Squamarina cartilaginea (With.) P. James f. cartilaginea

in Hawksworth & al., Lichenologist 12: 107, 1980. Basionym: Lichen cartilagineus With. - Bot. Arrang. Brit. Plants, 2: 708, 1776.
Synonyms: Lecanora benacensis (A. Massal.) Jatta; Lecanora cartilaginea (With.) A.L. Sm.; Lecanora crassa (Huds.) Ach.; Lecanora crassa var. caespitosa (Vill.) Rabenh.; Parmelia dufourei Fr.; Placodium crassum (Huds.) Link; Psoroma benacense A. Massal.; Psoroma crassum (Huds.) Gray; Squamaria crassa (Huds.) DC.; Squamarina cartilaginea (With.) P. James var. cartilaginea; Squamarina cartilaginea f. cartilaginea (With.) P. James; Squamarina cartilaginea f. iberica (Mattick) Clauzade & Cl. Roux; Squamarina crassa (Huds.) Poelt
Distribution: N - VG (Martellos & al. 2014), Frl (Tretiach 1996, Tretiach & Molaro 2007, Martellos & al. 2014), Ven (Nascimbene & Caniglia 1997, 2003c, Caniglia & al. 1999, Lazzarin 2000b, Nascimbene 2005c, 2008c, Thor & Nascimbene 2007, Nascimbene & Marini 2007, Martellos & al. 2014), TAA (Nascimbene 2001b, 2008b, Nascimbene & al. 2006, 2022), Lomb (Dalle Vedove & al. 2004, Gheza 2019), Piem (Clerc & al. 1999, Isocrono & al. 2003, Martellos & al. 2014), VA (Borlandelli & al. 1996, Piervittori & Isocrono 1997, 1999), Emil (Scarpa 1993, Nimis & al. 1996, Martellos & al. 2014, Fariselli & al. 2020), Lig (Valcuvia & al. 2000, Martellos & al. 2014, Giordani & al. 2016). C - Tosc (Benesperi 2000a, 2006, Loppi & al. 1998, 2004b, Benesperi & Lastrucci 2007, Tretiach & al. 2008, Lastrucci & al. 2009, Pasquinelli & Puccini 2010, Pasquinelli & al. 2013, Martellos & al. 2014), Marc (Nimis & Tretiach 1999, Martellos & al. 2014), Umb (Nimis & Tretiach 1999, Panfili 2000b, 2007, Ravera & al. 2006, Brackel 2015), Laz (Nimis & Tretiach 2004, Martellos & al. 2014, Brackel 2015, Di Pietro & al. 2022), Abr (Nimis & Tretiach 1999, Martellos & al. 2014, Caporale & al. 2016, Gheza & al. 2021), Mol (Garofalo & al. 1999, Nimis & Tretiach 1999, Caporale & al. 2008, Martellos & al. 2014, Genovesi & Ravera 2014), Sar (Nöske 2000, Rizzi & al. 2011, Martellos & al. 2014). S - Camp (Garofalo & al. 1999, 2010, Aprile & al. 2002, 2003, 2003b, Nimis & Tretiach 2004, Martellos & al. 2014), Pugl (Garofalo & al. 1999, Nimis & Tretiach 1999, Durini & Medagli 2002, Brackel 2011, Martellos & al. 2014, Gianfreda & Matino 2020), Bas (Nimis & Tretiach 1999, Potenza 2006, Potenza & al. 2010), Cal (Puntillo 1996, Lich. Graec. 297: Obermayer 2006, Martellos & al. 2014), Si (Nimis & al. 1994, 1996b, Ottonello & al. 1994, 2011, Ottonello 1996, Ottonello & Romano 1997, Grillo 1998, Grillo & al. 2002, 2009, Grillo & Caniglia 2004, Merlo 2004b, Caniglia & Grillo 2005, 2006, Brackel 2008b, Gianguzzi & al. 2009, Cataldo & Minissale 2013, Martellos & al. 2014, Ravera & al. 2023b).
Description: Thallus squamulose, 3-15 cm wide, 0.3-0.8 mm thick, the squamules imbricate, greenish when fresh, turning brownish in the herbarium, epruinose except in the young stages, indistinctly lobed except in young stages, the margins usually white, rarely blackening, the lower surface dark grey to black at least in central parts (pale in young squamules and in peripheral parts), attached by concolour rhizinoid hyphal strands. Upper cortex 20-50 µm thick, with an epinecral layer which is hyaline and without crystals in the upper 4-10 μm, and inspersed with crystals in the lower 8-15 µm; true cortex 15-35 µm thick, plectenchymatous, with elongated cells; medulla white (brownish in lower part), up to 0.6 mm thick, filled with crystals. Apothecia frequent, 0.6-3(-4) mm across, finally constricted at base, at first zeorine, then almost biatorine, with a yellowish white to medium brown, at first concave then flat to convex disc, and a 0.1-0.25 mm thick, pale brown to whitish, finally excluded thalline margin, the proper margin thin and poorly evident, level with disc. Thalline exciple 230-370 μm wide, corticate; proper exciple 200-260 μm wide laterally, filled with crystals, of branched-anastomosing hyphae arranged in a fan-way; epithecium yellowish brown, granular, 6-10 µm high; hymenium colourless, 60-80 µm high, hemiamyloid; paraphyses coherent, mostly simple, 1-1.5 μm thick at mid-level, the apical cells 2-4 μm wide; hypothecium colourless, 35-100 μm high, subprosoplectenchymatous, with large crystals soluble in K and insoluble in N, I- (the subhymenium is thin and devoid of crystals). Asci 8-spored, elongate-clavate, with a thin, outer amyloid layer and a thickened, amyloid tholus devoid of an ocular chamber, penetrated by an axial tube the sides of which stain I+ deeper blue, approaching the Porpidia-type. Ascospores 1-celled, hyaline, ellipsoid to subfusiform, (8-)11-15(-17) x (3.5-)4-5.5(-7) µm. Photobiont chlorococcoid. Spot tests: thallus K-, C-, KC- or KC+ faintly yellow, P-; medulla P+ yellow. Chemistry: cortex with usnic and isousnic acids; medulla with psoromic and 2ʹ-O-demethylpsoromic acids.
Note: a mainly southern, chemically variable species found on calcareous rocks, more rarely on soil, the most common species of the genus in Italy. Older records could refer to S. paradoxa (see note on that species). The f. iberica (Mattick) Clauzade & Cl. Roux, with a P+ red medulla, should be looked for in Italy. Strictly terricolouos specimens with a thicker cortex and somewhat larger spores have been recently described as Squamarina terricola Cl. Roux & Poumarat (Roux & Poumarat 2024): I prefer to await the results of an announced molecular analysis before accepting that species as distinct from S. cartilaginea.
Growth form: Squamulose
Substrata: rocks, soil, terricolous mosses, and plant debris
Photobiont: green algae other than Trentepohlia
Reproductive strategy: mainly sexual

Commonnes-rarity: (info)

Alpine belt: absent
Subalpine belt: extremely rare
Oromediterranean belt: very rare
Montane belt: rather common
Submediterranean belt: extremely common
Padanian area: very rare
Humid submediterranean belt: extremely common
Humid mediterranean belt: common
Dry mediterranean belt: common

pH of the substrata:

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Solar irradiation:

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Aridity:

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Eutrophication:

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Poleotolerance:

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Altitudinal distribution:

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Predictive model
Herbarium samples

Paolo Fontanesi; Owner: Paolo Fontanesi
Italy, Emilia-Romagna, Reggio Emilia, Sologno, nell'Appennino Reggiano.
Herbarium: Forum Natura Mediterraneo
2010



Andrea Moro; Owner: Department of Life Sciences, University of Trieste
italy, Friuli Venezia Giulia, Trieste, Trieste Karst near Borgo Grotta Gigante
16/02/2017



Andrea Moro; Owner: Department of Life Sciences, University of Trieste
italy, Friuli Venezia Giulia, Trieste, Trieste Karst near Borgo Grotta Gigante
16/02/2017



Andrea Moro; Owner: Department of Life Sciences, University of Trieste
italy, Friuli Venezia Giulia, Trieste, Trieste Karst near Borgo Grotta Gigante
16/02/2017



Andrea Moro; Owner: Department of Life Sciences, University of Trieste
italy, Friuli Venezia Giulia, Trieste, Trieste Karst near Borgo Grotta Gigante
16/02/2017


S. Martellos; Owner: Department of Life Sciences, University of Trieste
Herbarium: TSB (33367)
2003/03/04
Reaction P+ yellow


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Courtesy Danièle et Olivier Gonnet - Source: hhttps://www.afl-lichenologie.fr/Photos_AFL/Photos_AFL_S/Squamarina_cartilaginea.htm
France, 15/09/2014 - Mont-Cenis, chemin Chapelle St Barthélémy, alt. 2025 m - Savoie


Bernard Bougginier - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, Keremma


Bernard Bougginier - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Bernard Bougginier - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Bernard Bougginier - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, Treflez


Micghel David - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Micghel David - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Micghel David - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Micghel David - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, La Palue


Micghel David - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, Camaret Pen-Hat


Jacques Haine - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, Crozon


Jacques Haine - Source: http://www.lichensmaritimes.org/index.php?task=fiche&lichen=176&lang=en
France, Crozon


Marta González Garcia - Centro de Estudios Micologicos Asturianos
Spain


Marta González Garcia - Centro de Estudios Micologicos Asturianos
Spain


Marta González Garcia - Centro de Estudios Micologicos Asturianos
Spain