# Heterodermia

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***Heterodermia*** is a [genus](/source/Genus) of [foliose lichens](/source/Foliose_lichen) in the family [Physciaceae](/source/Physciaceae).[1] The genus has a widespread distribution, especially in [tropical](/source/Tropical) regions, and contains about 70 species. *Heterodermia* was historically confused with the related genus *[Anaptychia](/source/Anaptychia)*, but was distinguished in 1965 based on differences in [spore](/source/Spore) structure and chemical composition. These lichens can be identified in the field using simple chemical [spot tests](/source/Spot_test_(lichen)) that produce distinctive colour changes when applied to the [thallus](/source/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](/source/Cilia). The lichens can be identified by their thick-walled [ascospores](/source/Ascospore) and the presence of specific chemical compounds ([lichen products](/source/Lichen_product)) that produce colour reactions when [spot tested](/source/Spot_test_(lichen)). Most species are found in [tropical](/source/Tropical) and [subtropical](/source/Subtropical) regions around the world, commonly [growing on tree bark](/source/Corticolous_lichen) in [mountain forests](/source/Montane_ecosystem). *Heterodermia* species support a rich community of specialized [parasitic fungi](/source/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,[2] the first scientific study of the genus began in 1847 when [Thomas Taylor](/source/Thomas_Taylor_(botanist)) described *Parmelia diademata*,[3] a lichen that would eventually end up classified as a species of *Heterodermia*. The genus was formally [circumscribed](/source/Circumscription_(taxonomy)) in 1868 by Italian botanist [Vittore Benedetto Antonio Trevisan de Saint-Léon](/source/Vittore_Benedetto_Antonio_Trevisan_de_Saint-L%C3%A9on).[4] The generic name, which combines the [Greek](/source/Greek_language) *heteros* (other, different) and *derma* (a skin or hide), refers to the presence or absence of a lower [cortex](/source/Cortex_(botany)).[5]

Prior to this classification, *Heterodermia* species were placed in genus *[Anaptychia](/source/Anaptychia)*, an idea proposed by [Syo Kurokawa](/source/Syo_Kurokawa) in his 1962 monograph on the genus,[6] until some studies showed that the presence of thick-walled spores and the presence of [atranorin](/source/Atranorin) could be used as characters to separate the genera.[7][8] In 1965 [Josef Poelt](/source/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.[7] Although most contemporary lichenologists accepted Poelt's classification,[9][10][8][11] Kurokawa initially rejected these changes in 1973.[12] He later reversed his position and in 1998 accepted *Heterodermia* as distinct from *Anaptychia*, transferring several species accordingly.[13]

In North America, members of *Heterodermia* are [colloquially](/source/Common_name) known as fringe lichens or centipede lichens.[14]

## Description

*Heterodermia* typically forms a foliose [thallus](/source/Thallus)—occasionally shading towards a somewhat [fruticose](/source/Fruticose_lichen) habit—that is continuous, lobate and irregular, or arranged in loose [rosettes](/source/Rosette_(botany)) 2–10 cm (0.8–3.9 in) 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.[5]

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](/source/Isidia), [soredia](/source/Soredia), or , but never [pseudocyphellae](/source/Pseudocyphella). The cortex consists of longitudinally aligned hyphae (a , superficially like plant [collenchyma](/source/Collenchyma)). A is absent. The [photobiont](/source/Photobiont) forms a continuous band above a well-defined white [medulla](/source/Medulla_(lichenology)) 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](/source/Rhizine) are white to black, simple to densely branched—sometimes long enough to project beyond the lobe margins—and only rarely absent.[5]

The [ascomata](/source/Ascomata) (fruiting bodies) of *Heterodermia* are [apothecial](/source/Apothecia) 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.[5]

In section, the epihymenium is pale brown to brown-black, while the underlying [hymenium](/source/Hymenium) is colourless. The is usually colourless, only rarely tinged pale yellow. [Paraphyses](/source/Paraphyses) branch toward the top, where their terminal cells broaden and turn brown. The [asci](/source/Ascus) are cylindrical to somewhat club-shaped—*[Lecanora](/source/Lecanora)*-type—with eight [ascospores](/source/Ascospore). Their apex is [amyloid](/source/Amyloid_(mycology)) and thick-walled, enclosing a clear axial body. The spores develop one or more (small [vacuoles](/source/Vacuole) in the wall). They turn grey-brown to dark brown, are [ellipsoidal](/source/Ellipsoid) to oblong or fusiform, and have a single [septum](/source/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.[5]

[Conidiomata](/source/Conidiomata) lie immersed in the thallus at first, later becoming emergent. Their cells form short, branched chains and produce [conidia](/source/Conidia) enteroblastically (inner-wall budding). The resulting conidia are [bacilliform](/source/Bacilliform) to short-cylindrical.[5]

The genus *[Physcia](/source/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.[14]

## Species

Recent estimates have placed the number of species in *Heterodermia* at about 115.[15] As of July 2025, [Species Fungorum](/source/Species_Fungorum) accepts 66 species of *Heterodermia*, but does not yet account for several recent studies that have added many new species.[16]

- *[Heterodermia adunca](/source/Heterodermia_adunca)* [17]
- *[Heterodermia africana](/source/Heterodermia_africana)* [2]
- *[Heterodermia amphilacinulata](/source/Heterodermia_amphilacinulata)* [2] – Brazil
- *[Heterodermia andina](/source/Heterodermia_andina)* [18]
- *[Heterodermia angustiloba](/source/Heterodermia_angustiloba)* – Asia; Australia
- *[Heterodermia antillarum](/source/Heterodermia_antillarum)* – Australia; Central America; Caribbean; Africa; Galapagos Islands
- *[Heterodermia apicalis](/source/Heterodermia_apicalis)* [2] – Brazil
- *[Heterodermia archeri](/source/Heterodermia_archeri)* [19]
- *[Heterodermia arvidssonii](/source/Heterodermia_arvidssonii)* [18]
- *[Heterodermia badia](/source/Heterodermia_badia)* [18]
- *[Heterodermia barbifera](/source/Heterodermia_barbifera)*
- *[Heterodermia borphyllidiata](/source/Heterodermia_borphyllidiata)* [2]
- *[Heterodermia caesiosora](/source/Heterodermia_caesiosora)* [20]
- *[Heterodermia caneziae](/source/Heterodermia_caneziae)* [2] – Brazil
- *[Heterodermia comosa](/source/Heterodermia_comosa)*
- *[Heterodermia coralloides](/source/Heterodermia_coralloides)* [19] – Asia; Australia
- *[Heterodermia corcovadensis](/source/Heterodermia_corcovadensis)*
- *[Heterodermia delicatula](/source/Heterodermia_delicatula)* [2] – Brazil
- *[Heterodermia diademata](/source/Heterodermia_diademata)* – Australia; North America; Central America; South America; Africa; Asia
- *[Heterodermia dissecta](/source/Heterodermia_dissecta)* – Asia; Australia; Réunion
- *[Heterodermia dissecticodiademata](/source/Heterodermia_dissecticodiademata)* [2] – Brazil
- *[Heterodermia dissecticoflabellata](/source/Heterodermia_dissecticoflabellata)* [2] – Brazil
- *[Heterodermia domingensis](/source/Heterodermia_domingensis)*
- *[Heterodermia erecta](/source/Heterodermia_erecta)* [21]
- *[Heterodermia erinacea](/source/Heterodermia_erinacea)*
- *[Heterodermia exuberans](/source/Heterodermia_exuberans)* [20]
- *[Heterodermia flavodactyliza](/source/Heterodermia_flavodactyliza)* [2] – Brazil
- *[Heterodermia flavulifera](/source/Heterodermia_flavulifera)* [2] – Brazil
- *[Heterodermia follmannii](/source/Heterodermia_follmannii)*
- *[Heterodermia fragmentata](/source/Heterodermia_fragmentata)* [22]
- *[Heterodermia galactophylla](/source/Heterodermia_galactophylla)*
- *[Heterodermia granulifera](/source/Heterodermia_granulifera)*
- *[Heterodermia guzmaniana](/source/Heterodermia_guzmaniana)* [20]
- *[Heterodermia himalayana](/source/Heterodermia_himalayana)* [23]
- *[Heterodermia hybocarponica](/source/Heterodermia_hybocarponica)* [24] – Australia
- *[Heterodermia isidiophora](/source/Heterodermia_isidiophora)*
- *[Heterodermia isidiophorella](/source/Heterodermia_isidiophorella)* [19] – Australia
- *[Heterodermia kalbii](/source/Heterodermia_kalbii)* [25] – Brazil
- *[Heterodermia koyana](/source/Heterodermia_koyana)*
- *[Heterodermia koyanoides](/source/Heterodermia_koyanoides)* [19] – Australia
- *[Heterodermia labiata](/source/Heterodermia_labiata)* [2] – Brazil
- *[Heterodermia langdoniana](/source/Heterodermia_langdoniana)* [26]
- *[Heterodermia linearis](/source/Heterodermia_linearis)* [27] – Mexico
- *[Heterodermia macrosoraliata](/source/Heterodermia_macrosoraliata)* [2] – Brazil
- *[Heterodermia minor](/source/Heterodermia_minor)* [2] – Brazil
- *[Heterodermia mobergiana](/source/Heterodermia_mobergiana)* [28]
- *[Heterodermia namaquana](/source/Heterodermia_namaquana)*
- *[Heterodermia neocomosa](/source/Heterodermia_neocomosa)* [28]
- *[Heterodermia neocrocea](/source/Heterodermia_neocrocea)* [2] – Brazil
- *[Heterodermia neoleucomelaena](/source/Heterodermia_neoleucomelaena)*
- *[Heterodermia nigromarginata](/source/Heterodermia_nigromarginata)* [2] – Brazil
- *[Heterodermia obscurata](/source/Heterodermia_obscurata)*
- *[Heterodermia orientalis](/source/Heterodermia_orientalis)* [29] – China
- *[Heterodermia papuana](/source/Heterodermia_papuana)*
- *[Heterodermia parva](/source/Heterodermia_parva)* [18]
- *[Heterodermia phyllalbicans](/source/Heterodermia_phyllalbicans)* [2] – Brazil
- *[Heterodermia pindurae](/source/Heterodermia_pindurae)* [30] – Rwanda
- *[Heterodermia pinnata](/source/Heterodermia_pinnata)*
- *[Heterodermia podocarpa](/source/Heterodermia_podocarpa)*
- *[Heterodermia pseudospeciosa](/source/Heterodermia_pseudospeciosa)*
- *[Heterodermia queensberryi](/source/Heterodermia_queensberryi)* [31]
- *[Heterodermia ramosociliata](/source/Heterodermia_ramosociliata)* [28]
- *[Heterodermia rubrotricha](/source/Heterodermia_rubrotricha)* [32]
- *[Heterodermia sinocomosa](/source/Heterodermia_sinocomosa)* [29] – China
- *[Heterodermia sorediosa](/source/Heterodermia_sorediosa)* [28]
- *[Heterodermia spathulifera](/source/Heterodermia_spathulifera)*
- *[Heterodermia speciosa](/source/Heterodermia_speciosa)*
- *[Heterodermia spielmannii](/source/Heterodermia_spielmannii)* [2] – Brazil
- *[Heterodermia subcitrina](/source/Heterodermia_subcitrina)* [33]
- *[Heterodermia subcomosa](/source/Heterodermia_subcomosa)*
- *[Heterodermia sublinearis](/source/Heterodermia_sublinearis)* [2] – Brazil
- *[Heterodermia tabularis](/source/Heterodermia_tabularis)* [19] – Australia
- *[Heterodermia tasmanica](/source/Heterodermia_tasmanica)* [19] – Australia

- *[Heterodermia tremulans](/source/Heterodermia_tremulans)*
- *[Heterodermia upretii](/source/Heterodermia_upretii)* [34]
- *[Heterodermia urtasuni](/source/Heterodermia_urtasuni)* [35]
- *[Heterodermia velata](/source/Heterodermia_velata)* [36] – Brazil
- *[Heterodermia verdonii](/source/Heterodermia_verdonii)*
- *[Heterodermia verrucifera](/source/Heterodermia_verrucifera)*

Several former *Heteroderma* species have been moved to *[Polyblastidium](/source/Polyblastidium)*. This includes *[P. appendiculatum](/source/Polyblastidium_appendiculatum)*, *[P. casarettianum](/source/Polyblastidium_casarettianum)*, *[P. chilense](/source/Polyblastidium_chilense)*, *[P. corallophorum](/source/Polyblastidium_corallophorum)*, *[P. dendriticum](/source/Polyblastidium_dendriticum)*, *[P. fragilissimum](/source/Polyblastidium_fragilissimum)*, *[P. hypocaesium](/source/Polyblastidium_hypocaesium)*, *[P. propaguliferum](/source/Polyblastidium_propaguliferum)*, *[P. japonicum](/source/Polyblastidium_japonicum)*, *[P. magellanicum](/source/Polyblastidium_magellanicum)*, *[P. microphyllum](/source/Polyblastidium_microphyllum)*, *[P. neglectum](/source/Polyblastidium_neglectum)*, *[P. queenslandicum](/source/Polyblastidium_queenslandicum)*, *[P. squamulosum](/source/Polyblastidium_squamulosum)*, *[P. subneglectum](/source/Polyblastidium_subneglectum)*, *[P. togashii](/source/Polyblastidium_togashii)*, and *[P. violostriatum](/source/Polyblastidium_violostriatum)*.[15]

## Habitat and distribution

Regional [monographs](/source/Monograph) of *Heterodermia* have been published for several tropical regions, including Africa (26 species),[37][33] Australia (42 species),[5] South America (33 species),[18] Brazil (68 species),[2] Thailand (39 species),[15] Most species have a [pantropical](/source/Pantropical) or [subtropical](/source/Subtropical) distribution, although a few have ranges that extend into [temperate](/source/Temperate) or oceanic regions.[15] Tropical [montane](/source/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.[2]

## Ecology

*Heterodermia* species serve as [hosts](/source/Host_(biology)) for a diverse [community](/source/Community_(ecology)) of [lichenicolous fungi](/source/Lichenicolous_fungi)—specialized fungi that grow [parasitically](/source/Parasitism) or [commensally](/source/Commensalism) on lichens. These fungi represent one of the most species-rich groups of organisms associated with *Heterodermia*, with more than 20 documented species: *[Lichenostigma heterodermiae](/source/Lichenostigma_heterodermiae)*, *[Phoma heterodermiae](/source/Phoma_heterodermiae)*, *[Polycoccum heterodermiae](/source/Polycoccum_heterodermiae)*, *[Lichenotubeufia heterodermiae](/source/Lichenotubeufia_heterodermiae)*, *[Lichenopeltella heterodermiae](/source/Lichenopeltella_heterodermiae)*, *[Sphaerellothecium episoralium](/source/Sphaerellothecium_episoralium)*, *[Sphaerellothecium gallowayi](/source/Sphaerellothecium_gallowayi)*, *[Sphaerellothecium heterodermiae](/source/Sphaerellothecium_heterodermiae)*, *[Capronia muellerelloides](/source/Capronia_muellerelloides)*, *[Capronia solitaria](/source/Capronia_solitaria)*, *[Endococcus sipmanii](/source/Endococcus_sipmanii)*, *[Stigmidium heterodermiae](/source/Stigmidium_heterodermiae)*, *[Sclerococcum heterodermiae](/source/Sclerococcum_heterodermiae)*, *[Nanostictis heterodermiae](/source/Nanostictis_heterodermiae)*, *[Hyalopeziza heterodermiae](/source/Hyalopeziza_heterodermiae)*, *[Pleoscutula arsenii](/source/Pleoscutula_arsenii)*, *[Epicladonia heterodermiae](/source/Epicladonia_heterodermiae)*, *[Nectriopsis heterodermiae](/source/Nectriopsis_heterodermiae)*, *[Nectriopsis lichenophila](/source/Nectriopsis_lichenophila)*, *[Pronectria pycnidioidea](/source/Pronectria_pycnidioidea)*, and *[Neobarya ciliaris](/source/Neobarya_ciliaris)*.[38]

## Chemistry

Careful application of the K [spot test](/source/Spot_test_(lichen)) can be used to detect most of the diagnostic [lichen products](/source/Lichen_product) in *Heterodermia*, although [thin-layer chromatography](/source/Thin-layer_chromatography) is usually essential for a definite identification. Because they occur in all species of *Heterodermia*, [atranorin](/source/Atranorin) and [zeorin](/source/Zeorin) have no diagnostic value. [Norstictic acid](/source/Norstictic_acid) (K+ yellow → orange-red), with or without [connorstictic acid](/source/Connorstictic_acid) (or rarely only connorstictic), [salazinic acid](/source/Salazinic_acid) (K+ yellow → blood red), or [dissectic acid](/source/Dissectic_acid) are occasionally present, and their presence is a useful character at species level. [Pigments](/source/Biological_pigment) are also diagnostic, even though the certain identification of the [emodin](/source/Emodin) derivatives is not always possible. The pigments are clustered in chemosyndromes (a biogenetically related set of major and minor natural [metabolic products](/source/Secondary_metabolite) produced by a species) and there are only three different main chemosyndromes that can be distinguished by colour and K reaction. Additional [terpenoids](/source/Terpenoid) have also been recorded, including ([spathulene](/source/Spathulene), [japonene](/source/Japonene), [leucotylin](/source/Leucotylin), and several [hopane](/source/Hopane) derivatives.[2]

## Uses

There are some *Heterodermia* species that are used as components of [traditional medicine](/source/Traditional_medicine), and as spices or flavouring agent. *[Heterodermia diademata](/source/Heterodermia_diademata)* is used by the [Nepalese of Sikkim](/source/Sikkimese_people), who apply the thalli of this lichen to cuts to protect them from wetting and infection. This species is also used in [Uttar Pradesh](/source/Uttar_Pradesh), India as a perfume ingredient.[39] Several ethnic groups in [Madhya Pradesh](/source/Madhya_Pradesh) (a state in middle India) use the thallus of *[Heterodermia tremulans](/source/Heterodermia_tremulans)* as a spice and flavouring agent in various dishes of vegetables and meat.[40] The [Pankararu](/source/Pankararu) people of [Pernambuco State](/source/Pernambuco_State), Brazil, use *H. galactophylla* for treating digestive system-related problems such as diarrhea and vomiting and for treating [epilepsy](/source/Epilepsy).[41]

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