Dendrographa alectoroides, copyright Liam O’Brien.

Belongs within: Dothideomyceta.
Contains: Roccellina, Dirina, Roccella.

Lichens to dye for
Published 28 November 2022

Most lichen species, it is probably fair to say, are little recognized for their economic impact on humans. Among the most notable exceptions is Roccella tinctoria, a fruticose lichen found on rocks and cliffs on the coast of Portugal and northern Africa, as well as on the nearby Macaronesian islands. This species has long been used as the source of orchil, a red or purple dye. Orchil may be used to dye wool or silk without the need for a mordant and was particularly popular in Renaissance-era Europe. The Ruccellai family of Florence, one of that city’s wealthiest dynasties, derived their name from orchil (Tehler & Irestedt 2007). Prepared another way, the dye extracted from R. tinctoria forms litmus which many of you may recall being used as a pH indicator in high school chemistry.

Orchil Roccella tinctoria, copyright Norbert Nagel.

Roccella tinctoria is a representative of the Roccellaceae, a family of saxicolous and corticolous lichens found most abundantly in subtropical coastal habitats, particularly in arid and Mediterranean-climate regions. As currently circumscribed, Roccellaceae are typically characterised by a corticate thallus containing trentepohlioid photobionts, rounded ascomata with a thalline margin, a dark brown hypothecium, cylindrical-clavate asci, and ascospores lacking a gelatinous sheath (Ertz et al. 2015). Roccellaceae are representatives of the Arthoniomycetes, a distinct clade of fungi from the Lecanoromycetes to which belong the greater number of familiar lichens.

Dirina massiliensis, copyright Anita Gould.

The growth habit of Roccellaceae is varied, including both crustose and fruticose forms. Phylogenetic analyses indicate that changes in growth habit have occurred multiple times over the family’s evolution, from crustose to fruticose and back again (Tehler & Irestedt 2007). Species may reproduce sexually via the production of ascospores or asexually via the production of soredia, small ‘parcels’ containing fungal hyphae and associated algal photobionts. Production of both reproductive structures on a single thallus is rare, and often reproductive mode is used to distinguish species: for instance, Roccella tinctoria represents the asexual counterpart to the sexual R. canariensis.

Whereas lichen species are generally widespread, Roccellaceae are unusual in that species are often highly endemic. This may extend to higher taxa: for instance, the diverse genus Roccellina is almost entirely restricted to the Southern Hemisphere (Ertz et al. 2015). For Roccellaceae, at least, the classic proposal for micro-organisms that “everything is everywhere” simply does not apply.

Systematics of Roccellaceae
Roccellaceae [Bactrosporaceae, Lecanactidaceae, Opegraphaceae]
|--+--Llimonaea Egea & Torrente 1991TI07
| | |--*L. occulta Egea & Torrente 1991TI07
| | `--L. californicaTI07
| `--+--Combea Nylander 1860TI07
| | `--*C. mollusca (Ach.) Nylander 1860TI07
| `--+--Pentagenella Darbishire 1897 [incl. Camanchaca Follmann & Peine 1999]TI07
| | |--*P. fragillima Darbishire 1897 (see below for synonymy)TI07
| | |--P. corallina (Follmann & Peine) Tehler in Tehler & Irestedt 2007 (see below for synonymy)TI07
| | `--P. ligulata (Peine & Follmann) Tehler in Tehler & Irestedt 2007 [=Camanchaca ligulata]TI07
| `--+--Hubbsia Weber in Egan 1987TI07
| | |--*H. californica (Räs.) Weber in Egan 1987TI07
| | |--H. langeiTI07
| | `--H. parishiiTI07
| `--Ingaderia Darbishire 1897TI07, KC01 [incl. Darbishirella Zahlbr. 1898KC01, Dictyographa Müll. Arg. 1893KC01]
| |--*I. pulcherrima Darbishire 1897TI07
| |--I. friabillimaTI07
| |--I. gracillimaTI07
| `--I. troglodytisTI07
`--+--+--+--Lecanactis Körber 1855 (nom. cons.)TI07, KC01 (see below for synonymy)
| | | `--*L. abietina (Ach.) Körber 1855TI07
| | `--Schismatomma Flot. & Körb. ex Massal. 1852TI07, KC01 (see below for synonymy)
| | |--S. decoloransLK04
| | `--S. pericleum [incl. *S. dolosum Massalongo 1852]TI07
| `--+--RoccellinaTI07
| `--+--Syncesia Taylor 1836TI07, KC01
| | |--S. myrticola [incl. *S. albida Taylor 1836]TI07
| | |--S. farinaceaTI07
| | `--S. leprobolaTI07
| `--Dendrographa Darbishire 1895TI07, KC01
| |--*D. leucocephala (Tuck.) Darbishire 1895TI07
| | |--D. l. f. leucocephalaTI07
| | `--D. l. f. minorTI07
| `--D. alectoroidesTI07
| |--D. a. f. alectoroidesTI07
| `--D. a. f. parvaTI07

Roccellaceae incertae sedis:
Ancistroporella Thor 1995 [=Ancistrospora Thor 1991 non Menéndez & Azcuy 1972]KC01
Bactrospora Massal. 1852 (see below for synonymy)KC01
Chiodecton Ach. 1814 [incl. Melanodecton Massal. 1860]KC01
Cresponea Egea & Torrente 1993KC01
Dichosporidium Pat. 1903 [=Dichosporium Pat. 1899 non Nees 1816]KC01
Dirinastrum Müll. Arg. 1893 [=Dirinastromyces Cif. & Tomas. 1953]KC01
Dolichocarpus Sant. 1949KC01
Enterodictyon Müll. Arg. 1892 [=Enterodictyonomyces Cif. & Tomas. 1953, Enterodictyum Clem. & Shear 1931]KC01
Enterographa Fée 1824 (see below for synonymy)KC01
Erythrodecton Thor 1991KC01
Feigeana Mies, Lumbsch & Tehler 1995KC01
Follmanniella Peine & Werner 1995KC01
Gorgadesia Tav. 1964KC01
Graphidastra (Redinger) Thor 1990KC01
Halographis Kohlm. & Volkm.-Kohlm. 1988KC01
Haplodina Zahlbr. 1930KC01
Lecanographa Egea & Torrente 1994KC01
Mazosia Massal. 1854 (see below for synonymy)KC01
Minksia Müll. Arg. 1882 [incl. Cyrtographa Müll. Arg. 1894]KC01
Plectocarpon Fée 1825 (see below for synonymy)KC01
Pseudolecanactis Zahlbr. 1907KC01
Reinkella Darb. 1897 [=Reinkellomyces Cif. & Tomas. 1953]KC01
Roccellodea Darb. 1932KC01
Sagenidium Stirt. 1877KC01
Schizopelte Fr. 1875 [=Schizopeltomyces Cif. & Tomas. 1953]KC01
Sclerophyton Eschw. 1824 (see below for synonymy)KC01
Sigridea Tehler 1993KC01
Perigrapha Hafellner 1996 non Lederer 1857 (ICZN)KC01
Pulvinodecton Henssen & Thor 1998KC01
Sphaeromma Upadhyay 1964KC01
‘Streimannia’ Thor 1991 non Ochyra 1900KC01
Simonyella Steiner 1902SS09, KC01
`--S. variegataSS09
Opegrapha Ach. 1809 (nom. cons.)SS09, KC01 (see below for synonymy)
|--O. dolomiticaSS09
`--O. variaLH04
Roccellographa Steiner 1902SS09, KC01 [=Roccellographomyces Cif. & Tomas. 1953KC01]
`--R. cretaceaSS09

Bactrospora Massal. 1852 [=Bactrosporomyces Cif. & Tomas. 1953; incl. Melampydiomyces Cif. & Tomas. 1953, Melampydium Zahlbr. 1905, Melampylidium Stirt. ex Müll. Arg. 1894, Scolecactis Clem. 1909]KC01

Enterographa Fée 1824 [=Chiodectonomyces Cif. & Tomas. 1953; incl. Arthonaria Fr. 1825, Platygramma Leight. 1854 non Mey. 1825, Stigmatidium Mey. 1825]KC01

Lecanactis Körber 1855 (nom. cons.)TI07, KC01 [incl. Arthoniactis (Vain.) Clem. 1909KC01, Bacidiactis Choisy 1931KC01, Lecanactiomyces Cif. & Tomas. 1953KC01, Pyrenotea Fr. 1821 (nom. rej.)KC01]

Mazosia Massal. 1854 [=Rotula (Müll. Arg.) Müll. Arg. 1890 nec Lour. 1790 nec Raf. 1815 nec Schumacher 1817 (ICZN); incl. Micrographina Fink 1930, Rotularia (Vain.) Zahlbr. 1923 nec Sternb. 1825 nec Defrance 1827 (ICZN), Sporhaplus Upadhyay 1964]KC01

Opegrapha Ach. 1809 (nom. cons.)SS09, KC01 [incl. Alyxoria Gray 1821KC01, Aulaxinomyces Cif. & Tomas. 1953KC01, Fouragea Trevis. 1880KC01, Hysterina (Ach.) Gray 1821KC01, Leciographa Massal. 1854KC01, Lecoglyphis Clem. 1909KC01, Mycopegrapha Vain. 1921KC01, Opegraphella Müll. Arg. 1890KC01, Opegraphellomyces Cif. & Tomas. 1953KC01, Opegraphoidea Fink 1933KC01, Opegraphomyces Thomas ex Cif. & Tomas. 1953KC01, Phragmographum Henn. 1905KC01, Phyllographa (Müll. Arg.) Räsänen 1943 (nom. inv.)KC01, Plagiographis Knight & Mitt. 1860KC01, Scaphis Eschw. 1824KC01, Sclerographa (Vain.) Zahlbr. 1923KC01, Xylastra Massal. 1855KC01, Zwackhia Körb. 1855KC01]

Pentagenella corallina (Follmann & Peine) Tehler in Tehler & Irestedt 2007 [=*Camanchaca corallina Follmann & Peine 1999]TI07

*Pentagenella fragillima Darbishire 1897 [=Roccella fragillima (Darbishire) Follmann et al. 1998, R. fragilissima Mont. ex Follmann et al. 1998]TI07

Plectocarpon Fée 1825 [incl. Delisea Fée 1825 non Lamour. 1819, Epiphora Nyl. 1876 non Lindl. 1837, Lichenomyces Trevis. 1853]KC01

Schismatomma Flot. & Körb. ex Massal. 1852TI07, KC01 [=Schismatommatomyces Cif. & Tomas. 1953KC01; incl. Gomphospora Massal. 1852KC01, Platygrapha Nyl. 1855KC01, Platygraphis Hook. f. 1867KC01, Platygraphomyces Cif. & Tomas. 1953KC01]

Sclerophyton Eschw. 1824 [=Sclerophytonomyces Cif. & Tomas. 1953; incl. Stigmatella Mudd 1861 non Berk. & Curtis 1857]KC01

*Type species of generic name indicated


Ertz, D., A. Tehler, M. Irestedt, A. Frisch, G. Thor & P. van den Boom. 2015. A large-scale phylogenetic revision of Roccellaceae (Arthoniales) reveals eight new genera. Fungal Diversity 70: 31–53.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[LH04] Liu, Y. J., & B. D. Hall. 2004. Body plan evolution of ascomycetes, as inferred from an RNA polymerase II phylogeny. Proceedings of the National Academy of Sciences of the USA 101: 4507–4512.

[LK04] Lutzoni, F., F. Kauff, C. J. Cox, D. McLaughlin, G. Celio, B. Dentinger, M. Padamsee, D. Hibbett, T. Y. James, E. Baloch, M. Grube, V. Reeb, V. Hofstetter, C. Schoch, A. E. Arnold, J. Miadlikowska, J. Spatafora, D. Johnson, S. Hambleton, M. Crockett, R. Shoemaker, G.-H. Sung, R. Lücking, T. Lumbsch, K. O’Donnell, M. Binder, P. Diederich, D. Ertz, C. Gueidan, K. Hansen, R. C. Harris, K. Hosaka, Y.-W. Lim, B. Matheny, H. Nishida, D. Pfister, J. Rogers, A. Rossman, I. Schmitt, H. Sipman, J. Stone, J. Sugiyama, R. Yahr & R. Vilgalys. 2004. Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits. American Journal of Botany 91 (10): 1446–1480.

[SS09] Schoch, C. L., G.-H. Sung, F. López-Giráldez, J. P. Townsend, J. Miadlikowska, V. Hofstetter, B. Robbertse, P. B. Matheny, F. Kauff, Z. Wang, C. Gueidan, R. M. Andrie, K. Trippe, L. M. Ciufetti, A. Wynns, E. Fraker, B. P. Hodkinson, G. Bonito, J. Z. Groenewald, M. Arzanlou, G. S. de Hoog, P. W. Crous, D. Hewitt, D. H. Pfister, K. Peterson, M. Gryzenhout, M. J. Wingfield, A. Aptroot, S.-O. Suh, M. Blackwell, D. M. Hillis, G. W. Griffith, L. A. Castlebury, A. Y. Rossman, H. T. Lumbsch, R. Lücking, B. Büdel, A. Rauhut, P. Diederich, D. Ertz, D. M. Geiser, K. Hosaka, P. Inderbitzin, J. Kohlmeyer, B. Volkmann-Kohlmeyer, L. Mostert, K. O’Donnell, H. Sipman, J. D. Rogers, R. A. Shoemaker, J. Sugiyama, R. C. Summerbell, W. Untereiner, P. R. Johnston, S. Stenroos, A. Zuccaro, P. S. Dyer, P. D. Crittenden, M. S. Cole, K. Hansen, J. M. Trappe, R. Yahr, F. Lutzoni & J. W. Spatafora. 2009. The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits. Systematic Biology 58 (2): 224–239.

[TI07] Tehler, A., & M. Irestedt. 2007. Parallel evolution of lichen growth forms in the family Roccellaceae (Arthoniales, Ascomycota). Cladistics 23: 432–454.

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