New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia

Boom, P. P. G. van den; Alvarado, P.

doi:10.3390/d17030187

Open AccessArticle

New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia

by

P. P. G. van den Boom

^1,* and

P. Alvarado

^2,*

¹

Department of Research, Meise Botanic Garden, Nieuwelaan 38, B-1860 Meise, Belgium

²

Alvalab, Dr. Fernando Bongera st., Severo Ochoa bldg. S1.04, 33006 Oviedo, Spain

^*

Authors to whom correspondence should be addressed.

Diversity 2025, 17(3), 187; https://doi.org/10.3390/d17030187

Submission received: 12 September 2024 / Revised: 25 February 2025 / Accepted: 27 February 2025 / Published: 6 March 2025

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Abstract

:

In the present work, two species new to science, Bacidia subheterochroa and Bacidina terceirae, are reported among one hundred and five lichens and lichenicolous species reported from the islands Graciosa and Terceira (Azores Archipelago, Portugal). Updated phylogenetic studies of the family Ramalinaceae including these samples show that species of Bacidia are grouped in three different clades: Bacidia s. str. (type species B. rosella), Bacidina (type species Bn. phacodes), and Woessia (type species W. fusarioides). As a result, seven new combinations in the resurrected genus Woessia are here proposed. In addition, a comprehensive checklist of lichens and lichenicolous fungi of two of the islands (including locality and substrate) is provided, with five species being reported from Azores for the first time.

Keywords:

Macaronesia; Portugal; Ramalinaceae; taxonomy; biodiversity

1. Introduction

The present study deals with two species found during the catalogue of lichens and lichenicolous fungi of the islands of Graciosa and Terceira (Azores, Portugal). Graciosa Island belongs to the Portuguese archipelago of Azores (Figure 1). It only measures ca. 60 km², has its highest point at Pico do Facho (375 m asl), and is dominated by a 1.6 km wide caldera (The Caldera) in the southeast. It is one of the driest islands of the Azores, subjected to a mild climate with average daily temperatures of 14–20 °C, never going below 10 °C. The average annual precipitation is c. 840 mm, increasing from east to west. While the Azores archipelago is a major stronghold for remnants of the Laurisilvl Tertiary evergreen forest (formerly widely distributed in southern Europe) laurisilva is absent from Graciosa Island. For the location of the Azores archipelago in Macaronesia, see Figure 2.

The known lichen species of Graciosa were listed by Aptroot et al. [1]. Further recent information about the fungal species of Terceira can be found in van den Boom [2] and Etayo [3]. New records, as well as newly found species, from these islands were published after these authors, i.e., Crittendenia byssolomatis Diederich, van den Boom, and Millanes [4]. Other recent studies about lichens and lichenicolous fungi of the Azores include references [2,3,5,6,7,8]. Crustose lichens as well as many macro lichen genera are in need of revision for this archipelago. The few systematic studies published so far, dealing with the genera Usnea [9], Heterodermia [10], Anisomeridium [11], or Fissurina, Graphis and Topeliopsis [12], show that the taxonomical knowledge is still insufficient.

In the present work, the checklist of lichens and lichenicolous fungi of Graciosa and Terceira islands of the Azores is updated with new findings, thus showing increasing in biodiversity. To this end, the paper focuses on two species of the family.

Ramalinaceae not fitting any existing name. These are morphologically and phylogenetically compared with their closest relatives in the genera Bacidia and Bacidina in order to support the new names proposed for them.

The genus Bacidina was originally proposed because it differs morphologically and ecologically from Bacidia [13,14,15,16,17,18], but it was considered a synonym of Lichingoldia and Woessia on the basis of the one fungus one name policy, to the point that it was conserved against these names [16,19]. However, Kistenich et al. [20], using a five-gene dataset, found that the clades containing the type species of Bacidina and Woessia are not monophyletic (with a partial support with PP ≥ 0.95 but BS < 75). Ekman [17] recently reviewed the issue, pointing to the lack of information (especially from tropical regions) as the cause behind the failure to recover significant support for both clades, by other authors of [7,8,21,22,23].

2. Materials and Methods

Morphological studies—The specimens reported in the present study were recently (2021) collected by PvdB during a fieldtrip on the islands Graciosa and Terceira (Macaronesia), visiting 29 sites of lichenological interest (Tables S1 and S2), collecting lichens and lichenicolous fungi there. All specimens were studied by conventional macro- and microscopical techniques with hand-cut sections of the material mounted in tap water. Amyloid reactions were tested using Lugol’s iodine solution (K/I). Voucher specimens are kept in the herbarium of P. van den Boom, now deposited in BR herbarium (Meise Botanic Garden, Meise, Belgium), but a few specimens are deposited at B herbarium (ZE Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Berlin, Germany). Chorologic data of Macaronesia were taken from the existing checklists of the Azores [1,24], as well as other papers published after these works [2,3,7,25,26,27,28,29,30,31]. Species newly recorded from the Azores in the present work are marked with an asterisk.

Phylogenetic studies—Total DNA was extracted from dry specimens, employing a modified protocol based on Murray et al. [32]. PCR reactions [33] included 35 cycles with an annealing temperature of 54 °C. The primers, ITS1F and ITS4 [34,35], were employed to amplify the ITS rDNA region. LR0R and LR5 [36,37], were used for the 28S rDNA region, as well as mrSSU1 and mrSSU3R [38] for the mitochondrial small ribosomal subunit (mtSSU). PCR products were checked in 1% agarose gels, and amplicons were sequenced with one or both PCR primers. The sequences were manually corrected to remove reading errors in chromatograms.

Three different datasets were built: (1) one containing mtSSU, LSU, and RPB1 sequences of selected lineages of the family Ramalinaceae; (2) one containing only ITS sequences of Bacidia s. str. related to the samples; and (3) another one containing only ITS sequences of Bacidina related to the samples (datasets available at https://figshare.com/articles/dataset/READ1_sorted_BSF_0838_HTVCLDRXX_1_0023Bac_rub_1_S70637_fastqc_html/24298654). BLASTn [39] was used to select the sequences of the species most closely related to the samples from the International Nucleotide Sequence Database Collaboration public database ([INSDC, [40]). RPB1 sequences available were employed in the phylogenetic analyses of dataset 1 (despite that homologous sequences could not be produced from the samples studied), due to the valuable information to define generic limits provided by this gene. On the other hand, ITS rDNA was obtained from the samples but not employed in phylogenetic analyses of Ramalinaceae because this region is too variable to align distant genera correctly. The sequences retrieved are listed in Table 1, Table 2 and Table 3. The sequences first were aligned in MEGA 5.0 [41] with its Clustal W application and then realigned manually as needed to establish positional homology. Aligned loci were loaded in MrBayes 3.2.6 [42], where a Bayesian analysis was performed (dataset 1 partitions: mtSSU, LSU, RPB1; datasets 2–3 partitions: ITS; two simultaneous runs, four chains, temperature set to 0.2, sampling every 100th generation) until the average split frequencies between the simultaneous runs fell below 0.01 after 5.05 M (dataset 1), 0.22 M (dataset 2), and 3.17 M (dataset 3) generations. Finally, a full search for the best-scoring maximum likelihood tree was performed in RAxML 8.2.12 [43] using the standard search algorithm (same partitions, GTRCAT model, 2000 bootstrap replications). The significance threshold was set above 0.95 for posterior probability (PP) and 70% for bootstrap proportions (BPs).

3. Results

An annotated list of the 103 species found is reported in the Supplementary Materials (Table S2), with newly recorded taxa for at least one of the visited islands, including five species found for the first time in the Azores archipelago and two species new to science, described below. The phylogenetic studies of the family Ramalinaceae conducted to classify the new taxa (Figure 3) confirmed that species of Bacidia form, at least, three significantly different (and not directly related) clades, a result already suggested by previous works [17,20]. The first clade (1) contains the type species of Bacidia, B. rosella, and is therefore considered Bacidia s. str. The second one (2) contains the type species of two genera originally proposed for anamorphic states: Lichingoldia, (type species L. gyalectiformis = Bn. inundata) [44] and Woessia (type species W. fusarioides = Bn. Modesta) [45]. The third one (3) contains the type species of Bacidina Vězda [46], Bn. phacodes (Körb) Vězda. A sample identified as Bn. californica in public databases (H:Knudsen 6484, AFTOL-ID 4892) does not belong to any of these clades (it could be related to the clade formed by Toninia, Toniniopsis, Killiasia, Thalloidima, and Bibbya), so this species could represent a fourth genus different from the others, but the identity of this sample would need to be confirmed after appointing an epitype of Bn. californica.

The newly found species from Azores nested inside Bacidia and Bacidina, but they did not match any existing record in sequence databases (Figure 4 and Figure 5). Therefore, new names are proposed below for them.

3.1. Taxonomy

Bacidia subheterochroa van den Boom & P. Alvarado sp. nov. (Figure 6 and Figure 7)

MycoBank no.: MB 856424

Diagnosis: A species of Bacidia similar to B. heterochroa (Müll. Arg.) Zahlbr., which differs because of its squamulose thallus, the lack of enlarged cells at the outer rim of the excipulum; apothecia: dark brown to blackish; excipulum: prosoplectenchymatic with conglutinated hyphae, lacking crystals; epithecium: yellowish brown, K+ reddish, N−; hypothecium: pale to medium brown; ascospores acicular, often slightly curved, 35–50 × 2–3 μm, 6–8-septate; pycnidia: dark brown, up to ca. 50 μm diam., conidia filiform, strongly curved, 25–35 × 0.8 μm.

Type: Portugal Azores, Graciosa, E of Luz, Ribeira, Casa Perpétua, garden with mixed trees, shrubs, and wall of stones. Located at 39° 01.31′ N, 27° 59.23′ W, 85 m, 9 September 2021, P. & B. van den Boom 60426 (BR–holotype; AZU–isotype).

Description: Thallus rimose areolate, slightly squamulose, pale greenish grey to dark grey, up to 0.2 mm thick, upper surface slightly shiny to matt, ecorticated. Prothallus sometimes present and visible among the thallus parts as a dark rim. Photobiont chlorococcoid, cells 5–12 μm diam. Apothecia: Abundant, dispersed, or rarely confluent, roundish in outline, up to 0.8(–1) mm diam., thinly marginate when young; margin: 30–50 mm wide, paler than the disc, pale brown to yellowish brown, slightly shiny, sometimes flexuose; disc flat when young, becoming weakly convex, dark brown to blackish, never variously coloured; excipulum: hyaline to brownish, 20–40 μm wide, full of fine crystals, with conglutinated radiating cells at inner part, small prosoplectenchymatous cells with walls of 1.5–3 μm wide lumina, with isodiametric to ellipsoid lumina ca. 8 × 2.5 μm; epithecium: yellowish brown to dark brown, some parts hyaline, with fine crystals, K+ reddish, N–; hymenium: ca. 100 μm high, hyaline; hypothecium: pale to reddish brown; hamathecium of paraphyses, thin, 1–1.5 μm wide, septate, sometimes slightly branched, not anastomosed, mid-hymenium cells ca. 7–9 × 1–1.5 μm, tips usually slightly widened, up to 5 μm, yellow brown pigmented. Asci: Small cylindrical to slightly clavate, 45–55 × 8–14 μm, 8-spored, tholus staining dark blue in I with a paler blue, narrowly conical axial body, ocular chamber with a blunt body, surrounded by a rather small but strongly amyloid layer, not open at the apex. Ascospores: ± straight to rarely coiled in the ascus, acicular, rarely straight nearly always slightly curved, hyaline, 35–50 × 2–3 μm, 3–8-septate. Pycnidia: ca. 50 μm wide, ostiole margin medium brown in upper part; conidia: filiform, strongly curved, 25–35 × 0.8 μm, not septate.

Chemistry: K−, C−, P−, UV−, no chemical compounds detected. Arnoldiana—brown pigment in proper exciple, hypothecium, and epihymenium.

Etymology: The epithet refers to the similarity with Bacidia heterochroa.

Distribution and ecology: The new species is only known from the type locality where it grows abundantly on stones of an old wall.

Notes: Genetically, B. subheterochroa is close, but clearly different, from a group of specimens identified as B. heterochroa in databases (97.2% similarity in ITS rDNA, 98.86% in mtSSU). While some species in Bacidia seem to display great inter-individual variabilities in these genes (i.e., 96.6% among ITS sequences of B. schweinitzii), this is not the case for all of them (i.e., B. biatorina, B. laurocerasi, B. suffusa). In addition, the molecular identity of B. heterochroa needs to be confirmed, since there is more than one candidate lineage, and none of them come from the original locality (Buenos Aires, Argentina): the group of specimens named B. heterochroa—1 in the tree was collected in Georgia and Russia, while those named B. heterochroa—2 in the tree were found in South Korea. On the basis of the taxonomic uncertainty around B. heterochroa, as well as the deviant genetic profile of the sample studied in the present work and its morphological features, a new species name is proposed here for it.

Morphologically, B. subheterochroa is an inconspicuous species, easily mistaken for B. heterochroa, and microscopical control is needed to distinguish between both species with certainty. The latter has usually mottled apothecia with pale pink or pale orange parts on the disc, no crystals in the epithecium, and sometimes only a few small clusters of crystals in the excipulum; the purplish brown epithecium is N+ reddish and the outer rim of the excipulum contains a row of enlarged cells with lumina of up to 9 × 6 μm. Bacidia heterochroa is known from the USA mostly in coastal areas in the west and southeast, and it is widely distributed in tropical areas, Ekman [46]. Recently it has also been recorded from Macaronesia, Santa Maria, van den Boom [30], and found in the Caucasus by Gerasimova et al. [47]. Bacidia laurocerasi (Delise ex Duby) Zahlbr. is similar in habitus, but has a greyish thallus; epithecium: N−, hypothecium hyaline; ascospores: 2.5–4 μm wide; and conidia up to 17 μm long. Growing close to the new species is Lecania hutchinsiae (Nyl.) A.L. Sm. and an unknown Toninia sp. An unknown parasite is present on the thallus of the new species, with black pycnidia up to 0.15 mm diam., and brown conidia of simple cells of ca. 5 μm diam. or 1-septate conidia of ca. 8 × 5 μm.

Bacidina terceirae van den Boom & P. Alvarado sp. nov. (Figure 7 and Figure 8)

MycoBank no.: MB 856425

Diagnosis: Thallus thin minutely granular. Apothecia: 0.15–0.3(−0.35) mm diam., appressed to immersed, thinly marginate, sometimes persistent; margin: slightly paler than the disc, level with the disc in young apothecia; disc: pale cream, pale yellowish brown, plane to slightly convex. Ascospores: Fusiform, 1–3(−4)-septate, (12–)15–20(−22) × (1.5–)2–2.5(–2.8) μm. Pycnidia: Unknown.

Type: Portugal, Azores, Terceira, Angra do Heroismo, park in centre, with mixed trees and shrubs, including some exotic trees. Located at 38° 39.30′ N, 27° 12.57′ W, 275 m, 17 September 2021, P. & B. van den Boom 60754 (BR–holotype; AZU–isotype).

Description: Thallus, thin, minutely granular to scurfy granular-sorediate, granules 20–70 μm, ecorticate, greyish green to pale green, composed of spheroidal gonyocists covered with a cortex made of hyphae with a polygonal cell lumen, 4–5 μm wide, resembling a net; granules fixed to substrate by long and narrow hyphae. Photobiont chlorococcoid, 5–12 μm in diam. Apothecia: Abundantly present, biatorine, beige, greyish green, to pale green, 0.15–3(–0.35) mm in diam.; appressed to immersed in the thallus, margin is 40–60 μm wide, especially well developed in young apothecia, which is level with or slightly raised above the disc, usually slightly paler than the disc, but sometimes with pale brownish pigment, finally immarginate; disc is flat to slightly convex; excipular: hyphae radiating, the upper part made of branched and anastomosed hyphae with narrow lumina, 1 μm wide, with 1–2 outer layers of globose cells, 3–5 μm wide; basal part of the exciple made of branched and anastomosed hyphae with wide lumina, 3 μm wide and with 3–4 outer layers of globose to polygonal cell lumina, 4–8 μm wide; narrow hyphae looking like hair arise from the outer layer; without crystals, K+ yellowish, pale brownish pigment sometimes present; hypothecium: colourless, prosoplectenchymatous, with a dense textura intricata, cells 2–4 μm wide, with relative thick walls; subhymenium: colourless, no pigment present; hymenium: colourless, 35–50 μm thick, not gelatinized; epithecium: not differentiated; paraphyses: agglutinate, nor branched, neither anastomosed, filament 1–1.5 μm thick; apices: slightly swollen, 2–2.5 μm wide; asci: cylindrical to clavate, 8-spored, 18–32 × 8–11 μm, ascus Bacidia-type, with a wide conical axial masse; ascospores, fusiform with rounded ends to bacilliform, 1–3(–4)-septate, (12–)15–20(–22) × (1.5–)2–2.5(–2.8) μm. Pycnidia not seen.

Chemistry: Thallus K−, C−, P−, KC−. No crystals are present in apothecium or thallus. Inner exciple and subhymenium sometimes with pale brownish pigment, K + intensifying the pigmentation.

Etymology: The epithet refers to the island Terceira, a part of the Azores.

Distribution: This new Bacidina species is known from the westernmost European region. This species is found in hyperoceanic bioclimates, so far on only one island, Terceira.

Ecology: Bacidina terceirae is a corticolous species, grows in a parkland, on the bark of exposed roots very close to the soil, and often colonizes the surrounding substratum, such as the bark of tree bases. It has been found in the type locality abundantly on exposed roots of Ficus macrocarpa (holotype), on exposed roots of Chorisia speciosa, and on a trunk of Phoenix canariensis. Only few accompanying species have been found growing closer to the species: Lecania naegelii (Hepp) Diederich & van den Boom, Hyperphyscia adglutinata (Flörke) H. Mayrhofer & Poelt.

Notes: Genetically, Bn. terceirae is nested in a significantly monophyletic clade together with a sample of Bn. brittoniana (BG:Ekman 3657, from Kistenich et al. [20]) and several undetermined samples (PRA:JV24968, PRA:ZP30104, and PRA:JV24493, from Vondrák et al. [48]). However, the ITS rDNA sequence of Bn. terceirae is very different from that of Bn. brittoniana (92% similarity in 5.8S+ITS2 regions), so it is here supposed that these clades are reproductively isolated. Morphologically, Bn. brittoniana (Riddle) LaGreca & S. Ekman (= Bacidina varia S. Ekman) has an areolate, wrinkled or warted thallus, apothecia of 0.2–0.4(–0.6) mm diam., and much different ascospores of 20–50 × 1.5–2.5 μm, 3–7-septate. In the field, Bn. terceirae can be overlooked for Bn. delicata (Larbal. ex. Leight) V. Wirth & Vězda, as that species also has a finely granular, greenish thallus containing goniocysts, the excipulum is paraplectenchymatous and hyaline but pycnidia are usually abundantly present. Bacidina modesta (Vain.) S. Ekman is corticolous and grows often at the basis of trees, according to Smith et al. [49], as Bacidina sulphurella, but has a red-brown hypothecium and smaller ascospores. Within Bacidina, or Bacidia, a few species have a fine granular thallus, and all of them have apothecia much more prominently present, longer and wider ascospores, and grow on the bark of tree trunks (e.g., Bacidia biatorina (Körb.) Vain., B. rubella (Hoffm.) A. Massal. and B. iberica Aragón & I. Martinez).

Additional specimens examined: Portugal, Azores, Terceira, Angra do Heroismo, park in centre, with mixed trees and shrubs, including some exotic trees. Located at 38° 39.30′ N, 27° 12.57′ W, 275 m, 17 September 2021, P. & B. van den Boom 60761, 60813 (B; hb van den Boom).

3.2. New Combinations into Woessia:

Woessia adastra (Sparrius & Aptroot) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856433

Basionym: Bacidia adastra Sparrius & Aptroot, Lichenologist 35(4): 275 (2003) [MB#385914]

Specimen examined: Netherlands, Noord-Brabant, W of Mariahout, N of De Rijt, along small Salix forest, on wood of fence post, 51°22′25″ N, 5°32′32″ E, 12 m, 22 November 2009, P. & B. van den Boom 43635 (hb. v.d.Boom). Illustration: Sparrius & Aptroot, Lichenologist 35(4): 276 (2003).

Woessia brandii (Coppins & van den Boom) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856434

Basionym: Bacidia brandii Coppins & van den Boom, Lichenologist 34(4): 328 (2002) [MB#475096]

Specimen examined: The Netherlands; prov. Utrecht, SW of Baarn,

Pluismeer, on rotting Pinus stump among grassy vegetation, 51°10′54″ N, 5°14′29″ E, 12 m, 1 May 1994, P. van den Boom 15418 (hb. v.d. Boom) [isotypus]. Illustration: Ekman, Nordic Journal of botanic 5:16 (2023).

Woessia egenula (Nyl.) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856435

Basionym: Lecidea egenula Nyl., Flora, Regensburg 48: 147 (1865) [MB#390532}

Specimen examined: Netherlands, Noord-Brabant, Helmond, centre, nearby castle, along canal, on old brick wall, 51°28′39″ N, 5°39′ 10″ E, 20 m, 7 November 1987, P. van den Boom 5826 (hb. v.d. Boom). Illustration: Ekman, Nordic Journal of botanic 5:17 (2023)

Woessia neosquamulosa (Aptroot & Herk) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856436

Basionym: Bacidia neosquamulosa Aptroot & Herk, Lichenologist 31(2): 122 (1999) [MB#461080]

Specimen examined: Netherlands, Noord-Brabant, E of Valkenswaard, N of ‘t Leenderbos, ENE of Valkenhorst, row of mixed trees along field, on Sambucus, 51°21′45″ N, 5°31′34″ E, 25 m, 20 January 2008, P. & B. van den Boom 38889 (hb. v.d.Boom) [DNA]. Illustration: van den Boom, Bibliotheca Lichenologica 111: 49 (2021).

Woessia modesta (Zwackh ex Vain.) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 8564337

Basionym: Biatora modesta Zwackh, Lichenes exsiccati (Heidelberg): no. 332 (1850) [MB#830280]

syn. Woessia fusarioides D. Hawksw., Poelt & Tscherm.-Woess, in Hawksworth & Poelt, Pl. Syst. Evol. 154(3-4): 207 (1986)

Specimen examined: Netherlands, Noord-Brabant, N of Bergeyk, SW of Riethoven, N of road Eersel to Valkenswaard, mixed wood along field, on Quercus, 51°20′35″ N, 5°22′13″ E, 30 m, 24 June 2000, P. & B. van den Boom 24619 (hb. v.d.Boom). Illustration: Ekman, Nordic Journal of botanic 5: 16 (2023).

Woessia saxenii (Erichsen) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856438

Basionym: Bacidia saxenii Erichsen, Annls mycol. 39(2/3): 142 (1941) [MB#364539]

Specimen examined: Netherlands, Noord-Brabant, SW of Leende, ‘t Leenderbos, W side of main path, open area along Pinus forest, on stump of Picea, 51°18′55″ N, 5°30′38″ E, 30 m, 24 April 2000, P. & B. van den Boom 24283 (hb. v.d.Boom) Illustration: van den Boom, Herzogia 34(2): 521 (2021).

Woessia terricola (van den Boom & Alvarado) van den Boom & P. Alvarado comb. nov.

MycoBank no.: MB 856439

Basionym: Bacidina terricola van den Boom & Alvarado, Herzogia 32(2): 426 (2019) [MB#830042]

Specimen examined: Portugal, Azores: Faial, W of Castelo Branco, Morro de Castelo Branco, just east of the mountain, coastal area with abundantly Erica azorica and volcanic outcrops, 38°31′30″ N, 28°44′49″ W, 80 m, 4 April 2016, P. & B. van den Boom 55461 (hb. v.d. Boom) [isotype]. Illustration: van den Boom & Alvarado, Herzogia 32(2): 429 (2019).

4. Discussion

In the present work, the lichen checklist of Graciosa and Terceira islands was updated, adding new records and two newly discovered species, one nested inside Bacidia s. str., the other inside Bacidina s. str. The third clade of Bacidia-like species detected in previous works by Kistenich et al. [20] is here restored as the independent genus Woessia.

The present results, based on three genetic markers, mtSSU, LSU, and RPB1, produced significant Bayesian and ML support values for Bacidia, Bacidina, and Woessia using a dataset with 15, 11, and 9 putative species, respectively. The fully significant support found in the present work (not found in previous studies [17,20]) was maybe produced by the more complete dataset employed, which includes several unclassified sequences of Ramalinaceae generated by Kistenich et al. [50] from specimens collected in Indonesia, Malaysia, and Thailand.

Updated phylogenies of Bacidia and Bacidina were produced by Gerasimova et al. [47,51] and Czarnota and Guzow-Krzemińska et al. [52], respectively. Kondratyuk et al. [53] proposed four new genera in the family Ramalinaceae: Ivanpisutia, Coppinsidea, Vandenboomia, and Wolseleyidea. These authors also resurrected genera Lecaniella and Myrionora, but these were not included in the present analyses because of the scarce and incomplete sequence data in public databases.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17030187/s1. Table S1: Collecting sites at Graciosa and Terceira; Table S2: Annotated species list.

Author Contributions

Conceptualization, P.P.G.v.d.B. and P.A.; methodology, P.P.G.v.d.B. and P.A.; validation, P.P.G.v.d.B.; formal analysis, P.A.; investigation, P.P.G.v.d.B. and P.A.; data curation, P.P.G.v.d.B. and P.A.; writing—original draft preparation, P.P.G.v.d.B. and P.A.; writing—review and editing, P.P.G.v.d.B. and P.A.; visualization, P.P.G.v.d.B. and P.A.; supervision, P.P.G.v.d.B. and P.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not Applicable.

Data Availability Statement

All data generated for this study are contained in the text, figures, and tables.

Acknowledgments

An important contribution for this study was made by the laboratory http://www.alvalab.es and the Belgian botanical institute www.plantentuinmeise.be (accessed on 1 February 2025). We want to thank the two anonymous reviewers for their constructive comments, which were helpful for improving the manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. The islands of the Azores archipelago. C—Corvo; Fl—Floris; Fa—Faial; G—Graciosa; P—Pico; SJ—São Jorge; SM—São Miguel; SMa—Santa Maria; T—Terceira.

Figure 2. Location of Azores archipelago in Macaronesia.

Figure 3. A 50% majority rule mtSSU-LSU-RPB1 consensus phylogram of the family Ramalinaceae (Lecanorales) (with Scoliciosporum intrusum from the family Scoliciosporaceae as the outgroup) obtained using MrBayes from 37,875 sampled trees. Nodes were annotated if they were supported by ≥0.95 Bayesian posterior probability (left) or ≥70% maximum likelihood bootstrap proportions (right). Nonsignificant support values are exceptionally represented inside parentheses. The sequences newly generated in this study are in bold.

Figure 4. A 50% majority rule ITS consensus phylogram of clade I of genus Bacidia (Gerasimova et al. [47]) obtained using MrBayes from 1650 sampled trees. Nodes were annotated if they were supported by ≥0.95 Bayesian posterior probability (left) or ≥70% maximum likelihood bootstrap proportions (right). Nonsignificant support values are exceptionally represented inside parentheses. The sequences newly generated in this study are in bold.

Figure 5. A 50% majority rule ITS consensus phylogram of genus Bacidina obtained using MrBayes from 23,775 sampled trees. Nodes were annotated if they were supported by ≥0.95 Bayesian posterior probability (left) or ≥70% maximum likelihood bootstrap proportions (right). Nonsignificant support values are exceptionally represented inside parentheses. The sequences newly generated in this study are in bold.

Figure 6. Bacidia subheterochroa, holotype, habitus. Scale = 0.5 mm.

Figure 7. Details of apothecia: (A,B,D) Bacidina terceirae; (C,E) Bacidia subheterochroa. (A) = excipulum and hymenium; (B) = section through apothecium; (D) = ascospores; (C) = excipulum and hymenium; (E) = ascospores. Scale: (A,C) = 10 μm; (B) = 40 μm; (D,E) = 5 μm.

Figure 8. Bacidina terceirae, holotype, habitus. Scale = 1 mm.

Table 1. Specimens used in molecular phylogenetic studies of Ramalinaceae and their GenBank accession numbers. Samples sequenced in the present work are highlighted in bold characters.

Species	Specimen Voucher	mtSSU	LSU	RPB1
Aciculopsora longispora	CDS:Bungartz 3699	MN334224
Aciculopsora salmonea	BR:Lucking 17543	MG925842		MG926137
Aciculopsora srilankens	PDA:Weerakoon, G. WL15/2	MK400212
Bacidia absistens	BG:Ekman 3223, SE-93	MG925845		MG926139
Bacidia arceutina	BG:Ekman 3110, SE-23	MG925846	MG926041	MG926140
Bacidia cylindrophora	GZU:Ohmura 7091	MG925908
Bacidia ekmaniana	NY:1454, Lendemer 30488A	KX151746
Bacidia elongata	M:0182626	MW506352	MW493330
Bacidia fraxinea	GLM:0044145	MW506373	MW489428	MW540441
Bacidia rosella	BG:Ekman 3117	AY300877	AY300829	AY756412
Bacidia rubella	LG:DNA 581	JQ796831
Bacidia schweinitzii	AFTOL-ID 642	DQ972998	DQ782911	DQ782830
Bacidia schweinitzii	H:Lendemer 931 & Moody, AFTOL-ID 4969	KJ766354	KJ766527
Bacidia schweinitzii	MIN:Wetmore72619, SE-111		MG926045	MG926146
Bacidia sigmospora	BR:P.v.d.Boom 55090	MW622007
Bacidia sipmanii	LG:DNA 361	JQ796832	JQ796844
Bacidia sp. (as Phyllopsora sorediata)	BM:Wolseley & Aguirre-Hudson 3948, 1007	MG925905	MG926094	MG926196
Bacidia squamulosula	O:L-113543, SE-314	MG925856	MG926051	MG926152
Bacidia subheterochroa sp. nov.	P.v.d.B. 60426	PQ615684	PQ615682
Bacidia wellingtonii	O:L-204598	MG925853
Bacidina brittoniana	BG:Ekman 3657 SE-224		MG926050	MG926151
Bacidina californica	H:Knudsen 6484 AFTOL-ID 4892	KJ766356	KJ766529
Bacidina lacerata	O:L144583, Timdal 10213 519	MG925896
Bacidina medialis	BG:Ekman 3659 SE-225	MG925850	MG926044	MG926145
Bacidina phacodes	UPS:Ekman 3414 SE-78		MG926049
Bacidina sp.	O:Dahl, Kistenich, Timdal & Toreskaas AM-21	MW307634
Bacidina sp.	O: Barbosa, Haugan & Timdal 74	W307633
Bacidina sp.	PRA:JV24968	OK465496
Bacidina sp.	PRA:ZP30104	OK465497
Bacidina sp.	PRA:JV24493	OK465495
Bacidina sp. (as undetermined Ramalinaceae)	BM:1104063	MK400200
Bacidina sp. (as undetermined Ramalinaceae)	BORH: Wolseley P, Thues H & Vairappan CM 1.12.oQ	MK400201
Bacidina sp. (as undetermined Ramalinaceae)	BM:1104016	MK400192
Bacidina sp. (as undetermined Ramalinaceae)	BM:749824	MK400193
Bacidina sp. (as undetermined Ramalinaceae)	BORH: Wolseley P, Thues H & Vairappan CM 3.03.1	MK400206
Bacidina terceirae sp. nov.	P.v.d.B. 60754	PQ615685	PQ615683
Badimia dimidiata	BG:Lucking 16013	AY567774	MG926052
Badimia pallidula	HN:20170295_2	MW349653	MT791315
Badimia pallidula	HMAS:L146151, Wang 20192889	MT791324	MT791317
Badimia polillensis	Wang WWC386	MT791325	MT791318
Badimia sp.	20180134	MT791323	MT791316
Biatora meiocarpa	BG:Tonsberg 28317a BG	AM292710
Biatora subduplex	UBC:Bjork 13011 AFTOL-ID 4912	KJ766360	KJ766533	KJ766838
Biatora vernalis	O:L-165159, J.T. Klepsland JK09-L616	KF360418	KF360446
Biatora vernalis	BG:Tonsberg 23757	DQ838753	DQ838752
Biatora vetera	Knutsson 2002-080	AY567771
Bibbya albomarginata	O:L144851, Timdal 10481 5468	MG925927	MG926115	MG926212
Bibbya bullata	O:L68980, Elix & Streimann 40393 SE-251	MG925929	MG926116
Bibbya bullata	O:L144480, Bannister s.n., 4823	MG925928
Bibbya ruginosa	O:L139109, Timdal 10087 5469	MG925937	MG926121	MG926218
Bibbya vermifera	BG:Johansson 1619 SE-109	MG925852	MG926047	MG926148
Bilimbia lobulata	O:Rui Timdal 9169 O	AM292712	MG926062	MG926161
Cenozosia inanis	TROM:5478	MG925866	MG926066
Cliomegalaria symmictoides	P.v.d.Boom 55386	MW622006	MW621867
Cliostomum corrugatum	BG:Ekman 3115	AY567722	MG926067	MG926166
Cliostomum griffithii	LG:database 196	GU138667
Cliostomum sp. (as Ramalina fastigiata)	BG:Ekman 3616	AY756375	AY756360	AY756422
Cliostomum tenerum	UPS:Klinkenberg Jorgensen	AM292722
Coppinsidea alba (as Biatora veteranorum)	LG:database 24	GU138664
Coppinsidea pallens	UPS:Nordin 5640	AM292709
Coppinsidea sp. (as Biatora ligni-mollis)	LG:database 25	GU138665
Crocynia gossypina	BG:Andersen 91	AY567766
Crocynia pyxinoides	DUKE:0000020, AFTOL-ID 111	AY584615	AY584653
Eschatogonia angustiloba	O:SK1-286	MW307646
Eschatogonia angustiloba	O:AM-42	MW307643
Eschatogonia prolifera	O:L144799, Timdal 10429 447	MG925871		MG926170
Eschatogonia prolifera	O:L144577, Timdal 10207 446	MG925870	MG926070	MG926169
Killiasia sculpturata	O:L19092, Haugan & Timdal YAK17,30, 5470	MG925938	MG926122	MG926219
Krogia antillarum	O:L202829, AM39	MH174274
Krogia borneensis	B:2234	MH174277
Krogia coralloides	O:L21909, Krog & Timdal MAU51,83, SE-387	MG925875	MG926072	MG926173
Krogia coralloides	Diederich 18455	MH174278
Krogia isidiata	H:34385	MH174279
Lecania brialmontii	AAS:Convey 121	AM292726	MG926112	MG926209
Lecania cyrtella	BG:Ekman 3017	AY300891	AY300840	MG926175
Lecania dubitans	McCune herb. Hutchinson ID-933-08	AM292732
Lecania furfuracea	Palice herb. Palice 5995	AM292734
Lecania fuscella	LD:Arup L03046	AM292735
Lecania inundata	E:Coppins 19139 & Coppins	AM292740
Lecania naegelii	CBFS:Vondrak 247	AM292741
Lecania nylanderiana	UPS:Tibell 23168	AM292742
Lecania sp. (as Catillaria scotinoides)	E:Coppins 18298 & O’Dare	AM292720
Lopezaria versicolor	DUKE:0000023, AFTOL-ID 108	AY584622	AY584651
Lueckingia polyspora	BR:5483	MG925882	MG926082
Megalaria grossa	BG:Tonsberg 26038	AY762095	AY756356	AY756419
Megalaria laureri	BG:Ekman 3119, SE-125	MG925884		MG926182
Micarea sylvicola	BG:Ekman 3629	AY567769	AY756331	AY756392
Myxobilimbia sabuletorum	BG:Ekman 3091	AY567721	AY756346	AY756413
Parallopsora brakoae	O:L-144623, Timdal 10253	MG925891
Parallopsora labriformis	O:L-144789, Timdal 10419	MG925895		MG926188
Parallopsora leucophyllina	O:L-144645, 464	MG925897		MG926189
Parallopsora sp. (as undetermined Ramalinaceae)	PDA: SK1-667	MK400228
Parallopsora sp. (as undetermined Ramalinaceae)	PDA:Kistenich S. & Weerakoon G, Ne141	MK400210
Parallopsora sp. (as undetermined Ramalinaceae)	PDA:Weerakoon SK1-666	MK400225
Phyllopsora breviuscula	GZU:Kalb & Marcelli 515 1305	MG925893	MG926088	MG926186
Phyllopsora breviuscula	O:L-73990, 528	MG925892	MG926087	MG926185
Phyllopsora longiuscula	O:L-144803, 454	MG925899	MG926090	MG926191
Phyllopsora mauritiana	O:L-21214, SE-386	MG925900	MG926091	MG926192
Physcidia striata	ABL:Caceres & Aptroot 11640	MG925910	MG926098
Protoblastenia rupestris	BG:Johnsen 19.02.2002		AY756358
Psora decipiens	BG:Ekman 3327	AY567772	MG926102	AY756396
Psora rubiformis	BG:Ekman 3343	AY756374	AY756359
Rolfidium bumammum	O:L-202891, 4821	MG925930	MG926117	MG926213
Rolfidium coccocarpioides	O: Krog&Timdal MAU61/08	AY762096	AY756361
Scoliciosporum intrusum	BG:Ekman s.n.	AY567767	AY756329	AY756391
Scutula circumspecta	BG:Tonsberg 17554 SE-228	MG925848		MG926143
Scutula krempelhuberi	UPS:Wedin 6356	AY567789
Scutula miliaris	UPS:Wedin 6859	AY567790
Sporacestra pertexta	hb. Perez-Ortega 1040	MG925903	MG926093	MG926194
Sporacestra sp. (as Phyllopsora borbonica)	O:L-797	MG925890	MG926086	MG926184
Sporacestra sp. (as undetermined Ramalinaceae)	BM:1104062	MK400191
Sporacestra sp. (as undetermined Ramalinaceae)	PDA:Kistenich, S. & Weerakoon G. SK1-659	MK400223
Stirtoniella kelica	UPS:5757	MG925923
Thalloidima candidum	O:L25779, Bratli & Timdal 8733 SE-59	MG925932	MG926118	MG926215
Thalloidima physaroides	O:L15217, SE-250	MG925936	MG926120	MG926217
Thalloidima toninianum	O:L33356, SE-72		MG926125	MG926222
Toninia cinereovirens	O:Haugan & Timdal 7953	AY567724	AY756365	AY756429
Toninia populorum	BG:Ekman 3392 SE-274	MG925843	MG926039	MG926138
Toninia squalida	O:L29316, Haugan 4970 SE-70	MG925940	MG926123	MG926220
Toniniopsis aromatica	O:L19933, Haugan & Timdal 4819 SE-58	MG925926	MG926113	MG926210
Toniniopsis coelestina	O:L29319, Haugan 5985 SE-61	MG925933	MG926119
Toniniopsis obscura	UPS:Westberg TNW2182, SE-306	MG925943	MG926126	MG926223
Toniniopsis subincompta	BG:Ekman 3413 SE-100	MG925851	MG926046	MG926147
Tylothallia biformigera	BG:Ekman 3096, SE-28	MG925946	MG926129	MG926226
Waynea californica	UPS:Ekman L1486, SE-214	MG925947	MG926130
Woessia arnoldiana	BG:Ekman 3157 SE-118	MG925854	MG926048	MG926149
Woessia cf. sulphurella	LG:DNA 577	JQ796839
Woessia chloroticula	LG:DNA 580	JQ796833
Woessia chloroticula	B:Sipman 55079 AFTOL-ID 4990	KJ766357	KJ766530
Woessia delicata	LG:DNA 369	JQ796834	JQ796845
Woessia egenula	LG:DNA 489	JQ796836	JQ796846
Woessia inundata	BG:Ekman 3187 SE-87	MG925855		MG926150
Woessia modesta	DUKE:AFTOL-ID 1845	DQ986810	DQ986798	KJ766904
Woessia neosquamulosa	LG:DNA 491	JQ796838	JQ796848
Woessia terricola	BR:P.v.d.Boom 55461	MW622008

Table 2. Specimens used in molecular phylogenetic studies of Bacidia and their GenBank accession numbers. Samples sequenced in the present work are highlighted in bold characters.

Species	Herbarium	ITS
Bacidia areolata	Davydov 17428 & Yakovchenko (ALTB)	MW491455
Bacidia areolata	Gerasimova M-0182592 (M)	MH048614
Bacidia biatorina	Gerasimova M-0182570 (M)	MW523509
Bacidia biatorina	Knutsson 94-148 (hb. Knutsson)	AF282079
Bacidia biatorina	Otte GLM-0053198 (GLM)	MW523511
Bacidia biatorina	Urbanavichene & Urbanavichus M-0311922 (M)	MW523510
Bacidia caucasica	Otte GLM-0048447 (GLM)	MW523553
Bacidia heterochroa (1)	KBA-L-0000386	ON352606
Bacidia heterochroa (1)	KBA-L-0002727	ON352612
Bacidia heterochroa (1)	KBA-L-0002734	ON352613
Bacidia heterochroa (2)	Gagarina L-11635 (LE)	MW523516
Bacidia heterochroa (2)	Gerasimova M-0182575 (M)	MW523515
Bacidia heterochroa (2)	Otte GLM-0048864 (GLM)	MW523518
Bacidia heterochroa (2)	Otte GLM-0048909 (GLM)	MW523517
Bacidia kurilensis	Ezhkin M-0182620 (M)	MH048610
Bacidia kurilensis	Ezhkin M-0182621 (M)	MH048611
Bacidia kurilensis	Ezhkin M-0182622 (M)	MH048612
Bacidia laurocerasi	Alaska Spribille 26334 (KLGO)	MN483106
Bacidia laurocerasi	Ezhkin M-0308500 (M)	MW491460
Bacidia laurocerasi	Galanina M-0311952 (M)	MH048609
Bacidia laurocerasi	Otte GLM-0048211 (GLM)	MW523513
Bacidia laurocerasi	Otte GLM-0053624 (GLM)	MW523514
Bacidia laurocerasi	Urbanavichene & Urbanavichus M-0311924 (M)	MW523512
Bacidia laurocerasi	USA Wetmore 74318 (MIN)	AF282078
Bacidia lutescens	Ekman L1161 (LD)	AF282082
Bacidia schweinitzii	AFTOL-ID 642	DQ782850
Bacidia schweinitzii	Gerasimova M-0182579 (M)	MW491454
Bacidia schweinitzii	Gerasimova M-0182580 (M)	MH048613
Bacidia schweinitzii	Lendemer 29364 (NY1449)	KX151763
Bacidia schweinitzii	Lendemer 30548 (NY1452)	KX151761
Bacidia schweinitzii	Lendemer 31230A (NY1450)	KX151766
Bacidia schweinitzii	Lendemer 31238 (NY1451)	KX151764
Bacidia schweinitzii	Lendemer 31855 (NY1453)	KX151765
Bacidia schweinitzii	Shaheen (NY1451)	MG461696
Bacidia schweinitzii	Tripp 2614 (NY1448)	KX151762
Bacidia schweinitzii	Wetmore 72619 (MIN)	AF282080
Bacidia subheterochroa sp. nov.	P.v.d.B. 60426	PQ615680
Bacidia suffusa	Gerasimova M-0182593 (M)	MH048616
Bacidia suffusa	Gerasimova M-0182594 (M)	MH048617
Bacidia suffusa	Gerasimova M-0182601 (M)	MH048615
Bacidia suffusa	KBA-L-0000359	ON352605
Bacidia suffusa	KBA-L-0002776	ON352614
Bacidia suffusa	KBA-L-0002835	ON352616
Bacidia suffusa	Otte GLM-0048445 (GLM)	MW523551
Bacidia suffusa	Otte GLM-0048460 (GLM)	MW523552
Bacidia suffusa	Otte GLM-0048464 (GLM)	MW523555
Bacidia suffusa	Otte GLM-0048483 (GLM)	MW523554
Bacidia suffusa	Otte GLM-0052906 (GLM)	MW523548
Bacidia suffusa	Otte GLM-0052934 (GLM)	MW523547
Bacidia suffusa	Otte GLM-0052950 (GLM)	MW523549
Bacidia suffusa	Otte GLM-0055113 (GLM)	MW523550
Bacidia suffusa	Tucker 17000 (M)	MH048618
Bacidia suffusa	Wetmore 40219 (M)	MH048619
Bacidia suffusa	Wetmore 74771 (MIN)	AF282091
Bacidia wellingtoni	O:L-204598	MG925953

Table 3. Specimens used in molecular phylogenetic studies of Bacidina and their GenBank accession numbers. Samples sequenced in the present work are highlighted in bold characters.

Species	Herbarium	ITS
Bacidina adastra	PRA:JV24569	OK332938
Bacidina adastra	Malicek 16567	PQ524005
Bacidina adastra	Malicek 13928	OQ717318
Bacidina adastra	GPN:4961	MG461694
Bacidina arnoldiana	GPN:5301	JN972439
Bacidina arnoldiana	GPN:5384	JN972441
Bacidina arnoldiana	GPN:4895	JN972440
Bacidina arnoldiana	AFTOL-ID 1845	HQ650650
Bacidina arnoldiana	DF230	KX098347
Bacidina arnoldiana	DF207	KX098343
Bacidina arnoldiana	LIFU049-16	KX132958
Bacidina arnoldiana	DF89	KX098348
Bacidina arnoldiana	DF212	KX098346
Bacidina arnoldiana	LIFU063-16	KX132972
Bacidina arnoldiana	PRA:Vondrak 24753	OQ717717
Bacidina arnoldiana	Malicek 14733	OP730571
Bacidina arnoldiana	DF210	KX098344
Bacidina arnoldiana	PRA:JV24248	OK332939
Bacidina arnoldiana	P.v.d. Boom 47546a	KX239021
Bacidina arnoldiana	P.v.d. Boom 47546b	KX239020
Bacidina assulata	Malicek 15911	PP768145
Bacidina brandii	Czarnota 7599	KX239029
Bacidina brandii	P.v.d. Boom 44815	KX239025
Bacidina caligans	UPS:Johansson 21	AF282096
Bacidina caligans	GPN 4961	JN972442
Bacidina caligans	P.v.d. Boom 44973	KX239024
Bacidina chloroticula	PRA:Palice 32914	OQ717320
Bacidina chloroticula	BG:Toensberg 18642	AF282098
Bacidina chloroticula	PRA:Vondrak 26080	OQ717319
Bacidina delicata	BG:1996 Fritz	AF282097
Bacidina delicata	LG:DNA 369	JQ796854
Bacidina egenula	GPN:8337	KY379234
Bacidina egenula	GPN:8304	KY379235
Bacidina egenula	GPN:8314	KY379233
Bacidina egenula	PRA:JV24938	OL396592
Bacidina egenula	BG:Ekman 3003	AF282095
Bacidina flavoleprosa	PRA:Palice 32275	OQ717718
Bacidina flavoleprosa	GPN:5329	JN972443
Bacidina inundata	BG:Ekman 3187	AF282094
Bacidina iqbalii	LAH:36418-TYPE	MT952885
Bacidina lacerata	O:L144583	MG925993
Bacidina margallensis	LAH:36417	NR_182351
Bacidina mendax	GPN:4888-TYPE	JN972444
Bacidina mendax	Malicek 15875	OQ918722
Bacidina mendax	PRA:Palice 30714	OQ717321
Bacidina mendax	Malicek 1769	KX239016
Bacidina mendax	Palice 13638	KX239028
Bacidina mendax	PRA:Vondrak 24102	OQ717722
Bacidina mendax	PRA:Vondrak 24106	OQ717721
Bacidina mendax	Malicek 16485	PQ524006
Bacidina mendax	Czarnota 7406	KX239023
Bacidina neosquamulosa	UPS:L715988	OP256854
Bacidina neosquamulosa	P.v.d.Boom 41056	KX239026
Bacidina neosquamulosa	LG:DNA 490	JQ796855
Bacidina neosquamulosa	P.v.d.Boom44999	KX239018
Bacidina phacodes	Czarnota 8264	KX239036
Bacidina phacodes	BG:Ekman 3414	AF282100
Bacidina phacodes	PRA:Vondrak 25994	OQ717322
Bacidina phacodes	PRA:Vondrak 24463	OQ717723
Bacidina pycnidiata	Malicek 255	KX239034
Bacidina pycnidiata	Lubek s.n.	KX239032
Bacidina pycnidiata	Czarnota 4157-TYPE	KX239033
Bacidina sp.	WSL:DF223	KX098339
Bacidina sp.	WSL:DF80	KX098341
Bacidina sp.	WSL:DF72	KX098340
Bacidina sp.	PRA:Vondrak 24312	OQ717719
Bacidina sp.	PRA:Vondrak 25879	OQ717720
Bacidina sp.	PRA:JV24247	OK332883
Bacidina sp.	PRA:JV24915	OK332884
Bacidina sp.	PRA:JV24921	OK332885
Bacidina sp.	PRA:Palice 32056	OQ717726
Bacidina sp.	PRA:Palice 33646	OQ717327
Bacidina sp.	PRA:Palice 26142	OQ717326
Bacidina sp.	PRA:Palice 31392	OQ727426
Bacidina sp.	Malicek 14355	PP410490
Bacidina sp.	PRA:Vondrak 25004	OQ717324
Bacidina sp.	Vondrak 19891	MT803542
Bacidina sp. (as Bacidia viridifarinosa)	BV:MSCBU	PP889388
Bacidina sp. (as Lecidea circumpallens)	UPS:L062093	OP256856
Bacidina sulphurella	GPN:5275	JN972445
Bacidina sulphurella	GPN:5967	JN972446
Bacidina sulphurella	GPN:7246	JN972447
Bacidina terceirae	P.v.d.B. 60754	PQ615681
Bacidina terricola	BR: van den Boom 55461	NR_172202
Bacidina violacea	PRA:Palice 31173	OQ717328
Toninia cinereovirens	O:Haugan & Timdal 7953	AF282104

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Boom, P.P.G.v.d.; Alvarado, P. New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia. Diversity 2025, 17, 187. https://doi.org/10.3390/d17030187

AMA Style

Boom PPGvd, Alvarado P. New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia. Diversity. 2025; 17(3):187. https://doi.org/10.3390/d17030187

Chicago/Turabian Style

Boom, P. P. G. van den, and P. Alvarado. 2025. "New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia" Diversity 17, no. 3: 187. https://doi.org/10.3390/d17030187

APA Style

Boom, P. P. G. v. d., & Alvarado, P. (2025). New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia. Diversity, 17(3), 187. https://doi.org/10.3390/d17030187

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New Species of Bacidia s.l. from the Azores and the Resurrection of Genus Woessia

Abstract

1. Introduction

2. Materials and Methods

3. Results

3.1. Taxonomy

3.2. New Combinations into Woessia:

4. Discussion

Supplementary Materials

Author Contributions

Funding

Institutional Review Board Statement

Data Availability Statement

Acknowledgments

Conflicts of Interest

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