This is a key to fertile specimens of all non-fruticose species of Ramalinaceae known to occur in Italy (Nimis 2016), plus several taxa which are known from neighbouring countries (e.g. Nimis & al. 2018) and may occur in Italy. The key also includes the following genera which contain species which were previously assigned to the Ramalinaceae, but do not belong to this family: Adelolecia Hertel & Hafellner, Bryobilimbia Fryday, Printzen & S. Ekman, Catinaria Vain., Frutidella Kalb, Japewia Tønsberg, Porpidinia Timdal, and Schadonia Körb, plus Gilbertaria M. Svensson & Fryday (Sphaerophoraceae), for a total of 166 infrageneric taxa. A key to British species was published by Cannon & al. (2021). The present key, after a period of testing, will be eventually integrated into a general e-key to all lichens potentially occurring in Italy (Nimis & Martellos 2021).
The following genera do belong to the Ramalinaceae:
1) Bacidia De Not. - The delimitation of this large genus originally including more than 200 species has been problematic for quite a long time. The molecular phylogeny published by Kistenich & al. (2016) shows that Bacidia should be restricted to the highly supported B. rosella-group. The B. coprodes-group has been revised by Llop & Ekman (2007) and Ekman (2014), the B. rosella-group in Europe by Llop & al. (2007); the species of the Iberian Peninsula were treated by Llop & Hladun (2002) and Llop (2007). North American species were treated by Ekman (1996).
2) Bacidina Vězda - This genus was introduced to include a number of species previously referred to Bacidia, some of which were already earlier put in separate genera like Lichingoldia and Woessia because of their long, curved conidia. Although not recognised as a genus until 1991, these species had been informally treated by various authors as the "Bacidia phacodes-group". In the single-marker phylogeny of Ekman (2001), Bacidina formed an unsupported, monophyletic group. However, some evidence has accumulated that the genus may have to be split into Bacidina s.str. and Woessia (Kistenich & al. 2018). Bacidina was recovered as non-monophyletic also in the consensus trees of Sérusiaux & al. (2012), Miadlikowska & al. (1848), Kistenich & al. (2018b), van den Boom & Alvarado (2019) and van den Boom & Magain (2020), in all cases without enough branch support to draw any robust conclusions. According to Ekman (2023), the currently available sequence data cannot distinguish between a monophyletic and a non-monophyletic Bacidina s.lat. Until now, altogether c. 70 species have been formally named in Bacidina. The recent paper by Ekman (2023) recognizes 28 species as being present in Scandinavia. In Italy the genus is still very poorly studied, and would be well-worthy of a thorough monograph.
3) Bellicidia Kistenich, Timdal, Bendiksby & S. Ekman - This small genus was segregated from Bacidia by Kistenich & al. (2018) on the basis of both morphological and molecular data. It has a solitary position on a strongly supported branch as sister to the rest of the Toninia-clade.
4) Biatora Fr. - In its current circumscription, Biatora, included in the Ramalinaceae, comprises crustose species with a green algal photobiont, biatorine apothecia with an exciple composed of anticlinal-parallel hyphae, weakly branched, anastomosed and strongly conglutinated paraphyses, Biatora-type asci and simple to 3(-7)-septate, colourless ascospores. Until this modern delimitation of the genus, the species now accepted as Biatora were distributed over a heterogeneous group of taxa including e.g. Bacidia, Catillaria, and Lecidea. Since the second half of the XIX century, Biatora was treated as a subgenus of Lecidea until it was reinstalled as a monotypic genus based on B. vernalis. The circumscription of the genus has changed since then and the number of species increased from 17 to 42 (Printzen 2014). Molecular studies, based on a single or two gene loci, supported the monophyly of Biatora and its position within Ramalinaceae; a comprehensive study based on three gene loci also showed that the genus comprises at least six clades, which correspond to different morphological groups of species: the beckhausii-, hertelii-, meiocarpa-, ocelliformis-, rufidula- and vernalis-groups (Printzen 2014). Many species are chionophilous and extremely shade-tolerant. Three main distributional patterns can be observed: (sub-)arctic-alpine species, sub-oceanic-montane species, and species of montane coniferous or mixed coniferous forests (Printzen & Palice 1999). Keys to European species were published by Printzen (1995) and Printzen & Otte (2005).
5) Bibbya J.H. Willis - This recently resurrected genus (see Kistenich & al. 2018) was included in Toninia by Timdal (1992) and corresponds to his species groups 4 and 8 and partly group 5. Morphologically, it is characterized by a reddish brown, K+ red pigment in the epithecium and rim of the exciple.
6) Bilimbia De Not. - This genus, widely used in the XIX century, fell into disuse because of conflict with an earlier use of the name for a genus of higher plants (which, however, proved to be invalid), so that the species were subsumed into the “supergenus” Bacidia by Zahlbruckner. It differs from Bacidia, Biatora and Mycobilimbia by a slightly different tholus structure, the stout paraphyses, and the finely warted perispore, and presently includes c. 6 species. Molecular data suggest that it forms a well-supported group within the Ramalinaceae (see Reese Naesborg & al. 2007, and Miadlikowska & al. 2014).
7) Cliostomum Fr. - This genus presently includes c. 20 species, but several more species are expected, since several species may occur as sterile, sorediate crusts provisionally assigned to other genera (see Tønsberg 1992, Ekman 1997). The genus seems to be the sister group of Ramalina in the Ramalinaceae (Ekman 2001).
8) Kiliasia Hafellner - This genus, which was included in Toninia by Timdal (1992), includes species from his groups 1, 3, and 7 and is is a sister group to Bibbya and Thalloidima (Kistenich & al. 2018).
9) Lecania A. Massal. - This genus, whose saxicolous species were monographed by Mayrhofer (1988) in Europe, was found to be non-monophyletic by Reese Næsborg & al. (2007), which brought to the exclusion of some species; the closest genetic relatives are genera such as Bilimbia, Mycobilimbia, and Biatora. The phylogeny of the L. cyrtella-group was studied by Reese Næsborg (2008) with the resurrection of some species which were often considered as synonyms of L. cyrtella. In its present circumscription, the genus includes c. 50 species.
10) Megalaria Hafellner - This genus was separated from Catinaria to include the single species M. grossa, but it has subsequently been enlarged by the addition of numerous other morphologically similar taxa, and currently consists of c. 30 species from tropical, temperate and subpolar regions. The genus was originally characterised by asci with an axial body (barrel-shaped in the case of M. grossa but conical in other species), and by spores lacking a distinct perispore, until Ekman & Tønsberg (1996) included further characters, such as the texture of the exciple and the reactions in K and N of the insoluble apothecial pigments. See also Fryday & Lendemer (2010).
11) Mycobilimbia Rehm - This genus has faced several taxonomic complexities, and many species have been transferred to other genera. Three main groups were recognised: the first is now assigned to Bilimbia (formerly Myxobilimbia), which differs in having a warted perispore, the second contains M. tetramera (the type) and relatives, while the third, segregated into Bryobilimbia (not belonging into the Ramalinaceae), comprises the “Lecidea hypnorum”-group with simple ascospores (Fryday & al. 2014). M. berengeriana and M. olivacea still remain unassigned.
12) Phyllopsora Müll. Arg. - This pantropical and subtropical genus includes c. 55 species occurring in humid forests, primarily on bark, but also on decorticated wood, rock and bryophytes (Brako 1991). The genus is similar to Biatora, differing in the pubescent-squamulose thallus and in the slightly different ascus structure.
13) Scutula Tul. - This genus was treated by Triebel & al. (1997) and Wedin & al. (2007) to include a set of lichenicolous species. The association with Ramalinaceae was confirmed by Kistenich & al. (2018) who expanded the genus to include lichen-forming species without any obvious parasitic life-cycle stages.
14) Thalloidima A. Massal. - This recently resurrected genus (see Kistenich & al. 2018) was included in Toninia by Timdal (1992) and corresponds to his species groups 1 (except 4 species now placed in Kiliasia) and 10. Morphologically, it is characterized by the presence of a grey, K+ and N+ violet pigment in the epithecium and exciple (with the exception of T. toninianum). The thallus is mostly flattened-squamulose to bullate, but two species are non-lichenized. Most species are cyanotrophic at least when young.
15) Toninia A. Massal. - This genus was monographed by Timdal (1992), and also after the segregation of Porpidinia (Timdal 2010) it appeared to be highly polyphyletic (Ekman 2001, see also Miadlikowska & al. 2014). The molecular study by Kistenich & al. (2018) split Toninia in the sense of Timdal (1992) into segregates: the two new genera Bellicidia and Parallopsora, and the resurrected genera Bibbya, Kiliasia, Toniniopsis, Sporacestra, and Thalloidima. In its more restricted definition, Toninia corresponds to species groups 2, 6, 9 and partly 5 and 7 of Timdal (1992), with the inclusion of Arthrosporum and some species formerly assigned to Bacidia.
16) Toniniopsis Frey - This recently resurrected genus consists of species previously placed in Bacidia and Toninia, being morphologically similar to the latter, but differing in the generally stronger pigmentation of the exciple (see Kistenich & al. 2018). 17) Waynea Moberg - This genus, described to accommodate a single species from North America, presently comprises 7 species mainly occurring in areas with a Mediterranean climate (one species, however is limited to Siberia). The genus is characterised by a squamulose thallus, a well developed upper eucortex, laminal, biatorine apothecia with an algal layer below the hypothecium, asci of the Bacidia-type, and ellipsoidal to acicular, 0-5-septate ascospores. Important information of species of the genus is in Abassi Maaf & Roux (1985), van den Boom 2010, Bricaud & Roux (1993), Otte (2007), Roux & Giralt (1991), Roux & Clerc (1991), Roux & al. (1995), and Tretiach (1998).


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Last modified: June, 15, 2023

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