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Heterodermia

Heterodermia is a genus of foliose lichens in the family Physciaceae. The genus has a widespread distribution, especially in tropical regions, and contains about 70 species. Heterodermia was historically confused with the related genus Anaptychia, but was distinguished in 1965 based on differences in spore structure and chemical composition. These lichens can be identified in the field using simple chemical spot tests that produce distinctive colour changes when applied to the thallus. The lichens in this genus are small- to medium-sized, usually pale grey in colour, comprising narrow lobes with widened tips fringed with cilia. The lichens can be identified by their thick-walled ascospores and the presence of specific chemical compounds that produce colour reactions when spot tested. Most species are found in tropical and subtropical regions around the world, commonly growing on tree bark in mountain forests. Heterodermia species support a rich community of specialized parasitic fungi, with over 20 different species known to grow specifically on these lichens. Some species have traditional uses in medicine and cooking, particularly in India, Nepal, and Brazil.

Taxonomy
According to Fernanda de Souza and colleagues, the first scientific study of the genus began in 1847 when Thomas Taylor described Parmelia diademata, a lichen that would eventually end up classified as a species of Heterodermia. The genus was formally circumscribed in 1868 by Italian botanist Vittore Benedetto Antonio Trevisan de Saint-Léon. The generic name, which combines the Greek heteros (other, different) and derma (a skin or hide), refers to the presence or absence of a lower cortex. Prior to this classification, Heterodermia species were placed in genus Anaptychia, an idea proposed by Syo Kurokawa in his 1962 monograph on the genus, until some studies showed that the presence of thick-walled spores and the presence of atranorin could be used as characters to separate the genera. In 1965 Josef Poelt split Anaptychia into two genera. Lichens that remained in Anaptychia had thin-walled spores with sculptured surfaces, while those transferred to Heterodermia had thick-walled spores with smooth surfaces. Only nine species remained in the original Anaptychia classification. Although most contemporary lichenologists accepted Poelt's classification, Kurokawa initially rejected these changes in 1973. He later reversed his position and in 1998 accepted Heterodermia as distinct from Anaptychia, transferring several species accordingly. In North America, members of Heterodermia are colloquially known as fringe lichens or centipede lichens. ==Description==
Description
Heterodermia typically forms a foliose thallus—occasionally shading towards a somewhat fruticose habit—that is continuous, lobate and irregular, or arranged in loose rosettes across. Neighbouring thalli can merge, producing broad radiating mats or tangled clumps. The lobes may lie separate or touch; they range from closely adnate and appressed to partly ascending and loosely attached, and are linear to linear-cuneate or spoon-shaped (spathulate). Branching is mostly dichotomous but can be irregular, and the margins are often fringed with cilia that are simple or densely branched. The upper surface is whitish-, grey- or yellow-grey, flat to convex—sometimes concave near the edge—and may be dull or glossy. It can carry isidia, soredia, or , but never pseudocyphellae. The cortex consists of longitudinally aligned hyphae (a , superficially like plant collenchyma). A is absent. The photobiont forms a continuous band above a well-defined white medulla that may be tinged yellow, orange or brown. The lower surface, which may lack a cortex or bear a prosoplectenchymatous one, is white to whitish grey and often darkens to purple-grey, grey-black, or partly yellow, orange or brown. Rhizines are white to black, simple to densely branched—sometimes long enough to project beyond the lobe margins—and only rarely absent. The ascomata (fruiting bodies) of Heterodermia are apothecial and (ringed by thallus-like tissue). They sit on the thallus surface () and are rounded, either or borne on a short stalk. The exposed hymenial surface, or , ranges from pale to dark brown or black; it can be concave or nearly flat and may appear frosted () or smooth. A exciple rims the disc—prominent or —and remains distinct throughout the apothecium's life. In section, the epihymenium is pale brown to brown-black, while the underlying hymenium is colourless. The is usually colourless, only rarely tinged pale yellow. Paraphyses branch toward the top, where their terminal cells broaden and turn brown. The asci are cylindrical to somewhat club-shaped—Lecanora-type—with eight ascospores. Their apex is amyloid and thick-walled, enclosing a clear axial body. The spores develop one or more (small vacuoles in the wall). They turn grey-brown to dark brown, are ellipsoidal to oblong or fusiform, and have a single septum that often causes a slight constriction. Walls are very thick; internal apical thickenings only appear after the septum forms. A is thin or absent, and the surface remains smooth. Conidiomata lie immersed in the thallus at first, later becoming emergent. Their cells form short, branched chains and produce conidia enteroblastically (inner-wall budding). The resulting conidia are bacilliform to short-cylindrical. The genus Physcia most closely resembles Heterodermia. Unlike Heterodermia, however, it has a differently structured upper cortex consisting of (which gives it a uniform, unoriented appearance), and its ascospores are different. ==Species==
Species
Recent estimates have placed the number of species in Heterodermia at about 115. As of July 2025, Species Fungorum accepts 66 species of Heterodermia, but does not yet account for several recent studies that have added many new species. • Heterodermia aduncaHeterodermia africanaHeterodermia amphilacinulata – Brazil • Heterodermia andinaHeterodermia angustiloba – Asia; Australia • Heterodermia antillarum – Australia; Central America; Caribbean; Africa; Galapagos Islands • Heterodermia apicalis – Brazil • Heterodermia archeriHeterodermia arvidssoniiHeterodermia badiaHeterodermia barbiferaHeterodermia borphyllidiataHeterodermia caesiosoraHeterodermia caneziae – Brazil • Heterodermia comosaHeterodermia coralloides – Asia; Australia • Heterodermia corcovadensisHeterodermia delicatula – Brazil • Heterodermia diademata – Australia; North America; Central America; South America; Africa; Asia • Heterodermia dissecta – Asia; Australia; Réunion • Heterodermia dissecticodiademata – Brazil • Heterodermia dissecticoflabellata – Brazil • Heterodermia domingensisHeterodermia erectaHeterodermia erinaceaHeterodermia exuberansHeterodermia flavodactyliza – Brazil • Heterodermia flavulifera – Brazil • Heterodermia follmanniiHeterodermia fragmentataHeterodermia galactophyllaHeterodermia granuliferaHeterodermia guzmanianaHeterodermia himalayanaHeterodermia hybocarponica – Australia • Heterodermia isidiophoraHeterodermia isidiophorella – Australia • Heterodermia kalbii – Brazil • Heterodermia koyanaHeterodermia koyanoides – Australia • Heterodermia labiata – Brazil • Heterodermia langdonianaHeterodermia linearis – Mexico • Heterodermia macrosoraliata – Brazil • Heterodermia minor – Brazil • Heterodermia mobergianaHeterodermia namaquanaHeterodermia neocomosaHeterodermia neocrocea – Brazil • Heterodermia neoleucomelaenaHeterodermia nigromarginata – Brazil • Heterodermia obscurataHeterodermia orientalis – China • Heterodermia papuanaHeterodermia parvaHeterodermia phyllalbicans – Brazil • Heterodermia pindurae – Rwanda • Heterodermia pinnataHeterodermia podocarpaHeterodermia pseudospeciosaHeterodermia queensberryiHeterodermia ramosociliataHeterodermia rubrotrichaHeterodermia sinocomosa – China • Heterodermia sorediosaHeterodermia spathuliferaHeterodermia speciosaHeterodermia spielmannii – Brazil • Heterodermia subcitrinaHeterodermia subcomosaHeterodermia sublinearis – Brazil • Heterodermia tabularis – Australia • Heterodermia tasmanica – Australia • Heterodermia tremulansHeterodermia upretiiHeterodermia urtasuniHeterodermia velata – Brazil • Heterodermia verdoniiHeterodermia verrucifera Several former Heteroderma species have been moved to Polyblastidium. This includes P. appendiculatum, P. casarettianum, P. chilense, P. corallophorum, P. dendriticum, P. fragilissimum, P. hypocaesium, P. propaguliferum, P. japonicum, P. magellanicum, P. microphyllum, P. neglectum, P. queenslandicum, P. squamulosum, P. subneglectum, P. togashii, and P. violostriatum. ==Habitat and distribution==
Habitat and distribution
Regional monographs of Heterodermia have been published for several tropical regions, including Africa (26 species), Australia (42 species), South America (33 species), Brazil (68 species), Thailand (39 species), Most species have a pantropical or subtropical distribution, although a few have ranges that extend into temperate or oceanic regions. Tropical montane forest are a common habitat for many species, and many grow on tree bark or twigs. A few species however, also grow on rock or compacted soil, while fewer yet occur only on these substrates. ==Ecology==
Chemistry
Careful application of the K spot test can be used to detect most of the diagnostic lichen products in Heterodermia, although thin-layer chromatography is usually essential for a definite identification. Because they occur in all species of Heterodermia, atranorin and zeorin have no diagnostic value. Norstictic acid (K+ yellow → orange-red), with or without connorstictic acid (or rarely only connorstictic), salazinic acid (K+ yellow → blood red), or dissectic acid are occasionally present, and their presence is a useful character at species level. Pigments are also diagnostic, even though the certain identification of the emodin derivatives is not always possible. The pigments are clustered in chemosyndromes (a biogenetically related set of major and minor natural metabolic products produced by a species) and there are only three different main chemosyndromes that can be distinguished by colour and K reaction. Additional terpenoids have also been recorded, including (spathulene, japonene, leucotylin, and several hopane derivatives. ==Uses==
Uses
There are some Heterodermia species that are used as components of traditional medicine, and as spices or flavouring agent. Heterodermia diademata is used by the Nepalese of Sikkim, who apply the thalli of this lichen to cuts to protect them from wetting and infection. This species is also used in Uttar Pradesh, India as a perfume ingredient. Several ethnic groups in Madhya Pradesh (a state in middle India) use the thallus of Heterodermia tremulans as a spice and flavouring agent in various dishes of vegetables and meat. The Pankararu people of Pernambuco State, Brazil, use H. galactophylla for treating digestive system-related problems such as diarrhea and vomiting and for treating epilepsy. ==References==
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