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'''''Hydropunctaria amphibia''''' is a species of [[saxicolous lichen|saxicolous]] (rock-dwelling), [[crustose lichen]] in the family [[Verrucariaceae]].<ref name="CoL"/> One of several [[Marine fungi#Lichens|marine lichens]] in the genus ''[[Hydropunctaria]]'', it is found in Europe from Norway to the [[Mediterranean Basin]], and on both the east and west coasts of North America. The black, more or less thick, crust-like [[thallus]] grows on seashore rocks – both [[siliceous rock]]s and [[limestone]] – in the lower [[supralittoral zone]], an area also known as the splash zone. ''Hydropunctaria amphibia'' sets itself apart from other species in ''Hydropunctaria'' through the distinct shape of the {{lichengloss|perithecium}} (fruiting body) apex, which is either flat-topped or scalloped, in contrast to the typically rounded or immersed apex seen in its relatives.
'''''Hydropunctaria amphibia''''' is a species of [[saxicolous lichen|saxicolous]] (rock-dwelling), [[crustose lichen]] in the family [[Verrucariaceae]].<ref name="CoL"/> One of several [[Marine fungi#Lichens|marine lichens]] in the genus ''[[Hydropunctaria]]'', is widely distributed across Europe, extending from Norway to the [[Mediterranean Basin|Mediterranean]] and the [[Iberian Peninsula|Iberian]] coasts, and has a nearly ubiquitous presence along the [[Costa Brava|Catalan coast]] of Spain. In North America, it is found along the Atlantic Coast from [[Nova Scotia]] to the [[Boston Harbor]] islands, where its presence in low-[[pollution]] areas indicates its potential as a [[bioindicator]] for marine lichen community health, and on the west coast in [[British Columbia]], particularly in the [[Gwaii Haanas]]'s upper [[littoral zone|littoral]] fringe. The black, crust-like [[thallus]] grows on seashore rocks – both [[siliceous rock]]s and [[limestone]] – in the lower [[supralittoral zone]], an area also known as the splash zone. Originally described more than two centuries ago as a species of ''[[Verrucaria]]'', ''Hydropunctaria amphibia'' sets itself apart from other species in ''Hydropunctaria'' through the distinct shape of the {{lichengloss|perithecium}} (fruiting body) apex, which is either flat-topped or scalloped, in contrast to the typically rounded or immersed apex seen in its relatives.


==Taxonomy==
==Taxonomy==
The lichen was originally [[species description|described]] in 1807 by the Spanish botanist [[Simón de Roxas Clemente y Rubio]], who classified it as a species of ''[[Verrucaria]]''. His [[Latin]] description of the organism was brief: {{lang|la|Crusta subtartarea suborbiculari uniformi contigua aequabili picea nitida, tuberculis subcylindricis tandem subpatellulaeformibus}} ({{gloss|The crust is almost tartarean, suborbicular, uniform, contiguous, even, pitch-black, shiny, with tubercles initially subcylindrical then eventually somewhat in the form of a small patella (kneecap)}}).<ref name="Clemente y Rubio 1807"/>{{#tag:ref|Passage translated by [[GPT-4]].|group=note}} In 2011, [[Claude Roux]] transferred the [[taxon]] to the ''[[Hydropunctaria]]'',<ref name="Roux et al. 2011"/> a segregate genus of ''Verrucaria'' that had been [[circumscription (taxonomy)|circumscribed]] in 2009.<ref name="Gueidan et al. 2009"/> [[Alan Orange]], apparently unaware of this transfer, proposed a similar transfer himself the following year.<ref name="Orange 2012"/>
The lichen was originally [[species description|described]] in 1807 by the Spanish botanist [[Simón de Roxas Clemente y Rubio]], who classified it as a species of ''[[Verrucaria]]''. His [[Latin]] description of the organism was brief: {{lang|la|Crusta subtartarea suborbiculari uniformi contigua aequabili picea nitida, tuberculis subcylindricis tandem subpatellulaeformibus}} ({{gloss|The crust is almost tartarean, suborbicular, uniform, contiguous, even, pitch-black, shiny, with tubercles initially subcylindrical then eventually somewhat in the form of a small patella (kneecap)}}).<ref name="Clemente y Rubio 1807"/>{{#tag:ref|Passage translated by [[GPT-4]].|group=note}} In 2011, [[Claude Roux]] reclassified the [[taxon]] to ''[[Hydropunctaria]]'',<ref name="Roux et al. 2011"/> a genus previously branched out from ''Verrucaria'' in 2009 to form its own distinct group.<ref name="Gueidan et al. 2009"/> [[Alan Orange]], apparently unaware of this transfer, proposed a similar transfer himself the following year.<ref name="Orange 2012"/>


==Description==
==Description==
''Hydropunctaria amphibia'' is characterised by a [[prothallus#In lichens|prothallus]] that is typically inconspicuous, though occasionally it may appear whitish and is very locally developed. The [[thallus]], which is on the surface of the rock,<ref name="Orange et al. 2023"/> presents a translucent green to amber colouration when wet, and black when dry.<ref name="Taylor 1982"/> It has an abundantly cracked surface that often separates into discrete {{lichengloss|areoles}} (small, distinct patches), making the surface appear uneven with numerous blackish ridges. The actively growing edge of the thallus is clearly defined, with ridges measuring between 40 and 400&nbsp;[[micrometre|μm]] in length and 40 to 50&nbsp;μm in width, generally aligned perpendicular to the margin. Within the interior of the thallus, these ridges are shorter, often branched, and do not follow a uniform direction.<ref name="Orange et al. 2023"/>
The [[crustose lichen]] ''Hydropunctaria amphibia'' is characterised by a [[prothallus#In lichens|prothallus]] that is typically inconspicuous, but in some specific, small areas, it can become visible as a whitish layer. The [[thallus]], which is on the surface of the rock,<ref name="Orange et al. 2023"/> presents a translucent green to amber colouration when wet, and black when dry.<ref name="Taylor 1982"/> It has an abundantly cracked surface that often separates into discrete {{lichengloss|areoles}} (small, distinct patches), making the surface appear uneven with numerous blackish ridges. The actively growing edge of the thallus is clearly defined, with ridges measuring between 40 and 400&nbsp;[[micrometre]]s (μm) in length and 40 to 50&nbsp;μm in width, generally aligned perpendicular to the margin. Within the interior of the thallus, these ridges are shorter, often branched, and do not follow a uniform direction.<ref name="Orange et al. 2023"/>
[[File:Hydropunctaria amphibia.jpg|thumb|''Hydropunctaria amphibia'' growing on aerohaline (sea-sprayed) rocks.]]
[[File:Hydropunctaria amphibia.jpg|thumb|''Hydropunctaria amphibia'' growing on aerohaline (sea-sprayed) rocks.]]
A {{lichengloss|pseudocortex}}, a layer beneath the surface of the thallus, contains brown pigment, contributing to the lichen's dark appearance. {{lichengloss|Perithecia}}, the reproductive structures of the lichen, project from the thallus surface, typically measuring 220 to 360&nbsp;μm in diameter, often with a distinct flat top and sides that may be uneven or have protrusions (bosses); the [[ostiole]], or opening, of the perithecia may be visible as a small pit.<ref name="Orange et al. 2023"/>
The {{lichengloss|pseudocortex}}, a layer beneath the surface of the thallus, contains brown pigment, contributing to the lichen's dark appearance. {{lichengloss|Perithecia}}, the reproductive structures of the lichen, project from the thallus surface. They typically measure 220 to 360&nbsp;μm in diameter, and often have a distinct flat top. The sides may be uneven or have protrusions (bosses). The [[ostiole]], or opening, of the perithecia may be visible as a small pit.<ref name="Orange et al. 2023"/>


The {{lichengloss|involucrellum}}, another structural component of the perithecia, is well-developed, providing additional protection to the spore-producing parts. The {{lichengloss|ascospores}}, which are the reproductive cells produced within the perithecia, are narrowly cylindrical to [[ellipsoid]]al in shape, measuring between 13 and 19.5&nbsp;μm in length and 5 to 7.5&nbsp;μm in width, with a length to width ratio ranging from 2.1 to 3.5, indicating their elongated form.<ref name="Orange et al. 2023"/>
The {{lichengloss|involucrellum}}, another structural component of the perithecia, is well-developed, providing additional protection to the spore-producing parts. The {{lichengloss|ascospores}}, which are the reproductive cells produced within the perithecia, are narrowly cylindrical to [[ellipsoid]]al in shape, measuring between 13 and 19.5&nbsp;μm in length and 5 to 7.5&nbsp;μm in width, with a length to width ratio ranging from 2.1 to 3.5, indicating their elongated form.<ref name="Orange et al. 2023"/>
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==Similar species==
==Similar species==


''Hydropunctaria amphibia'' can be distinguished from other species within the genus ''Hydropunctaria'' by several characteristics. One of the key differences lies in the structure of the perithecium, which in ''Hydropunctaria amphibia'' is either flat-topped or {{lichengloss|crenulate}} (having a scalloped or notched edge), as opposed to the more commonly rounded or immersed apex found in other ''Hydropunctaria'' species. Furthermore, ''Hydropunctaria amphibia'' has a more extensive development of densely pigmented tissue throughout its thallus. This pigmentation manifests as elongated bars, up to 60&nbsp;μm wide, that reach the surface of the thallus. These pigmented areas are particularly noticeable near the thallus margin, where they align perpendicularly to the margin and parallel to the elongated cracks that characterise the species.<ref name="Orange et al. 2023"/>
''Hydropunctaria amphibia'' can be distinguished from other species within the genus ''Hydropunctaria'' by several characteristics. One of the key differences is the structure of the perithecium, which in ''Hydropunctaria amphibia'' is either flat-topped or {{lichengloss|crenulate}} (having a scalloped or notched edge), as opposed to the more commonly rounded or immersed apex found in other ''Hydropunctaria'' species. Furthermore, ''Hydropunctaria amphibia'' has a more extensive development of densely pigmented tissue throughout its thallus. This pigmentation manifests as elongated bars, up to 60&nbsp;μm wide, that reach the surface of the thallus. These pigmented areas are particularly noticeable near the thallus margin, where they align perpendicularly to the margin and parallel to the elongated cracks that characterise the species.<ref name="Orange et al. 2023"/>


One study suggested that a feasible way to distinguish between ''H.&nbsp;amphibia'' and ''H.&nbsp;maura'' was the thickness of the thallus edge and the width of the ascospores.<ref name="Renobales Scheifler & Noya 1991"/> A later publication, however, found that these {{lichengloss|characters}} overlapped in the studied population, and that it was more reliable to use the shape of the perithecia, the extent of thallus cover over the perithecia, and the shape of the {{lichengloss|carbonaceous}} thickenings.<ref name="Paz-Bermúdez et al. 2005"/> Another source suggests that when wet, ''H.&nbsp;amphibia'' becomes more light coloured and translucent compared to ''H.&nbsp;maura'', and so its ridges become more distinctly contrasted.<ref name="Taylor 1982"/>
One study suggested that a feasible way to distinguish between ''H.&nbsp;amphibia'' and ''H.&nbsp;maura'' was the thickness of the thallus edge and the width of the ascospores.<ref name="Renobales Scheifler & Noya 1991"/> A later publication, however, found that these {{lichengloss|characters}} overlapped in the studied population, and that it was more reliable to use the shape of the perithecia, the extent of thallus cover over the perithecia, and the shape of the {{lichengloss|carbonaceous}} thickenings.<ref name="Paz-Bermúdez et al. 2005"/> Another source suggests that when wet, ''H.&nbsp;amphibia'' becomes more light coloured and translucent compared to ''H.&nbsp;maura'', and so its ridges become more distinctly contrasted.<ref name="Taylor 1982"/>
Line 35: Line 35:
==Habitat and distribution==
==Habitat and distribution==


''Hydropunctaria amphibia'' is one of several marine lichens in the genus ''Hydropunctaria''.<ref name="Jones et al. 2015"/> It grows on seashore rocks, both [[siliceous rock|siliceous]] and [[limestone]].<ref name="Orange et al. 2023"/> Rarely, it has been recorded growing on [[concrete]].<ref name="Gilbert 2001"/> It usually grows in the [[supralittoral zone]], the area above the spring [[high tide]] line that is regularly, but infrequently splashed. It typically gets exposure to direct sunlight in these locations.<ref name="Gasulla et al. 2019"/> It is one of few species able to survive under these conditions, due to its extreme resistance to desiccation. Other organisms commonly found in this habitat include the [[barnacle]] species ''[[Microeuraphia depressa]]'', and the [[sea snail]] species ''[[Melarhaphe neritoides]]'', ''[[Echinolittorina punctata]]''.<ref name="Chappuis et al. 2014"/> ''Hydropunctaria amphibia'' grows in the belt underneath its close relative ''[[Hydropunctaria maura|H.&nbsp;maura]]''.<ref name="Paz-Bermúdez et al. 2005"/>
''Hydropunctaria amphibia'' is one of several marine lichens in the genus ''Hydropunctaria''.<ref name="Jones et al. 2015"/> It grows on seashore rocks, both [[siliceous rock|siliceous]] and [[limestone]].<ref name="Orange et al. 2023"/> Rarely, it has been recorded growing on [[concrete]].<ref name="Gilbert 2001"/> It usually grows in the [[supralittoral zone]], the area above the spring [[high tide]] line that is regularly but infrequently splashed. It typically gets exposure to direct sunlight in these locations.<ref name="Gasulla et al. 2019"/> It is one of few species able to survive under these conditions, due to its extreme resistance to desiccation. Other organisms commonly found in this habitat include the [[barnacle]] species ''[[Microeuraphia depressa]]'', and the [[sea snail]] species ''[[Melarhaphe neritoides]]'' and ''[[Echinolittorina punctata]]''.<ref name="Chappuis et al. 2014"/> ''Hydropunctaria amphibia'' grows in the belt underneath its close relative ''[[Hydropunctaria maura|H.&nbsp;maura]]''.<ref name="Paz-Bermúdez et al. 2005"/>


The lichen is widely distributed in Europe, from Norway south to the [[Mediterranean]], and also on the [[Iberian Peninsula|Iberian]] coasts.<ref name="Paz-Bermúdez et al. 2005"/> Detailed [[cartography]] of the littoral habitats found along the [[Catalan Coastal Range|Catalan coast]] of Spain show its almost ubiquitous presence there.<ref name="Mariani et al. 2014"/> In North America, ''Hydropunctaria amphibia'' has been documented from the [[East Coast of the United States|Atlantic Coast]], including the [[littoral zone]] of [[Nova Scotia]]'s [[Bay of Fundy]] coast,<ref name="Vail & Walker 2021"/> the coast of [[Newfoundland and Labrador]],<ref name="Brodo 1995"/> and the [[Boston Harbor Islands National Recreation Area]]. In the latter location, it only grows in low-[[pollution]] areas, suggesting low pollution tolerance, and potential for use as a [[bioindicator]] of marine lichen community health.<ref name="Nokes et al. 2022"/> On the west coast, it is known to occur in [[British Columbia]]. In the [[Gwaii Hanaas]], it grows in the upper littoral fringe–defined as {{cvt|0.3|–|0.8|m}} above the ''[[Fucus]]''-[[barnacle]] zone–most commonly on limestone.<ref name="Brodo and Sloan 2004"/>
The lichen is widely distributed in Europe, from Norway south to the [[Mediterranean]] and also on the [[Iberian Peninsula|Iberian]] coasts.<ref name="Paz-Bermúdez et al. 2005"/> Detailed [[cartography]] of the littoral habitats found along the [[Catalan Coastal Range|Catalan coast]] of Spain show its almost ubiquitous presence there.<ref name="Mariani et al. 2014"/> In North America, ''Hydropunctaria amphibia'' has been documented from the [[East Coast of the United States|Atlantic Coast]], including the [[littoral zone]] of [[Nova Scotia]]'s [[Bay of Fundy]] coast,<ref name="Vail & Walker 2021"/> the coast of [[Newfoundland and Labrador]],<ref name="Brodo 1995"/> and the [[Boston Harbor Islands National Recreation Area]]. In the latter location, it only grows in low-[[pollution]] areas, suggesting low pollution tolerance, and potential for use as a [[bioindicator]] of marine lichen community health.<ref name="Nokes et al. 2022"/> On the west coast, it is known to occur in [[British Columbia]]. In the [[Gwaii Hanaas]], it grows in the upper littoral fringe (defined as {{cvt|0.3|–|0.8|m}} above the ''[[Fucus]]'' [[barnacle]] zone), most commonly on limestone.<ref name="Brodo and Sloan 2004"/>


==Notes==
==Notes==

Revision as of 17:37, 4 March 2024

Hydropunctaria amphibia
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
Class: Eurotiomycetes
Order: Verrucariales
Family: Verrucariaceae
Genus: Hydropunctaria
Species:
H. amphibia
Binomial name
Hydropunctaria amphibia
(Clemente) Cl.Roux (2011)
Synonyms[1]
  • Verrucaria amphibia Clemente (1807)

Hydropunctaria amphibia is a species of saxicolous (rock-dwelling), crustose lichen in the family Verrucariaceae.[2] One of several marine lichens in the genus Hydropunctaria, is widely distributed across Europe, extending from Norway to the Mediterranean and the Iberian coasts, and has a nearly ubiquitous presence along the Catalan coast of Spain. In North America, it is found along the Atlantic Coast from Nova Scotia to the Boston Harbor islands, where its presence in low-pollution areas indicates its potential as a bioindicator for marine lichen community health, and on the west coast in British Columbia, particularly in the Gwaii Haanas's upper littoral fringe. The black, crust-like thallus grows on seashore rocks – both siliceous rocks and limestone – in the lower supralittoral zone, an area also known as the splash zone. Originally described more than two centuries ago as a species of Verrucaria, Hydropunctaria amphibia sets itself apart from other species in Hydropunctaria through the distinct shape of the perithecium (fruiting body) apex, which is either flat-topped or scalloped, in contrast to the typically rounded or immersed apex seen in its relatives.

Taxonomy

The lichen was originally described in 1807 by the Spanish botanist Simón de Roxas Clemente y Rubio, who classified it as a species of Verrucaria. His Latin description of the organism was brief: Crusta subtartarea suborbiculari uniformi contigua aequabili picea nitida, tuberculis subcylindricis tandem subpatellulaeformibus ('The crust is almost tartarean, suborbicular, uniform, contiguous, even, pitch-black, shiny, with tubercles initially subcylindrical then eventually somewhat in the form of a small patella (kneecap)').[3][note 1] In 2011, Claude Roux reclassified the taxon to Hydropunctaria,[4] a genus previously branched out from Verrucaria in 2009 to form its own distinct group.[5] Alan Orange, apparently unaware of this transfer, proposed a similar transfer himself the following year.[6]

Description

The crustose lichen Hydropunctaria amphibia is characterised by a prothallus that is typically inconspicuous, but in some specific, small areas, it can become visible as a whitish layer. The thallus, which is on the surface of the rock,[7] presents a translucent green to amber colouration when wet, and black when dry.[8] It has an abundantly cracked surface that often separates into discrete areoles (small, distinct patches), making the surface appear uneven with numerous blackish ridges. The actively growing edge of the thallus is clearly defined, with ridges measuring between 40 and 400 micrometres (μm) in length and 40 to 50 μm in width, generally aligned perpendicular to the margin. Within the interior of the thallus, these ridges are shorter, often branched, and do not follow a uniform direction.[7]

Hydropunctaria amphibia growing on aerohaline (sea-sprayed) rocks.

The pseudocortex, a layer beneath the surface of the thallus, contains brown pigment, contributing to the lichen's dark appearance. Perithecia, the reproductive structures of the lichen, project from the thallus surface. They typically measure 220 to 360 μm in diameter, and often have a distinct flat top. The sides may be uneven or have protrusions (bosses). The ostiole, or opening, of the perithecia may be visible as a small pit.[7]

The involucrellum, another structural component of the perithecia, is well-developed, providing additional protection to the spore-producing parts. The ascospores, which are the reproductive cells produced within the perithecia, are narrowly cylindrical to ellipsoidal in shape, measuring between 13 and 19.5 μm in length and 5 to 7.5 μm in width, with a length to width ratio ranging from 2.1 to 3.5, indicating their elongated form.[7]

The photobiont of H. amphibia specimens collected in the Iberian Peninsula was Halofilum ramosum (a species of green alga in the order Ulvales). In an ecological study conducted in Anglesey, off the north-west coast of Wales, H. amphibia was shown to have a distribution similar to the brown algal species Pelvetia canaliculata, and species of the red algal genus Hildenbrandia.[9]

Similar species

Hydropunctaria amphibia can be distinguished from other species within the genus Hydropunctaria by several characteristics. One of the key differences is the structure of the perithecium, which in Hydropunctaria amphibia is either flat-topped or crenulate (having a scalloped or notched edge), as opposed to the more commonly rounded or immersed apex found in other Hydropunctaria species. Furthermore, Hydropunctaria amphibia has a more extensive development of densely pigmented tissue throughout its thallus. This pigmentation manifests as elongated bars, up to 60 μm wide, that reach the surface of the thallus. These pigmented areas are particularly noticeable near the thallus margin, where they align perpendicularly to the margin and parallel to the elongated cracks that characterise the species.[7]

One study suggested that a feasible way to distinguish between H. amphibia and H. maura was the thickness of the thallus edge and the width of the ascospores.[10] A later publication, however, found that these characters overlapped in the studied population, and that it was more reliable to use the shape of the perithecia, the extent of thallus cover over the perithecia, and the shape of the carbonaceous thickenings.[11] Another source suggests that when wet, H. amphibia becomes more light coloured and translucent compared to H. maura, and so its ridges become more distinctly contrasted.[8]

In comparison, Wahlenbergiella striatula, another marine lichen species that may present ridges on its thallus, differs primarily in the size of its ascospores, which are smaller than those of Hydropunctaria amphibia.[7] Also, its thallus has a dot matrix that imparts texture to the surface.[12]

Habitat and distribution

Hydropunctaria amphibia is one of several marine lichens in the genus Hydropunctaria.[13] It grows on seashore rocks, both siliceous and limestone.[7] Rarely, it has been recorded growing on concrete.[14] It usually grows in the supralittoral zone, the area above the spring high tide line that is regularly but infrequently splashed. It typically gets exposure to direct sunlight in these locations.[15] It is one of few species able to survive under these conditions, due to its extreme resistance to desiccation. Other organisms commonly found in this habitat include the barnacle species Microeuraphia depressa, and the sea snail species Melarhaphe neritoides and Echinolittorina punctata.[16] Hydropunctaria amphibia grows in the belt underneath its close relative H. maura.[11]

The lichen is widely distributed in Europe, from Norway south to the Mediterranean and also on the Iberian coasts.[11] Detailed cartography of the littoral habitats found along the Catalan coast of Spain show its almost ubiquitous presence there.[17] In North America, Hydropunctaria amphibia has been documented from the Atlantic Coast, including the littoral zone of Nova Scotia's Bay of Fundy coast,[18] the coast of Newfoundland and Labrador,[19] and the Boston Harbor Islands National Recreation Area. In the latter location, it only grows in low-pollution areas, suggesting low pollution tolerance, and potential for use as a bioindicator of marine lichen community health.[20] On the west coast, it is known to occur in British Columbia. In the Gwaii Hanaas, it grows in the upper littoral fringe (defined as 0.3–0.8 m (1 ft 0 in – 2 ft 7 in) above the Fucus barnacle zone), most commonly on limestone.[21]

Notes

  1. ^ Passage translated by GPT-4.

References

  1. ^ "Synonymy. Current Name: Hydropunctaria amphibia (Clemente) Cl. Roux, in Roux, Masson, Bricaud, Coste & Poumarat, Bull. Soc. linn. Provence, num. spéc. 14: 108 (2011)". Species Fungorum. Retrieved 7 February 2024.
  2. ^ "Hydropunctaria amphibia (Clemente) Cl. Roux". Catalogue of Life. Species 2000: Leiden, the Netherlands. Retrieved 7 February 2024.
  3. ^ Clemente y Rubio, S.R. (1807). Ensayo sobre las variedades de la vid comun que vegetan en Andalucia [Essay on the varieties of the common grapevine that grow in Andalusia] (PDF) (in Latin). Madrid: Imprenta de Villalpando. p. 299.
  4. ^ Roux, C.; Masson, D.; Bricaud, O.; Coste, C.; Poumarat, S. (2011). "Flore et végétation des lichens et champignons lichénicoles de quatre réserves naturelles des Pyrénées-Orientales (France)" (PDF). Bulletin de la Société Linnéenne de Provence (in French). 14 (spécial): 3–151 [108].
  5. ^ Gueidan, Cécile; Savić, Sanja; Thüs, Holger; Roux, Claude; Keller, Christine; Tibell, Leif; Prieto, Maria; Heiðmarsson, Starri; Breuss, Othmar; Orange, Alan; Fröberg, Lars; Wynns, Anja Amtoft; Navarro-Rosinés, Pere; Krzewicka, Beata; Pykälä, Juha; Grube, Martin; Lutzoni, François (2009). "Generic classification of the Verrucariaceae (Ascomycota) based on molecular and morphological evidence: recent progress and remaining challenges". Taxon. 58 (1): 184–208. doi:10.1002/tax.581019.
  6. ^ Orange, Alan (2012). "Semi-cryptic marine species of Hydropunctaria (Verrucariaceae, lichenized Ascomycota) from north-west Europe". The Lichenologist. 44 (3): 299–320. doi:10.1017/S0024282911000867.
  7. ^ a b c d e f g Orange, A.; Cannon, P.; Prieto, M.; Coppins, B.; Sanderson, N.; Simkin, J. (2023). Verrucariales: Verrucariaceae, including the genera Agonimia, Atla, Bagliettoa, Catapyrenium, Dermatocarpon, Endocarpon, Henrica, Heteroplacidium, Hydropunctaria, Involucropyrenium, Merismatium, Nesothele, Normandina, Parabagliettoa, Placidopsis, Placidium, Placopyrenium, Polyblastia, Psoroglaena, Sporodictyon, Staurothele, Thelidium, Trimmatothele, Verrucaria, Verrucula, Verruculopsis and Wahlenbergiella (PDF). Revisions of British and Irish Lichens. Vol. 31. p. 35.
  8. ^ a b Taylor, Ronald M. (1982). Marine Flora and Fauna of the Northeastern United States. Lichens (Ascomycetes) of the Intertidal Region (Report). NOAA Technical Report NMFS Circular. Vol. 446. U.S. Department of Commerce. National Oceanic and Atmospheric Administration. National Marine Fisheries Service. p. 17.
  9. ^ Fletcher, A. (1973). "The Ecology of Marine (Littoral) Lichens on some Rocky shores of Anglesey". The Lichenologist. 5 (5–6): 368–400. doi:10.1017/s0024282973000459.
  10. ^ Renobales Scheifler, Gustavo; Noya, Rosario (1991). "Estudio morfológico comparado de Verrucaria maura y V. amphibia en la Costa Vasca" [Comparative morphological study of Verrucaria maura and V. amphibia on the Basque Coast]. Acta Botanica Malacitana. 16: 149–156.
  11. ^ a b c Paz-Bermúdez, Graciela; Eugenia Lopez de Silanes, Maria; Carballal, Regina (2005). "The lichen genera Collemopsidium Nyl. and Verrucaria Schrader from the Galician seashores (NW Spain)". Nova Hedwigia. 80 (1–2): 73–88.
  12. ^ So, Jae Eun; Halda, Josef P.; Hong, Soon Gyu; Hur, Jae-Seoun; Kim, Ji Hee (2023). "The Revision of Lichen Flora Around Maxwell Bay, King George Island, Maritime Antarctic". Journal of Microbiology. 61 (2): 159–173. doi:10.1007/s12275-023-00015-x. PMC 9998320. PMID 36847971.
  13. ^ Jones, E.B. Gareth; Suetrong, Satinee; Sakayaroj, Jariya; Bahkali, Ali H.; Abdel-Wahab, Mohamed A.; Boekhout, Teun; Pang, Ka-Lai (2015). "Classification of marine Ascomycota, Basidiomycota, Blastocladiomycota and Chytridiomycota". Fungal Diversity. 73 (1): 1–72 [51]. doi:10.1007/s13225-015-0339-4.
  14. ^ Gilbert, Oliver (2001). "The Lichen flora of coastal saline lagoons in England". The Lichenologist. 33 (5): 409–417. doi:10.1006/lich.2001.0344.
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  16. ^ Chappuis, Eglantine; Terradas, Marc; Cefalì, Maria Elena; Mariani, Simone; Ballesteros, Enric (2014). "Vertical zonation is the main distribution pattern of littoral assemblages on rocky shores at a regional scale". Estuarine, Coastal and Shelf Science. 147: 113–122. doi:10.1016/j.ecss.2014.05.031. hdl:10045/48363.
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