# Graphis (lichen)

Graphis is a genus of lichen-forming fungi in the family Graphidaceae. Historically, Graphis was used as a broad category for species with colourless, transversely septate ascospores within the Graphidaceae. However, with advances in genetic research, this classification has become more refined. As a result, species previously classified under Graphina have been re-assigned to Allographa or Graphis. The species complex around Graphis scripta has also been recognised, leading to the identification of several new species, many of which may have been previously overlooked.

## Description

Genus Graphis includes crustose lichens, which have a crust-like appearance that can range from being fully embedded in the substrate to sitting on the surface. The lichen's symbiotic partner, or photobiont, is green algae from the genus Trentepohlia. The apothecia (fruiting bodies) of Graphis can also be immersed or superficial. These structures are typically elongated, resembling slits (referred to as lirelliform), and can be simple, branched, or star-shaped. The disc of the apothecium, where spores are released, often remains slit-like or closed. Unlike some lichens, Graphis lacks a thalline exciple, which is a rim of tissue derived from the lichen thallus surrounding the apothecium. Instead, it has a true exciple, which is usually black, opaque, and well-developed. This exciple is sometimes marked by longitudinal grooves.
Inside the apothecium, the hymenium (a spore-bearing layer) is colourless and does not react to staining with iodine (I-). Beneath this layer, the hypothecium can be either pale or dark, and it is relatively thin. The hamathecium, which supports the developing spores, consists of unbranched filaments called paraphyses. The spore-producing structures, the asci, typically contain up to eight spores. These asci are club-shaped to slightly cylindrical and release their spores through an apical split. The spores themselves are divided by transverse walls (septate) or have a muriform structure (multiple divisions), and they turn violet when stained with iodine as they mature. Initially colourless, these spores may darken to brown if they become over-mature. Graphis also produces conidia, which are asexual spores, within flask-shaped structures called pycnidia. The conidia are usually cylindrical to ellipsoidal and remain colourless.
Chemically, some species of Graphis contain compounds known as β-orcinol depsidones. Additionally, older apothecia in certain species might react with potassium hydroxide solution (K+) to produce a purple colouration, indicating the presence of anthraquinones.
Graphis can be distinguished from the genus Phaeographis by its colourless spores, which may become brown with age. Additionally, in the field, Graphis species may resemble Opegrapha species, but the latter can be differentiated by their distinctively structured asci and spore-bearing tissues.

## Habitat and distribution

Graphis has a cosmopolitan distribution, which includes most continents across the world. This includes Florida in North America. Ecologically, Graphis lichens are mostly found growing on bark, though they are occasionally found on rock. The genus is primarily distributed in tropical and subtropical regions, with a few extending into temperate zones.

## Species

As accepted by Species Fungorum;
* Graphis alba J.C.Dantas, Lücking & M.Cáceres (2017)
* Graphis albidofarinacea Adaw. & Makhija (2006)
* Graphis albissima Müll.Arg. (1895)
* Graphis albocarpa J.Kalb, Lücking & Kalb (2018)
* Graphis alboglaucescens Adaw. & Makhija (2007)
* Graphis alboscripta Coppins & P.James (1992)
* Graphis allugallenensis Weerakoon, Wijey. & Lücking (2012)
* Graphis alpestris (Zahlbr.) Staiger (2002)
* Graphis amaliana Amórtegui, B.Moncada & Lücking (2019)
* Graphis analoga Nyl. (1859)
* Graphis andamanica Swarnal. (2017)
* Graphis anfractuosa (Eschw.) Eschw. (1833)
* Graphis apertella A.W.Archer (2001)
* Graphis aphaneomicrospora (Makhija & Adaw.) J.Kalb & Kalb (2017)
* Graphis aphanes Mont. & Bosch (1856)
* Graphis apoda (Zahlbr.) Lücking (2009)
* Graphis appendiculata Common & Lücking (2011)
* Graphis arbusculiformis (Vain.) Lücking (2012)
* Graphis archeri Dal-Forno & Eliasaro (2010)
* Graphis argentea Makhija & Adaw. (2005)
* Graphis australosiamensis Sutjar. & Kalb (2014)
* Graphis awasthii Kr.P. Singh & Swarnal. (2008)
* Graphis balaghatensis (Adaw. & Makhija) J.Kalb & Kalb (2017)
* Graphis bernadetiae van den Boom & Sipman (2014)
* Graphis bipartita (Müll.Arg.) Lücking (2008)
* Graphis boliviana Kukwa & Flakus (2013)
* Graphis borealis (A.W.Archer) A.W.Archer (2005)
* Graphis brahmanensis Aptroot (2009)
* Graphis breussii G.Neuwirth & Lücking (2009)
* Graphis brevicarpa M.Nakan., Kashiw. & K.H. Moon (2002)
* Graphis brittoniae Seavey & J.Seavey (2011)
* Graphis bungartzii A.B.Peña, Lücking, Herrera-Camp. & R.Miranda (2014)
* Graphis cambodiensis M.Nakan., Kashiw. & K.H.Moon (2010)
* Graphis caribica Lücking (2011)
* Graphis carmenelisana B.Moncada, Motta & Lücking (2019)
* Graphis catherinae A.W.Archer (2001)
* Graphis centrifuga Räsänen (1949)
* Graphis chondroplaca (Redinger) Lücking (2008)
* Graphis cincta (Pers.) Aptroot (2009)
* Graphis cinnamomea Adaw. & Makhija (2006)
* Graphis coenensis A.W.Archer (2007)
* Graphis colliculoides Adaw. & Makhija (2006)
* Graphis collinsiae Lücking & Lumbsch (2010)
* Graphis consociata (Makhija & Dube) Lücking & Kalb (2018)
* Graphis crassilabra Müll.Arg. (1882)
* Graphis crebra
* Graphis cremicolor (H.Magn.) Lücking & A.W. Archer (2009)
* Graphis crocea (Makhija & Adaw.) Lücking & Kalb (2018)
* Graphis crystallifera (Redinger) Staiger (2002)
* Graphis cupei Vain. ex Lücking (2009)
* Graphis cycadicola A.W.Archer & Elix (2007)
* Graphis darjeelingensis (Dharne & Roych.) Kr.P. Singh & Swarnal. (2008)
* Graphis dendrogramma
* Graphis descissa Müll.Arg. (1895)
* Graphis desquamescens (Fée) Zahlbr. (1909)
* Graphis dichotoma (Müll.Arg.) Lücking (2008)
* Graphis discarpa A.W.Archer (2004)
* Graphis dispersa Makhija & Adaw. (2005)
* Graphis distincta Makhija & Adaw. (2005)
* Graphis dracaenae Vain. (1901)
* Graphis duplicata Ach. (1814)
* Graphis duplicatoinspersa Lücking (2008)
* Graphis eburnea Adaw. & Makhija (2007)
* Graphis eimeoensis A.W.Archer & Elix (2007)
* Graphis elegans (Borrer ex Sm.) Ach. (1814)
* Graphis elevata Seavey & J.Seavey (2011)
* Graphis emersa Müll.Arg. (1893)
* Graphis emersella J.Kalb, Lücking & Kalb (2018)
* Graphis endoxantha Nyl. (1861)
* Graphis enteroleuca (Ach.) Lücking (2009)
* Graphis epimelaeana Müll.Arg. (1893)
* Graphis evirescens (Redinger) Lücking (2009)
* Graphis ferrugineodisca Seavey & J.Seavey (2017)
* Graphis filiformis Adaw. & Makhija (2007)
* Graphis fissurinoides van den Boom & Sipman (2013)
* Graphis flavopalmicola Y.Joshi, Lücking & Hur (2010)
* Graphis flavovirens Makhija & Adaw. (2005)
* Graphis flosculifera Weerakoon, Lücking & Aptroot (2019)
* Graphis fulvescens M.Nakan., Kashiw. & K.H. Moon (2015)
* Graphis galactoderma (Zahlbr.) Lücking (2009)
* Graphis gloriosensis A.W.Archer & Elix (2007)
* Graphis gonimica
* Graphis guangdongensis Z.F.Jia & J.C. Wei (2009)
* Graphis halonata Kalb & Aptroot (2018)
* Graphis hatschbachii S.C.Feuerst. & Eliasaro (2014)
* Graphis hinnulea Seavey & J.Seavey (2011)
* Graphis hodgesiana Lendemer (2010)
* Graphis hongkongensis Wei Guo & Hur (2015)
* Graphis hunanensis (Zahlbr.) M.Nakan. & Kashiw. (2003)
* Graphis hyphosa Staiger (2002)
* Graphis hypocrellina Lücking & Chaves (2008)
* Graphis immersella Müll.Arg. (1895)
* Graphis immersicans A.W.Archer (2001)
* Graphis implicata Fée (1874)
* Graphis inamoena Zahlbr. (1928)
* Graphis indica J.Kalb & Kalb (2017)
* Graphis inopinata S.C.Feuerst. & Eliasaro (2014)
* Graphis inspersolongula Aptroot (2009)
* Graphis insulana (Müll.Arg.) Lücking & Sipman (2008)
* Graphis insularis Makhija & Adaw. (2005)
* Graphis inustuloides Lücking (2012)
* Graphis isidiza Adaw. & Makhija (2004)
* Graphis japonica (Müll.Arg.) A.W.Archer & Lücking (2009)
* Graphis jeanmuelleri Lücking (2009)
* Graphis jejuensis K.H.Moon, M.Nakan. & Kashiw. (2012)
* Graphis jinhuana Kalb & Z.F.Jia (2014)
* Graphis kakaduensis A.W.Archer (2001)
* Graphis kavintuca Motta, B.Moncada & Lücking (2019)
* Graphis khaojoneana J.Kalb, Lücking & Kalb (2018)
* Graphis khaoyaiensis Poengs. & Lumbsch (2019)
* Graphis kollaimalaiensis Adaw. & Makhija (2006)
* Graphis koltermaniae Seavey & J.Seavey (2017)
* Graphis koratensis Pitakpong, Kraichak & Lücking (2015)
* Graphis koreana S.Joshi & Hur (2013)
* Graphis kousyuensis (Horik. & M.Nakan.) Lücking (2009)
* Graphis kurokawae M.Nakan., Kashiw. & K.H. Moon (2008)
* Graphis leptaleocarpa A.W.Archer (2002)
* Graphis leptocarpa Fée (1825)
* Graphis leptocarpoides Makhija & Adaw. (2005)
* Graphis leptoclada Müll.Arg. (1882)
* Graphis leptotremoides Papong, Lücking & Kraichak (2014)
* Graphis librata C.Knight (1884)
* Graphis lindsayana Lücking & Diederich (2017)
* Graphis lineola Ach. (1810)
* Graphis litoralis Lücking, Sipman & Chaves (2008)
* Graphis lueckingiana S.Joshi, Upreti & Hur (2018)
* Graphis lueckingii Dal-Forno & Eliasaro (2010)
* Graphis luluensis A.W.Archer (2004)
* Graphis lurizana Lücking, B.Moncada & Celis (2018)
* Graphis lutea (Chevall.) Aptroot (2008)
* Graphis maharashtrana Chitale, Makhija & B.O.Sharma (2011)
* Graphis manipurensis Pushpi Singh & Kr.P.Singh (2014)
* Graphis maomingensis Meng Z.Yang & Lu L.Zhang (2018)
* Graphis maritima (A.W.Archer) A.W.Archer (2007)
* Graphis marusae A.B.Peña & Lücking (2011)
* Graphis medusula (Spreng.) Kremp. (1876)
* Graphis meghalayensis Adaw. & Makhija (2007)
* Graphis mellis-insulae S.C.Feuerst. & Eliasaro (2014)
* Graphis microsperma (Chitale, Makhija & B.O.Sharma) Lücking & Kalb (2018)
* Graphis mikuraensis Y.Ohmura & M.Nakan. (2016)
* Graphis mucronata
* Graphis murali-elegans Sipman (2014)
* Graphis muscicola (Kalb) Staiger (2002)
* Graphis myrtacea (Müll.Arg.) Lücking (2008)
* Graphis neeladriensis Mohabe, Bodicherla, Nayaka & A.M.Reddy (2016)
* Graphis negrosina (Vain.) Lücking (2009)
* Graphis neoelongata Lücking (2009)
* Graphis neoraensis Jagad. Ram & G.P. Sinha (2009)
* Graphis nerurensis Makhija, A.Dube, Adaw. & Chitale (2006)
* Graphis nigririmis (Nyl.) Müll.Arg. (1895)
* Graphis nigrocarpa Adaw. & Makhija (2006)
* Graphis nigroglobosa Sipman (2014)
* Graphis nilgiriensis Adaw. & Makhija (2006)
* Graphis nitida Nyl. (1858)
* Graphis norfolkensis A.W.Archer & Elix (2019)
* Graphis norsorediata Aptroot (2020)
* Graphis norstictica A.W.Archer & Lücking (2009)
* Graphis nudanorsticta Z.F.Jia & Q.D.Wang (2018)
* Graphis omiana J.Kalb, Lücking & Kalb (2018)
* Graphis palmicola Makhija & Adaw. (2005)
* Graphis pananensis Kalb & Z.F.Jia (2014)
* Graphis panhalensis (Patw. & C.R.Kulk.) Chitale, Makhija & B.O.Sharma (2011)
* Graphis papillifera Lücking, N.Marín & B.Moncada (2023) – Colombia
* Graphis paradisserpens Sipman & Lücking (2008)
* Graphis paradussii Z.F.Jia (2011)
* Graphis paralleloides M.Cáceres & Lücking (2009)
* Graphis paranaensis Dal-Forno & Eliasaro (2010)
* Graphis paraschiffneri Lücking & Breuss (2015)
* Graphis paraserpens Lizano & Lücking (2008)
* Graphis pernambucoradians M.Cáceres & Lücking (2007)
* Graphis persulcata Stirt. (1879)
* Graphis pertricosa (Kremp.) A.W.Archer (2005)
* Graphis plumbeidisca M.Nakan., Kashiw. & K.H. Moon (2014)
* Graphis propinqua Müll.Arg. (1882)
* Graphis pseudoglyphis Lücking, N.Marín & B.Moncada (2023) – Colombia
* Graphis pseudoserpens Chaves & Lücking (2008)
* Graphis pseudosophistica
* Graphis puiggarii (Müll.Arg.) Lücking (2008)
* Graphis pustulosa M.Cáceres, Aptroot & Lücking (2014)
* Graphis pyrrhocheiloides
* Graphis rajapakshana Weerakoon, Lücking & Aptroot (2019)
* Graphis ramosa Nyl. (1891)
* Graphis rimosothallina Weerakoon, Lücking & Aptroot (2019)
* Graphis riopiedrensis (Fink) Lücking (2009)
* Graphis robertusii A.B.Peña, Herrera-Camp. & R.Miranda (2015)
* Graphis rondoniana M.Cáceres, Aptroot & Lücking (2014)
* Graphis rongklaensis Sutjar. (2012)
* Graphis rosae-emiliae A.B.Peña & Lücking (2014)
* Graphis rosalbinana B.Moncada, Amórtegui & Lücking (2019)
* Graphis salacinilongiramea Adaw. & Makhija (2007)
* Graphis santanderiana Motta, B.Moncada & Lücking (2019)
* Graphis saxicola (Müll.Arg.) A.W.Archer (2005)
* Graphis schummiana J.Kalb, Lücking & Kalb (2018)
* Graphis scripta (L.) Ach. (1809)
* Graphis seawardii Flakus & Kukwa (2013)
* Graphis semiaperta Müll.Arg. (1891)
* Graphis sirohiensis Pushpi Singh & Kr.P.Singh (2014)
* Graphis sitapurensis Makhija & Adaw. (2005)
* Graphis slendrae Hale ex Lücking (2009)
* Graphis solmariana Motta, B.Moncada & Lücking (2019)
* Graphis srilankensis Weerakoon, Wijey. & Lumbsch (2012)
* Graphis stellata M.Cáceres & Lücking (2007)
* Graphis stenospora Müll.Arg. (1882)
* Graphis stenotera Vain. (1921)
* Graphis stipitata A.W.Archer (2001)
* Graphis streimannii A.W.Archer (2001)
* Graphis subalbostriata Lücking (2009)
* Graphis subaltamirensis Passos, M.Cáceres & Lücking (2018)
* Graphis subargentata S.C.Feuerst.,Dal-Forno & Eliasaro (2014)
* Graphis subcontorta (Müll.Arg.) Lücking & Chaves (2008)
* Graphis subcupei S.C.Feuerst. & Eliasaro (2014)
* Graphis subelongata Papong, Lücking & Kraichak (2014)
* Graphis subfiliformis E.L.Lima & Lücking (2019)
* Graphis subhiascens (Müll.Arg.) Lücking (2008)
* Graphis subinsulana Poengs. & Kalb (2014)
* Graphis subintermedians Hale ex Lücking (2009)
* Graphis sublitoralis J.Kalb, Lücking & Kalb (2018)
* Graphis submarginata Lücking (2008)
* Graphis subregularis A.W.Archer (2001)
* Graphis subschroederi J.Kalb, Lücking & Kalb (2018)
* Graphis subtecta (Nyl.) Lücking (2009)
* Graphis subtenella Müll.Arg. (1887)
* Graphis subvittata Adaw. & Makhija (2007)
* Graphis sulphurella (Zahlbr.) Lücking (2009)
* Graphis sundarbanensis Jagad.Ram & G.P.Sinha (2007)
* Graphis supracola A.W.Archer (2001)
* Graphis suzanae Koch & S.C.Feuerst. (2016)
* Graphis syzygii Aptroot (2009)
* Graphis taneina M.Nakan., Kashiw. & K.H.Moon (2011)
* Graphis tenella Ach. (1814)
* Graphis tenoriensis Chaves & Lücking (2008)
* Graphis tenuirima (Shirley) A.W.Archer (2005)
* Graphis tetracarbonisata L.I.Ferraro, Lücking, Aptroot & M.Cáceres (2014)
* Graphis tetralocularis C.Bock & M.Hauck (2005)
* Graphis thunsinhalayensis Weerakoon, Arachchige & Lücking (2019)
* Graphis treubii Zahlbr. (1928)
* Graphis tricolor Aptroot & Feuerstein (2020)
* Graphis turgidula Müll.Arg. (1895)
* Graphis valparaiensis Adaw. & Makhija (2007)
* Graphis vermifera Müll.Arg. (1887)
* Graphis verrucata Z.F.Jia & Kalb (2014)
* Graphis verruciformis Adaw. & Makhija (2006)
* Graphis verrucoserpens Lücking (2019)
* Graphis vinosa Müll.Arg. (1895)
* Graphis xylophaga (R.C.Harris) Lendemer (2008)
* Graphis yunnanensis S.Joshi, Upreti & Hur (2015)