# Strigulaceae

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Family of lichen-forming fungi

Strigulaceae Strigula species on plant foliage in Hawaii (possibly Strigula elegans) Scientific classification Kingdom: Fungi Division: Ascomycota Class: Dothideomycetes Order: Strigulales Family: Strigulaceae Zahlbr., 1898 Type genus Strigula Zahlbr. (1898) Synonyms[1] Phyllobatheliaceae Bitter & F.Schill. (1927)

***Strigulaceae*** is a [family](/source/Family_(taxonomy)) of mostly [lichen](/source/Lichen)-forming [fungi](/source/Fungi), one of two families in the order [Strigulales](/source/Strigulales) (class [Dothideomycetes](/source/Dothideomycetes)).[2] A [molecular analysis](/source/Molecular_phylogenetics) of the [type genus](/source/Type_genus), *[Strigula](/source/Strigula)*, has led to a reallocation of the [foliicolous](/source/Foliicolous_lichen) species into six genera that correspond to well-delimited [clades](/source/Clade) with diagnostic [phenotype](/source/Phenotype) features. These lichens live almost exclusively in [tropical](/source/Tropical) rainforests, where they grow as thin films on the surface of living leaves rather than on bark or rock like many other lichens. The family includes around 140 species distributed across multiple continents, making them one of the most widespread groups of leaf-dwelling organisms. DNA studies have revealed that what scientists once thought was a single large genus actually represents several distinct evolutionary [lineages](/source/Lineage_(evolution)).

## Taxonomy

The family Strigulaceae was erected by [Alexander Zahlbruckner](/source/Alexander_Zahlbruckner) in 1898 for a mixed assortment of leaf-inhabiting [pyrenocarpous](/source/Glossary_of_lichen_terms#pyrenocarpous) lichens. Mid-twentieth-century revisions restricted the concept to three epiphyllous genera—*[Strigula](/source/Strigula)*, *[Phylloporis](/source/Phylloporis)* and *[Raciborskiella](/source/Raciborskiella)*—but subsequent authors merged the latter two into an expanded *Strigula* and went on to add [corticolous](/source/Corticolous_lichen) (bark-dwelling) and [saxicolous](/source/Saxicolous_lichen) (rock-dwelling) species as well as the genera *[Phyllobathelium](/source/Phyllobathelium)*, *[Phyllocratera](/source/Phyllocratera)* and *[Flavobathelium](/source/Flavobathelium)* on [morphological](/source/Morphology_(biology)) grounds. Although multigene surveys in the late 2000s showed that these satellite taxa belonged inside Strigulaceae, they also hinted that *Strigula* *sensu lato* (in the loose sense) was [paraphyletic](/source/Paraphyletic) and that the delimitation of genera within the family required re-examination. Strigulaceae is placed in the order [Strigulales](/source/Strigulales) (class [Dothideomycetes](/source/Dothideomycetes)) and forms the core lichenised [lineage](/source/Lineage_(evolution)) of that order; its closest relative is the recently described family [Tenuitholiascaceae](/source/Tenuitholiascaceae).[3]

A broad multilocus [phylogeny](/source/Phylogeny) published in 2020 analysed 65 specimens representing 27 foliicolous species of *Strigula sensu lato* using various genetic markers. The study recovered six strongly supported [clades](/source/Clade) that correlate with clear [morphological](/source/Morphology_(biology)) and ecological characters, and it treated each clade at generic rank. The [type species](/source/Type_species), *[S. smaragdula](https://en.wikipedia.org/w/index.php?title=Strigula_smaragdula&action=edit&redlink=1)*, anchors *Strigula* *sensu stricto* (in the strict sense); the novel *[Serusiauxiella](/source/Serusiauxiella)* groups species with rapidly elongating macroconidial appendages and a *Trentepohlia*-like photobiont; *[Raciborskiella](/source/Raciborskiella)* comprises hypophyllous taxa with large ascospores; *[Puiggariella](/source/Puiggariella)* accommodates species with pale, non-carbonised perithecia and papillose thalli; *[Racoplaca](/source/Racoplaca)* covers the *S. subtilissima* complex characterised by olive-brown, finely lobed thalli edged by a black line; and *[Phylloporis](/source/Phylloporis)* was reinstated for supracuticular species allied to a *[Phycopeltis](/source/Phycopeltis)* photobiont. Together these six genera constitute a monophyletic foliicolous lineage within Strigulaceae, and the authors transferred or recombined more than a dozen species names to reflect the revised framework.[3]

The same analysis showed that the only sequenced non-foliicolous taxon, *[Strigula jamesii](https://en.wikipedia.org/w/index.php?title=Strigula_jamesii&action=edit&redlink=1)*, does not belong to this core lineage but instead forms a separate clade together with *Flavobathelium* and *Phyllobathelium*. This finding implies that additional saxicolous and corticolous species presently kept in *Strigula* will require reassignment once their DNA data become available. As of 2020, therefore, Strigulaceae included at least the nine lichenised genera *Strigula*, *Serusiauxiella*, *Raciborskiella*, *Puiggariella*, *Racoplaca*, *Phylloporis*, *Flavobathelium*, *Phyllobathelium* and *Phyllocratera*, while the status of non-lichenised *[Oletheriostrigula](/source/Oletheriostrigula)* remained doubtful pending further study.[3]

## Description

*[Racoplaca melanobapha](/source/Racoplaca_melanobapha)*

Members of Strigulaceae are tiny, leaf-inhabiting lichens that usually form thin, crust-like films rather than the [leafy](/source/Foliose_lichen) or [shrubby](/source/Fruticose_lichen) growth seen in many [temperate](/source/Temperate) lichens. Most species live inside the outer [cuticle](/source/Plant_cuticle) of living leaves (a [habit](/source/Habit_(biology)) called subcuticular), so their thalli show up only as faint green to grey patches or delicate olive-brown lobes on the leaf surface; in a few genera the lichen sits on top of the cuticle (supracuticular) and can be peeled away with a needle . Four easily recognised thallus patterns match the main genetic lineages: thick bright-green crusts (*Strigula* sensu stricto and *Serusiauxiella*), paper-thin bluish films on the leaf underside (*Raciborskiella*), folded pale-green mats with tiny white pimples (*Puiggariella*), and olive-brown, lobe-edged patches outlined by a hair-line black border (*Racoplaca*). Because these lichens are so inconspicuous and confined to tropical forests, many casual observers overlook them entirely.[3]

Microscopically the family is unified by a *Strigula*-type [ascus](/source/Ascus)—a tiny, double-walled ([bitunicate](/source/Glossary_of_lichen_terms#bitunicate)) spore sac whose tip contains a short clear plug (the ocular chamber); this character separates Strigulaceae from its closest relative, [Tenuitholiascaceae](/source/Tenuitholiascaceae). Each ascus releases eight colourless [ascospores](/source/Ascospore), usually with a single cross-wall ([septum](/source/Septum)) and measuring 7–70 [micrometres](/source/Micrometre) (μm) depending on the genus; in *Raciborskiella* the spores are at the larger end of this range and bear gelatinous "tails" that probably aid in sticking to new leaves . The flask-shaped fruit bodies ([perithecia](/source/Perithecia)) that house the asci are minute black dots, their walls commonly [carbonised](/source/Glossary_of_lichen_terms#carbonised) (hardened and darkened) except in *Puiggariella*. Asexual spores ([macroconidia](/source/Conidium)) are rod-shaped and tipped with mucilaginous appendages; in most genera these swell slowly after a day in water, but in *Serusiauxiella* they elongate to roughly 70 μm within an hour—a rapid-fire trick thought to help the spores glue themselves to the slippery leaf surface. Chemical screening shows no detectable [lichen products](/source/Lichen_product), so chemical [spot tests](/source/Spot_test_(lichen)) and [thin-layer chromatography](/source/Thin-layer_chromatography) are uninformative for this family.[3]

## Photobiont

Strigulaceae lichens partner with just a few groups of [green algae](/source/Green_alga). The majority harbour *[Cephaleuros](/source/Cephaleuros)*, a filamentous alga that threads through the leaf cuticle and makes it difficult to lift the lichen away; *Serusiauxiella* instead teams up with a *[Trentepohlia](/source/Trentepohlia_(alga))*-like alga, while the supracuticular genus *Phylloporis* associates with *[Phycopeltis](/source/Phycopeltis)*, whose flat, radiating plates let the lichen detach cleanly.[3]

## Genera

*[Phylloporis palmae](https://en.wikipedia.org/w/index.php?title=Phylloporis_palmae&action=edit&redlink=1)*

- *[Dichoporis](/source/Dichoporis)* Clem. (1909)[4] – 20 spp.

- *[Flagellostrigula](/source/Flagellostrigula)* Lücking, S.H.Jiang & Sérus. (2020)[5] – 2 spp.

- *[Flavobathelium](/source/Flavobathelium)* Lücking, Aptroot & G.Thor (1997)[6] – 1 sp.

- *[Oletheriostrigula](/source/Oletheriostrigula)* Huhndorf & R.C.Harris (1996)[7] – 1 sp.

- *[Phyllobathelium](/source/Phyllobathelium)* (Müll.Arg.) Müll.Arg. (1890)[8] – 8 spp.

- *[Phyllocharis](/source/Phyllocharis_(lichen))* Fée – 1 sp.

- *[Phyllocraterina](/source/Phyllocraterina)* Sérus. & Aptroot (2020)[5]

- *[Phylloporis](/source/Phylloporis)* Clem. (1909) – 6 spp.

- *[Puiggariella](/source/Puiggariella)* Speg. (1881)[9] – 3 spp.

- *[Raciborskiella](/source/Raciborskiella)* Höhn. (1909)[10] – 2 spp.

- *[Racoplaca](/source/Racoplaca)* Fée (1825)[11] – 5 spp.

- *[Serusiauxiella](/source/Serusiauxiella)* S.H.Jiang, Lücking & J.C.Wei (2020)[3] – 3 spp.

- *[Strigula](/source/Strigula)* Fr. (1823)[12] – ca. 60 spp.

- *[Swinscowia](/source/Swinscowia)* S.H.Jiang, Lücking & Sérus. (2020)[5] – 34 spp.

## References

1. **[^](#cite_ref-2024_Outline_1-0)** Hyde, K.D.; Noorabadi, M.T.; Thiyagaraja, V.; He, M.Q.; Johnston, P.R.; Wijesinghe, S.N.; et al. (2024). ["The 2024 Outline of Fungi and fungus-like taxa"](https://www.researchgate.net/publication/385939154). *Mycosphere*. **15** (1): 5146–6239 [5225]. [doi](/source/Doi_(identifier)):[10.5943/mycosphere/15/1/25](https://doi.org/10.5943%2Fmycosphere%2F15%2F1%2F25). [hdl](/source/Hdl_(identifier)):[11584/429245](https://hdl.handle.net/11584%2F429245).

1. **[^](#cite_ref-Wijayawardene_et_al._2020_2-0)** 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"](https://doi.org/10.5943%2Fmycosphere%2F11%2F1%2F8). *Mycosphere*. **11**: 1060–1456. [doi](/source/Doi_(identifier)):[10.5943/mycosphere/11/1/8](https://doi.org/10.5943%2Fmycosphere%2F11%2F1%2F8). [hdl](/source/Hdl_(identifier)):[10481/61998](https://hdl.handle.net/10481%2F61998).

1. ^ [***a***](#cite_ref-Jiang_et_al._2020_3-0) [***b***](#cite_ref-Jiang_et_al._2020_3-1) [***c***](#cite_ref-Jiang_et_al._2020_3-2) [***d***](#cite_ref-Jiang_et_al._2020_3-3) [***e***](#cite_ref-Jiang_et_al._2020_3-4) [***f***](#cite_ref-Jiang_et_al._2020_3-5) [***g***](#cite_ref-Jiang_et_al._2020_3-6) Jiang, Shu-Hua; Lücking, Robert; Xavier-Leite, Amanda Barreto; Cáceres, Marcela E. S.; Aptroot, André; Portilla, Carlos Viñas; Wei, Jiang-Chun (2020). ["Reallocation of foliicolous species of the genus *Strigula* into six genera (lichenized Ascomycota, Dothideomycetes, Strigulaceae)"](https://doi.org/10.1007%2Fs13225-020-00445-7). *Fungal Diversity*. **102** (1): 257–291. [doi](/source/Doi_(identifier)):[10.1007/s13225-020-00445-7](https://doi.org/10.1007%2Fs13225-020-00445-7).

1. **[^](#cite_ref-Clements_1909_4-0)** Clements, Frederic E. (1909). [*The genera of Fungi*](https://www.biodiversitylibrary.org/page/2892734). Minneapolis: H.W. Wilson Co. pp. 40, 173.

1. ^ [***a***](#cite_ref-Hongsanan_et_al._2020_5-0) [***b***](#cite_ref-Hongsanan_et_al._2020_5-1) [***c***](#cite_ref-Hongsanan_et_al._2020_5-2) Hongsanan, Sinang; Hyde, Kevin D.; Phookamsak, Rungtiwa; Wanasinghe, Dhanushka N.; McKenzie, Eric H. C.; Sarma, V. Venkateswara; et al. (2020). ["Refined families of Dothideomycetes: orders and families incertae sedis in Dothideomycetes"](https://doi.org/10.1007%2Fs13225-020-00462-6). *Fungal Diversity*. **105** (1): 17–318. [doi](/source/Doi_(identifier)):[10.1007/s13225-020-00462-6](https://doi.org/10.1007%2Fs13225-020-00462-6).

1. **[^](#cite_ref-Lücking_et_al._1997_6-0)** Lücking, Robert; Aptroot, André; Thor, Göran (1997). "New species or interesting records Of foliicolous lichens. II. *Flavobathelium epiphyllum* (Lichenized Ascomycetes: Melanommatales)". *The Lichenologist*. **29** (3): 221–228. [doi](/source/Doi_(identifier)):[10.1006/lich.1996.0079](https://doi.org/10.1006%2Flich.1996.0079).

1. **[^](#cite_ref-Huhndorf_&_Harris_1996_7-0)** Huhndorf, S.M.; Harris, R.C. (1996). "*Oletheriostrigula*, a new genus for *Massarina papulosa* (Fungi, ascomycetes)". *Brittonia*. **48** (4): 551–555. [Bibcode](/source/Bibcode_(identifier)):[1996Britt..48..551H](https://ui.adsabs.harvard.edu/abs/1996Britt..48..551H). [doi](/source/Doi_(identifier)):[10.2307/2807875](https://doi.org/10.2307%2F2807875). [JSTOR](/source/JSTOR_(identifier)) [2807875](https://www.jstor.org/stable/2807875).

1. **[^](#cite_ref-Müller_1890_8-0)** Müller, J. (1890). ["Lichenologische Beiträge XXXIII"](https://www.biodiversitylibrary.org/page/71824). *Flora (Regensburg)* (in Latin). **73** (2): 187–202 [195].

1. **[^](#cite_ref-Spegazzini_1881_9-0)** Spegazzini, C. (1881). "Fungi argentini additis nonnullis brasiliensibus montevideensibusque. Pugillus quartus (Continuacion)". *Anales de la Sociedad Científica Argentina*. **12** (3): 97–117.

1. **[^](#cite_ref-Höhnel_1909_10-0)** Höhnel, F. von (1909). "Fragmente zur Mykologie: VIII. Mitteilung (Nr. 354 bis 406)". *Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften. Wien. Mathematisch-naturwissenschaftliche Classe. Abteilung 1* (in German). **118**: 1157–1246.

1. **[^](#cite_ref-Fée_1825_11-0)** Fée, A.L.A. (1825). *Essai sur les cryptogames des écorces exotiques ofcinales* (in French). Paris: F. Didot. pp. lxviii, xciv, xcix.

1. **[^](#cite_ref-Fries_1823_12-0)** Fries, E.M. (1823). *Systema Mycologicum* (in Latin). Vol. 2. p. 535.

Taxon identifiers Strigulaceae Wikidata: Q10681823 Wikispecies: Strigulaceae AusLichen: 30016213 CoL: GQZ EoL: 6188 EPPO: 1SRGLF GBIF: 1945 iNaturalist: 174950 IndexFungorum: 81870 IRMNG: 110339 ITIS: 500137 MycoBank: 81870 NBN: NHMSYS0001499165 NCBI: 714593 NZOR: 3310e13c-4166-4e76-9907-1d8f1921ab9a Open Tree of Life: 994303 SpeciesFungorum: 81870 Tropicos: 100474473 Phyllobatheliaceae Wikidata: Q7188649 Wikispecies: Phyllobatheliaceae EoL: 6198 EPPO: 1PYBTF GBIF: 2470 IndexFungorum: 81159 IRMNG: 108138 ITIS: 612992 MycoBank: 81159 Open Tree of Life: 4082896 SpeciesFungorum: 81159

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