# Ingaderia

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***Ingaderia*** is a [genus](/source/Genus) of [lichen](/source/Lichen)-forming [fungi](/source/Fungi) in the family [Opegraphaceae](/source/Opegraphaceae).[1] Species in this genus form pale, [crusty patches](/source/Crustose_lichen) that often crack when thick, and produce elongated [fruiting bodies](/source/Fruiting_bodies) that appear as narrow slits on the surface. The genus was originally established with a single species from South America, but [molecular](/source/Molecular_phylogenetics) studies published in 2023 expanded its scope to include several species previously classified in other genera such as *[Llimonaea](/source/Llimonaea)* and *Paraingaderia*. Species are distinguished by their thick, dark [fruiting body](/source/Fruiting_body) walls, elongated [ascospores](/source/Ascospore) divided by [cross-walls](/source/Septum), and the presence of various [lichen products](/source/Lichen_product) including [erythrin](/source/Erythrin), [gyrophoric acid](/source/Gyrophoric_acid), or [lecanoric acid](/source/Lecanoric_acid). The genus comprises eight species found primarily in warmer climates, growing on rocks or bark.

## Taxonomy

The genus was [circumscribed](/source/Circumscription_(taxonomy)) by the British botanist [Otto Vernon Darbishire](/source/Otto_Vernon_Darbishire) in 1897. In his original description, Darbishire characterized *Ingaderia* as having a more or less rounded thallus without sharply defined margins, with lateral, oblong-pyriform [apothecia](/source/Apothecia) (fruiting bodies) that are or branched, possessing a hollow and but lacking a thallus sheath, and with apothecia often situated directly under the hypothecium in the gonidial medulla. He established the genus with the single species *Ingaderia pulcherrima*, citing *Roccella intricata* var. *alectoroides* as a [synonym](/source/Synonym_(taxonomy)), based on material from South America.[2]

Earlier work had suggested that the lirellae described on the [type species](/source/Type_species), *Ingaderia pulcherrima*, might belong to a [lichenicolous](/source/Lichenicolous_fungus) (lichen-dwelling) parasite rather than the lichen-forming fungus itself, so the true ascomata of the genus were often treated as unknown. Using new [molecular](/source/Molecular_phylogenetics) data, Ertz and Tehler showed that *I. pulcherrima* belongs near lirellate taxa with a thick black and , and argued that Darbishire's original description probably refers to the fruit bodies of the ; they noted that the ascospores match those seen in related members of the *Ingaderia* clade.[3]

In a two-locus phylogenetic study of the family [Opegraphaceae](/source/Opegraphaceae) published in 2023, Damien Ertz and Anders Tehler used nuclear large subunit [ribosomal DNA](/source/Ribosomal_DNA) (nuLSU) and RPB2 sequences to investigate relationships among lirellate genera in the family. Their [phylogenetic tree](/source/Phylogenetic_tree) recovered *Ingaderia pulcherrima* in a strongly supported [clade](/source/Clade)] together with species that had been placed in the separate genera *[Llimonaea](/source/Llimonaea)*, *Paraingaderia* and *[Fulvophyton](/source/Fulvophyton)*, including *Llimonaea flexuosa*, *L. occulta*, *L. sorediata*, *Paraingaderia placodioidea* and *Fulvophyton sorediatum*. Because these taxa share a thick dark excipulum and hypothecium, elongate multiseptate [ascospores](/source/Ascospore) and similar secondary chemistry, Ertz and Tehler adopted an enlarged generic concept in which *Llimonaea* and *Paraingaderia* are treated as [synonyms](/source/Synonym_(taxonomy)) of *Ingaderia*. They accordingly made the [new combinations](/source/New_combination) *Ingaderia flexuosa*, *I. occulta*, *I. placodioidea* and *I. sorediata*, and introduced the [replacement name](/source/Replacement_name) *Ingaderia vandenboomii* for *Llimonaea sorediata*; under this treatment *Paraingaderia* is no longer maintained as a separate genus.[3]

## Description

*Ingaderia* species form crust-like growths that adhere tightly to the rock or bark, but the margin can develop into small, shrubby projections and in some taxa the entire thallus becomes distinctly bushy ([fruticose](/source/Fruticose_lichen)). The surface is usually a pale cream or chalk-white, though fresh material may show a faint pink flush that weathers to grey-brown with age. Where the lichen grows thickly it often cracks, and many specimens are dusted with powdery reproductive granules called [soredia](/source/Soredia) that help the partner organisms disperse together. Under the [hand lens](/source/Hand_lens) the thallus looks rather than glossy because it is laden with minute [calcium oxalate](/source/Calcium_oxalate) crystals; these can form a continuous frost beneath or within the outer skin (), though some species lack a cortex entirely. The internal algal partner belongs to the orange-tinged genus *[Trentepohlia](/source/Trentepohlia_(alga))*, a common in tropical crusts.[4]

[Sexual](/source/Sexual_reproduction) fruit bodies are usually abundant and elongated. At first they lie buried in the thallus, but they soon erupt to form narrow slits or that may remain partially covered by surrounding tissue. Depending on the species the fruit bodies can remain straight, curve sinuously, or branch repeatedly into a tangled network; a light dusting of surface crystals () is optional. The dark outer wall () reacts with a drop of [potassium hydroxide](/source/Potassium_hydroxide) solution (the [K test](/source/K_test)) by becoming slightly darker or olive-tinged. Inside, the clear [hymenium](/source/Hymenium) contains eight slender [asci](/source/Ascus) whose tips stain blue with iodine, each releasing spindle-shaped [ascospores](/source/Ascospore) divided by one to several cross-walls; the spores thicken and turn brown over time and are wrapped in a gelatinous sheath that aids dispersal. Tiny, flask-shaped [pycnidia](/source/Pycnidia) immersed in the thallus provide an [asexual](/source/Asexual_reproduction) route: they exude colourless, hair-like [conidia](/source/Conidia) that curve gently like a sickle. [Secondary chemistry](/source/Secondary_metabolism) is variable but many species produce the [depside](/source/Depside) compounds [erythrin](/source/Erythrin), [gyrophoric acid](/source/Gyrophoric_acid), or [lecanoric acid](/source/Lecanoric_acid), while a few synthesise [psoromic acid](/source/Psoromic_acid).[4]

Ertz and Tehler argued that there are too few stable anatomical differences to justify keeping several small lirellate genera separate around *Ingaderia*. In their material, all species in the "*Llimonaea flexuosa–L. occulta*" lineage share a strongly developed dark brown to black excipulum and hypothecium, medium-sized, thick-walled, transversely septate ascospores and slender, curved conidia, whereas thallus [growth form](/source/Lichen_growth_forms), and secondary chemistry vary even within single species. They concluded that splitting this clade into several mostly [single-species](/source/Monotypic_taxon) genera would not be [morphologically](/source/Morphology_(biology)) meaningful, and instead adopted a single, phylogenetically defined *Ingaderia* that brings together crustose, , subfruticose and fruticose species with broadly similar lirellate ascomata and chemistry.[3]

## Species

- *[Ingaderia dendritica](/source/Ingaderia_dendritica)* [3]
- *[Ingaderia flexuosa](/source/Ingaderia_flexuosa)* [3]
- *[Ingaderia friabillima](/source/Ingaderia_friabillima)* [5]
- *[Ingaderia occulta](/source/Ingaderia_occulta)* [3]
- *[Ingaderia placodioidea](/source/Ingaderia_placodioidea)* [3]
- *[Ingaderia pulcherrima](/source/Ingaderia_pulcherrima)*
- *[Ingaderia sorediata](/source/Ingaderia_sorediata)* [3]
- *[Ingaderia vandenboomii](/source/Ingaderia_vandenboomii)* [3]

The species once named *Ingaderia troglodytica* is synonymous with *[Paralecanographa grumulosa](/source/Paralecanographa_grumulosa)*. The taxon once known as *Ingaderia gracillima* [6] has since been transferred to the genus *[Pentagenella](/source/Pentagenella)* as *[Pentegenella gracillima](/source/Pentegenella_gracillima)*.[7]

## References

1. Wijayawardene, Nalin; Hyde, Kevin; Al-Ani, Laith Khalil Tawfeeq; Somayeh, Dolatabadi; Stadler, Marc; Haelewaters, Danny et al. (2020). "Outline of Fungi and fungus-like taxa". *Mycosphere*. **11**: 1060–1456. [doi:10.5943/mycosphere/11/1/8](https://doi.org/10.5943/mycosphere/11/1/8). [hdl:10481/61998](https://hdl.handle.net/10481/61998)

1. Darbishire, O.V. (1897). ["Über die Flechtentribus der Roccellei"](https://www.biodiversitylibrary.org/page/5001353) [On the lichen tribe Roccellei] (in German). *Berichte der Deutschen Botanischen Gesellschaft*. **15**: 2–10.

1. Ertz, Damien & Tehler, Anders (2023). "New species of Arthoniales from Cape Verde with an enlarged concept of the genus *Ingaderia*". *The Lichenologist*. **55** (1): 1–15. [Bibcode:2023ThLic..55....1E](https://ui.adsabs.harvard.edu/abs/2023ThLic..55....1E). [doi:10.1017/S0024282922000408](https://doi.org/10.1017/S0024282922000408)

1. Cannon, P.; Coppins, B.; Sanderson, N.; Simkin, J. (2025). [*Various Arthoniales, including*Andreiomyces*(Andreiomycetaceae),*Chrysothrix*(Chrysotrichaceae),*Ingaderia*(Opegraphaceae),*Roccellographa*(Roccellographaceae) and*Bactrospora*,*Bryostigma*,*Felipes*,*Mixtoconidium*,*Perigrapha*and*Phacothecium*(family unassigned)*](https://britishlichensociety.org.uk/sites/default/files/Other%20Arthoniales.pdf). Vol. 53. Revisions of British and Irish Lichens. p. 5.

1. Follmann, G.; Schulz, M.; Werner, B. (1998). "On the identity and position of *Pentagenella fragillima*, *Roccellodea nigerrima*, and some related species (Roccellaceae, Opegraphales)". *Journal of the Hattori Botanical Laboratory*. **85**: 245–265.

1. Feige, G.B. & Lumbsch, T. (1993). "A European species of the lichen genus *Ingaderia* and comments on the relationships of the genera *Darbishirella* and *Ingaderia* (Roccellaceae)". *Mycotaxon*. **48**: 381–387.

1. ["Record Details: *Ingaderia gracillima* (Kremp.) Feige & Lumbsch, Mycotaxon 48: 386 (1993)"](https://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=357667). [Index Fungorum](/source/Index_Fungorum). Retrieved 25 November 2025.

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Adapted from the Wikipedia article [Ingaderia](https://en.wikipedia.org/wiki/Ingaderia) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Ingaderia?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.
