# Phacopsis

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This article is about the genus of fungi. For the defunct legume genus, see [Astragalus (plant)](/source/Astragalus_(plant)).

***Phacopsis*** is a [genus](/source/Genus) of [lichenicolous](/source/Lichenicolous_fungi) ([lichen](/source/Lichen)-dwelling) [fungi](/source/Fungi). They are [parasites](/source/Parasitism) of members of the large lichen [family](/source/Family_(taxonomy)) [Parmeliaceae](/source/Parmeliaceae), of which they are also a member. Originally proposed by [Edmond Tulasne](/source/Edmond_Tulasne) in 1852 to contain 3 species, *Phacopsis* now contains 10 species, although historically, 33 [taxa](/source/Taxon) have been described in the genus. Many of the species are poorly known, some of them having been documented only from the [type specimen](/source/Type_(biology)).

*Phacopsis* species appear as partially immersed, shiny brown to black [apothecia](/source/Apothecia) that cause [gall](/source/Gall)-like deformations on the [thallus](/source/Thallus) of the [host](/source/Host_(biology)) lichen. Features of *Phacopsis* used to distinguish species from each other include the shape of their [spores](/source/Ascospore), and the colour and reaction of the hypothecium (a tissue layer under the spore-bearing [hymenium](/source/Hymenium)) when [stained](/source/Staining) using an [iodine–starch test](/source/Iodine%E2%80%93starch_test). Since *Phacopsis* fungi are usually restricted to living on hosts belonging to a single phylogenetic [clade](/source/Clade), the identity of the host lichen is another diagnostic [character](/source/Phenotypic_trait) used to identify species.

## Systematics

*Phacopsis* was [circumscribed](/source/Circumscription_(taxonomy)) by French [mycologist](/source/Mycology) [Edmond Tulasne](/source/Edmond_Tulasne) in 1852 with multiple species originally classified in genus *[Abrothallus](/source/Abrothallus)*. Tulasne included three species: *P. clemens*, *P. varia*, and *P. vulpina*.[1] The last-mentioned species was later selected as the [type species](/source/Type_species) by [Frederic Clements](/source/Frederic_Clements) and [Cornelius Shear](/source/Cornelius_Lott_Shear) (1931),[2] and by [John Axel Nannfeldt](/source/John_Axel_Nannfeldt) (1932).[3] Of the other two species originally included in *Phacopsis* by Tulasne, *P. clemens* has since been referred to the genus *[Arthonia](/source/Arthonia)*,[4] while *P. varia* is now known as *[Opegrapha physciaria](/source/Opegrapha_physciaria)*.[5]

In 1988, Dagmar Triebel and Gerhard Rambold proposed that *Phacopsis* should be considered [synonymous](/source/Synonym_(taxonomy)) with *[Nesolechia](/source/Nesolechia_(fungus))* (another genus of lichenicolous fungi in the Parmeliaceae), owing to similarities both in the structure of their apothecia and the characteristics of their hymenia.[6] This proposed synonymy, however, was not accepted by several authors in the years following,[7][8][9] until 1995 when Triebel, Rambold, and [John Elix](/source/John_Alan_Elix) showed that the supposed differences that separated the two genera were not consistent.[10] Following this, Ove Eriksson and [David L. Hawksworth](/source/David_L._Hawksworth) used an expanded generic concept of *Phacopsis* (*[sensu lato](/source/Sensu_lato)*) in the next update of their regular publication on [ascomycete](/source/Ascomycota) systematics, *Systema Ascomycetum*.[11]

[Molecular phylogenetic](/source/Molecular_phylogenetic) analysis published in 2015 demonstrated that the group of species then considered to be members of *Phacopsis* actually comprised three distinct [lineages](/source/Lineage_(evolution)), representing three genera: *Phacopsis*, *Nesolechia*, and the newly circumscribed *[Raesaenenia](/source/Raesaenenia)*.[12] In 2017, Divakar and colleagues used a then-recently developed "temporal phylogenetic" approach in an attempt to make family- and genus-level classification more consistent with evolutionary history. They proposed to synonymize genus *Nesolechia* with *[Punctelia](/source/Punctelia)*, and *Raesaenenia* with *[Protousnea](/source/Protousnea)*, because the lichenicolous genera originated relatively recently and fell under the timeframe threshold for genus level.[13] This proposed synonymy was not accepted in a later critical analysis of the temporal phylogenetic approach for fungal classification.[14] In a 2018 review of taxonomic developments in the family Parmeliaceae, the authors recommended not synonymizing *Nesolechia* and *Phacopsis*, suggesting that the separation between these two genera has not yet been sufficiently established.[15]

The higher-level classification of *Phacopsis* has changed several times in its taxonomic history. It has been placed in the order [Arthoniales](/source/Arthoniales),[16] in the families [Graphidaceae](/source/Graphidaceae),[2] and the [Acarosporaceae](/source/Acarosporaceae).[17] Using [electron microscopy](/source/Electron_microscopy), [Josef Hafellner](/source/Josef_Hafellner) examined the [asci](/source/Ascus) (spore-bearing cells) of the type species *P. vulpina*, showing them to be similar to those commonly found in members of the [Lecanoraceae](/source/Lecanoraceae), and so he considered the genus most appropriately classified in this family.[18] This familial placement, however, was not corroborated in Triebel and colleagues' 1995 review and revision of the genus.[10] Molecular phylogenetic analysis eventually revealed its true [phylogenetic](/source/Phylogenetic) placement in the family [Parmeliaceae](/source/Parmeliaceae). This relationship is an example of adelpho-parasitism, in which the host species is closely related to the parasite;[19] this type of relationship is fairly common in the Lecanorales. The genus *Protousnea* has a [sister taxon](/source/Sister_taxon) relationship with *Phacopsis*. It has been estimated that the lichenicolous *Phacopsis* [diverged evolutionarily](/source/Divergent_evolution) from the lichen-forming *Protousnea* during the [Miocene](/source/Miocene) (23.03 to 5.333 [mya](/source/MYA_(unit))).[12]

## Description

All species are endokapylic, meaning they possess a [thallus](/source/Thallus) in which no morphologically distinct lichenized structure is formed. Their [apothecia](/source/Apothecia) are circular to irregular in shape, and may be dispersed or aggregated. The excipulum (the outer margin of the apothecium) range from colourless to blackish-brown, and often comprises "a few rows of cells which may be difficult to recognize". The hypothecium (a layer of dense hyphal tissue just below the [hymenium](/source/Hymenium)) is colourless to blackish-brown. In some species, the hyphae in the hypothecium have a violet [staining](/source/Staining) reaction with an [iodine–starch test](/source/Iodine%E2%80%93starch_test); this characteristic can be used to distinguish between some species. The hymenium is colourless, pale brownish, or pale [olivaceous](/source/Olivaceous). Asci are more or less club-shaped, and contain eight spores. They are surrounded by a thin outer [amyloid](/source/Amyloid_(mycology)) wall layer and a thicker, non-amyloid inner wall layer; a non-amyloid zone rests above the axial body of the [ascus](/source/Ascus).[10]

The [ascospores](/source/Ascospore) are colourless, lack [septa](/source/Septum), and have smooth walls with occasional thickening at either end. The spores have a range of shapes; depending on the species, the following shapes have been recorded: [ellipsoid](/source/Ellipsoid), ovoid (egg-shaped), fusiform (spindle-shaped), lemon-shaped, falciform ([sickle](/source/Sickle)-shaped), fabiform (bean-shaped), and partly curved. The [pycnidia](/source/Pycnidia) of *Phacopsis* are immersed in the thallus of the host. [Pycnospores](/source/Pycnospore) have a [bacilliform](/source/Bacilliform) shape.[10]

## Distribution

Members of the host family, Parmeliaceae, are found worldwide and the family has centres of distribution in Asia and in the [Southern Hemisphere](/source/Southern_Hemisphere); *Phacopsis* probably occurs wherever the host does. A 1995 survey of *Phacopsis* accepted 13 [taxa](/source/Taxon) (eight species and several [varieties](/source/Variety_(botany))) found on 41 host species on 20 genera. Little is known about several *Phacopsis* species, and some have only been found at their [type locality](/source/Type_(biology)#type_locality).[10] The most southernly distributed member of the genus is *Phacopsis usneae*, known to occur only in southern Chile and Antarctica.[20]

Like many lichenicolous fungi, *Phacopsis* species are usually restricted to living on a host belong to a single phylogenetic [clade](/source/Clade). In 2002, [André Aptroot](/source/Andr%C3%A9_Aptroot) and Triebel suggested a possible close phylogenetic relationship between *Paraparmelia* and *[Xanthoparmelia](/source/Xanthoparmelia)*, since *[Phacopsis australis](/source/Phacopsis_australis)* was noted to grow on representatives from both of those lichen genera.[21] Since then, [molecular phylogenetic](/source/Molecular_phylogenetic) analysis has demonstrated that they are equivalent, and *Paraparmelia* is now placed in synonymy with *Xanthoparmelia*.[22]

## Species

*Phacopsis* species are distinguished from each other by the characteristics of their ascospores, the colour and amyloid reaction of the hypothecium, and the identity of their host.[10] According to [Index Fungorum](/source/Index_Fungorum), 33 taxa have been described under the name *Phacopsis* (28 species and 5 varieties).[23] A recent (2022) estimate places 10 species in genus *Phacopsis*.[24] The following list of *Phacopsis* species indicates its name, [taxonomic authority](/source/Author_citation_(botany)) ([standard abbreviations](/source/List_of_botanists_by_author_abbreviation_(A)) are used) and year of publication, type locality, and host species (or genus).

- *[Phacopsis australis](/source/Phacopsis_australis)* Aptroot & Triebel (2002)[21] – southern Africa; host=*[Xanthoparmelia](/source/Xanthoparmelia)*
- *[Phacopsis cephalodioides](/source/Phacopsis_cephalodioides)* (Nyl.) Triebel & Rambold (1988)[6] – Denmark; host=*[Hypogymnia physodes](/source/Hypogymnia_physodes)*
- *[Phacopsis lethariellae](/source/Phacopsis_lethariellae)* Hafellner & Rambold (1995)[10] – [Canary Islands](/source/Canary_Islands); host=*[Lethariella intricata](/source/Lethariella_intricata)*
- *[Phacopsis oroarcticae](/source/Phacopsis_oroarcticae)* Zhurb. (2010)[25] – [Severnaya Zemlya](/source/Severnaya_Zemlya); host=*[Brodoa oroarctica](/source/Brodoa_oroarctica)*
- *[Phacopsis prolificans](/source/Phacopsis_prolificans)* (Müll.Arg.) Triebel & Rambold (1992)[26] – Japan; host=*[Platismatia interrupta](/source/Platismatia_interrupta)*
- *[Phacopsis rufa](/source/Phacopsis_rufa)* (Müll.Arg.) Triebel (1992)[26] – Australia; hosts=*[Cetrelia](/source/Cetrelia)* and *[Punctelia](/source/Punctelia)*
- *[Phacopsis thallicola](/source/Phacopsis_thallicola)* (A.Massal.) Triebel & Rambold (1988)[6] – New Zealand; hosts=*[Cetrelia sanguinea](/source/Cetrelia_sanguinea)*, *[Flavoparmelia caperata](/source/Flavoparmelia_caperata)*, *[Flavoparmelia praesignis](/source/Flavoparmelia_praesignis)*, *[Parmotrema eurysacum](/source/Parmotrema_eurysacum)*, *[Rimelia cetraria](/source/Rimelia_cetraria)*
- *[Phacopsis usneae](/source/Phacopsis_usneae)* C.W.Dodge (1948)[27] – [Kerguelen Islands](/source/Kerguelen_Islands); host=*[Usnea trachycarpa](/source/Usnea_trachycarpa)*
- *[Phacopsis vulpicidae](/source/Phacopsis_vulpicidae)* Zhurb. & Diederich (2019)[28] – [Renchinlkhümbe](/source/Renchinlkh%C3%BCmbe) (Mongolia); host=*[Vulpicida](/source/Vulpicida)*
- *[Phacopsis vulpina](/source/Phacopsis_vulpina)* Tul. (1852) – France; host=*[Letharia](/source/Letharia)*

In 1995, Triebel and colleagues described *Phacopsis menegazziae* for a [Nepalese](/source/Nepal) fungus similar to *Phacopsis oxyspora*, but with smaller spore dimensions and growing on the host *[Menegazzia terebrata](/source/Menegazzia_terebrata)*.[10] Paul Diederich synonymised *P. menegazziae* with *P. oxyspora* in 2003, based on specimens he found growing on *Menegazzia* with ascospores similar in size to typical *P. oxyspora*.[29] *Phacopsis oxyspora* (Tul.) Triebel & Rambold (1988) is now *[Nesolechia oxyspora](/source/Nesolechia_oxyspora)* (Tul.) A.Massal. (1856), the type species of *Nesolechia*.

Several species formerly placed in *Phacopsis* have since been transferred to other genera. These include:

- *Phacopsis crustulosae* Creveld (1981) now *[Gyrophthorus crustulosae](/source/Gyrophthorus_crustulosae)* (Creveld) Hafellner & Sancho (1990)[30]
- *Phacopsis doerfeltii* Alstrup & P.Scholz (1998) now *[Nesolechia doerfeltii](/source/Nesolechia_doerfeltii)* (Alstrup & P.Scholz) Diederich (2018)[31]
- *Phacopsis ericetorum* (Flot. ex Körb.) Vouaux (1914) now *[Rhymbocarpus ericetorum](/source/Rhymbocarpus_ericetorum)* (Flot. ex Körb.) Etayo (2010)[32]
- *Phacopsis falcispora* Triebel & Rambold (1995) now *[Nesolechia falcispora](/source/Nesolechia_falcispora)* (Triebel & Rambold) Diederich (2018)[33]
- *Phacopsis huuskonenii* Räsänen (1948) now *[Raesaenenia huuskonenii](/source/Raesaenenia_huuskonenii)* (Räsänen) D.Hawksw., C.Boluda & H.Lindgren (2015)[12]
- *Phacopsis lesdainii* Vouaux (1914) now *[Echinodiscus lesdainii](/source/Echinodiscus_lesdainii)* (Vouaux) Etayo & Diederich (2000)[34]
- *Phacopsis macrospora* Uloth (1861) now *[Cercidospora macrospora](/source/Cercidospora_macrospora)* (Uloth) Hafellner & Nav.-Ros. (2004)[35]
- *Phacopsis muelleri* Willey (1892) now *[Calloria muelleri](/source/Calloria_muelleri)* (Willey) Seaver (1951)[36]
- *Phacopsis mulleri* Willey (1892) now *[Skyttella mulleri](/source/Skyttella_mulleri)* (Willey) D.Hawksw. & R.Sant. (1988)[37]

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1. ["Record Details: *Phacopsis ericetorum* (Flot. ex Körb.) Vouaux, Bull. Soc. mycol. Fr. 30(2): 144 (1914)"](http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=191263). [Index Fungorum](/source/Index_Fungorum). Retrieved 30 March 2022.

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1. ["Record Details: *Phacopsis muelleri* Willey, Enum. Lich. New Bedford: 34 (1892)"](http://www.indexfungorum.org/names/NamesRecord.asp?RecordID=191160). [Index Fungorum](/source/Index_Fungorum). Retrieved 30 March 2022.

1. Hawksworth, David L. & Santesson, Rolf (1988). ["*Skyttella*, a new genus for *Phacopsis mulleri* Willey (syn. *Agyrium flavescens* Rehm)"](http://nhm2.uio.no/botanisk/lav/Graphis/2_2/GS_2_33.pdf). *Graphis Scripta*. **2** (2): 33–37.

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