compound fumarprotocetraric acid. The specific epithet inflata refers to the inflated thallus of the lichen.
Cladonia compressa is a species of lichen in the family Cladoniaceae. Found in Bolivia, it was formally described as a new species in 2016 by lichenologists Teuvo Ahti and Adam Flakus. The type specimen was collected by the second author near Siniari colony at an altitude of 2,186 m (7,172 ft). Here, in a Yungas secondary cloud forest, the lichen was found growing on the ground, in humus-rich mineral soil. The specific epithet compressa refers to the compressed podetia. Secondary compounds that occur in the lichen include fumarprotocetraric acid (major), and minor to trace amounts of protocetraric acid and physodalic acid.
Cladonia graeca is a species of saxicolous (rock-dwelling), fruticose lichen in the family Cladoniaceae. Found in Greece, it was formally described as a new species in 2011 by lichenologists Harrie Sipman and Teuvo Ahti. The type specimen was collected by Sipman and Thomas Raus from the summit area of Mount Ochi (Euboea), at an altitude of 1,370 m (4,490 ft); it has also been recorded on the island Thasos. The small and inconspicuous lichen grows on boulders and cliffs of siliceous schist, especially in areas where downward-trickling water collects in dwarf shrub vegetation. It is similar to C. macrophylla and C. decorticata, but differs chemically from those species, as it contains fumarprotocetraric acid rather than psoromic acid or perlatolic acid.
Cladonia trassii is a species of fruticose lichen in the family Cladoniaceae. It has a circumpolar distribution and is found in arctic/alpine and subarctic habitats.
Cladonia vescula is a species of fruticose lichen in the family Cladoniaceae. It is known for its small size and unique chemical composition. It can be found in the montane cloud forests and pre-Andean Amazonian forests of Bolivia and Peru, where it grows on mineral soil mixed with humus. This species closely resembles Cladonia peziziformis and Cladonia corymbosula in morphology, but it is more slender and almost entirely lacking a cortex.
Cladonia lutescens is a species of fruticose lichen in the family Cladoniaceae. It occurs in high-altitude conditions of the Himalayas.
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 monomorpha is a species of terricolous (ground-dwelling), fruticose lichen in the family Cladoniaceae. It is part of the Cladonia pyxidata group, known for brown apothecia on cup-shaped podetia.
Cladonia flavocrispata is a species of fruticose lichen in the family Cladoniaceae. Found in Venezuela and described as a new species in 2013, it is closely related to Cladonia hians, but is distinguished by its unique chemical composition and morphology.
Cladonia incerta is a species of fruticose lichen in the family Cladoniaceae. Found in New Zealand, it was formally described as a new species in 2003 by the lichenologist Samuel Hammer. He collected the type specimen near the Dome Forest walkway north of Warkworth. The species epithet incertain refers to the uncertain ontogeny (development) of the podetia, which take various forms.
Cladonia nitidella is a species of fruticose lichen in the family Cladoniaceae. It is found in New Zealand, where it grows on bare soil and among mosses in often moist, partially sun-exposed habitats. It was formally described as a new species in 2003 by the lichenologist Samuel Hammer. He collected the type specimen from Rainbow Reach above Waiau River, about 10 km south of Te Anau, at an elevation of 200 m.
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.
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