Introduction
The southern polar regions include not only the Antarctic continent, with less than c. 1% surface area suitable for terrestrial vegetation (Peat et al. Reference Peat, Clarke and Convey2007), but also the climatically milder maritime Antarctic Peninsula and the islands in the South Atlantic, as well as the islands in the southern Indian Ocean, the New Zealand shelf islands (Auckland Islands and Campbell Island), Macquarie Island (Lauer et al. Reference Lauer, Rafiqpoor and Frankenberg1996; Brummitt et al. Reference Brummitt, Pando, Hollis and Brummitt2001) and the subantarctic subregions along the mountain ranges in southern South America (Morrone Reference Morrone2000).
These milder and often ice-free areas are characterized by the absence of arboreal vegetation and are colonized by specialized and cold-adapted pioneer vegetation, including a high density of mostly rock-dwelling crustose lichens acting as pioneer organisms (Hertel Reference Hertel1984; Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020). Lichens, as the dominant elements of Antarctic terrestrial vegetation, are represented by 300–400 species in the continent, according to the BAS (British Antarctic Survey) database, and they have been recorded in many studies during the last hundred years (e.g. Øvstedal & Smith Reference Øvstedal and Smith2001; Castello Reference Castello2003; Hertel Reference Hertel2007; Peat et al. Reference Peat, Clarke and Convey2007; Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2010, Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012; Colesie et al. Reference Colesie, Green, Türk, Hogg, Sancho and Büdel2014; Halıcı et al. Reference Halıcı, Bartak and Güllü2018, Reference Halıcı, Kahraman, Kistenich and Timdal2021; Wagner et al. Reference Wagner, Brunauer, Bathke, Cary, Fuchs, Sancho, Türk and Ruprecht2021). An even higher diversity of lichens can be estimated for southern South America (sSA, including Falkland Islands: Fryday et al. Reference Fryday, Orange, Ahti, Øvstedal and Crabtree2019; Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020; Etayo et al. Reference Etayo, Sancho, Gómez-Bolea, Søchting, Aguirre and Rozzi2021; Fryday Reference Fryday2022). However, the biodiversity of lichens in the Antarctic regions is still not fully known. New species are continuing to be described, which has become possible especially through the use of modern techniques such as DNA barcoding (Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020; Halıcı et al. Reference Halıcı, Kahraman, Kistenich and Timdal2021, Reference Halıcı, Güllü, Yiğit and Barták2022; Lagostina et al. Reference Lagostina, Andreev, Dal Grande, Grewe, Lorenz, Lumbsch, Rozzi, Ruprecht, Sancho and Søchting2021). These techniques also have the potential to reveal whether species for which only an endemic distribution was previously indicated are in fact bipolar, along continents (South to North America) or even globally distributed. This also applies to a small number of lecideoid lichen species such as Lecidella siplei (C. W. Dodge & G. E. Baker) May. Inoue or Lecidea polypycnidophora U. Rupr. & Türk which were previously described as endemic to Antarctica (Castello Reference Castello2003; Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2010). More recent studies have demonstrated that Lecidella siplei has a bipolar distribution and Lecidea polypycnidophora an alpine (North American)-southern polar distribution (Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012; Hale et al. Reference Hale, Fisher, Keuler, Smith and Leavitt2019). These unresolved biogeographical connections between Antarctica and Svalbard (Norway; L. siplei), as well as montane regions of western North America (L. polypycnidophora, L. andersonii Filson), show that potential colonization routes seem to exist and that the timing of dispersal events may also be highly relevant (e.g. Buschbom Reference Buschbom2007; Fernandez-Mendoza & Printzen Reference Fernandez-Mendoza and Printzen2013; Hale et al. Reference Hale, Fisher, Keuler, Smith and Leavitt2019; Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020).
The globally distributed genus Lecidella Körb. is represented by c. 50 recognized species and belongs to the family Lecanoraceae (Knoph & Leuckert Reference Knoph, Leuckert, Nash, Ryan, Diederich, Gries and Bungartz2004). This is not only clearly supported by molecular data (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015), but also by the quite similar ascus type. However, the Lecidella-type ascus also occurs in the genus Japewiella (Printzen Reference Printzen1999). It differs from the Lecanora-type ascus by an amyloid tholus and a narrow axial body converging towards the apex (Knoph & Leuckert Reference Knoph, Leuckert, Nash, Ryan, Diederich, Gries and Bungartz2004), in contrast to a thickened and also amyloid tholus at the apex, but with a broad non-amyloid axial mass (Wirth et al. Reference Wirth, Hauck and Schultz2013). The genus is better distinguished by the extremely lax and rarely capitate paraphyses (Wirth et al. Reference Wirth, Hauck and Schultz2013). Initially, Hertel & Leuckert (Reference Hertel and Leuckert1967) treated Lecidella as a subgenus of the genus Lecidea, but two years later they elevated it to genus level (Hertel & Leuckert Reference Hertel and Leuckert1969) because of its secondary chemistry, which differs from Lecidea by the presence of chlorinated norlichexanthones in many species. This genus is considered taxonomically difficult because of its high variation and/or plasticity in diagnostic characters (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015). The genus Lecidella is currently represented by nine species from the subantarctic region (Knoph & Leuckert Reference Knoph and Leuckert1994; Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020; Etayo et al. Reference Etayo, Sancho, Gómez-Bolea, Søchting, Aguirre and Rozzi2021; Fryday Reference Fryday2022) and around six for maritime and continental Antarctica (Castello Reference Castello2003; Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012; Fryday Reference Fryday2022).
Globally distributed genera in lichen-forming fungi repeatedly show disjunct distributions caused, for instance, by vicariance and mid-distance dispersal (Lücking et al. Reference Lücking, Papong, Thammathaworn and Boonpragob2008), or by transition from the Arctic to Patagonia (sSA) in the Pleistocene, resulting in cryptic speciation (Fernandez-Mendoza & Printzen Reference Fernandez-Mendoza and Printzen2013). Although this is not a common occurrence, Ruprecht et al. (Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012, Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020) reported a small number of endemic taxa and locally differentiated subgroups for the southern polar regions (southern South America, continental Antarctica) for the genera Lecidea and Lecidella.
Here we describe two new species of the genus Lecidella from maritime Antarctica and southern Patagonia, one with a North American-maritime Antarctic distribution and one with a so far exclusively southern South American-maritime Antarctic distribution, as well as several other maritime Antarctic accessions.
Material and Methods
Site descriptions (Fig. 1)
James Ross Island, located in the North-East Antarctic Peninsula region, has a cold, polar-continental climate (Martin & Peel Reference Martin and Peel1978) because of the Trinity Peninsula Mountains (Antarctic Peninsula) that shield the island from precipitation (Davies et al. Reference Davies, Glasser, Carrivick, Hambrey, Smellie and Nývlt2013). Precipitation estimates range from 200 to 500 mm per year (van Lipzig et al. Reference van Lipzig, King, Lachlan-Cope and van den Broeke2004) and, therefore, James Ross Island is considered a semi-arid environment.
Figure 1. Collection sites of Lecidella species in maritime Antarctica, the subantarctic areas of southern South America (Chile), and North America. Turquoise triangles = Lecidella ayazii; pink circles = Lecidella drakensis. In colour online.
Horseshoe Island is located in Marguerite Bay within the West Antarctic Peninsula. Gaul Cove is located on the east, while Lystad Bay is on the west coast of the island. The glacier-free regions are mainly composed of plutonic rocks consisting of granite and gabbro, banded gneiss and granitic gneiss belonging to the metamorphic complex. Sediments and moraines are also formed because of glacial movements on the island. The north of the island consists of rocks of a more diverse origin, whereas the mountains in the south are ponderous granites.
The two areas in southern Patagonia (Chile, sSA) are located in the Región de Magallanes y de la Antártica Chilena: Laguna Blanca at the volcanic Rock Morro Chico and the western part of the Torres del Paine National Park. Both areas are dominated by siliceous substrata and are influenced by the unique climate conditions for Patagonia, caused by strong westerly winds from the Pacific Ocean (Silva et al. Reference Silva, Rojas and Fedele2009).
The specimens are deposited at the Erciyes University Herbarium Kayseri, Turkey (ERCH), at the herbarium of the University of Salzburg, Austria (SZU) and at the herbarium of Michigan State University (MSC; Table 1).
Table 1. Voucher information and GenBank Accession numbers of the investigated specimens of the genus Lecidella collected in the subantarctic areas of southern South America (Chile), maritime Antarctica and northern Canada.
Morphological analyses
Light microscopic investigations were carried out with a Leica DVM6 digital microscope, a LEITZ Laborlux S and an Olympus BX53 (with an Olympus OM-D, E-M1 mark II camera). Photographic images were made of thallus morphology and anatomical features of asci and ascospores. Cross-sections of apothecia (12 μm thick) were prepared using a Leitz Kryomat 1703 freezing microtome. Measurements of anatomical structures always refer to water mounts, with at least 20 measurements made for all investigated anatomical structures. Spore measurements are given as: (minimum) most frequent (80%) (maximum).
Chemical analyses
Secondary metabolites were identified by thin-layer chromatography (TLC) using solvents A (toluene:1,4-dioxane:glacial acetic acid, 36:9:1) and C (toluene:glacial acetic acid, 20:3 (Huneck et al. Reference Huneck, Yoshimura, Huneck and Yoshimura1996; Orange Reference Orange2001). Spot tests were as follows: KOH, 10% (K), nitric acid solution, 35% (N), saturated potassium hypochlorite solution (C) and Lugol's iodine solution, 50% (I). KC: at first K was applied and then C within 10 s.
DNA amplification, sequencing and phylogenetic analyses
Total DNA was extracted from individual thalli using the DNeasy Plant Mini Kit (Qiagen) following manufacturer instructions. The internal transcribed spacer (ITS) regions of the mycobionts’ nuclear ribosomal DNA (nrITS) were sequenced and amplified using the primers ITS1F (Gardes & Bruns Reference Gardes and Bruns1993) and ITS4 (White et al. Reference White, Bruns, Lee, Taylor, Innis, Gelfand, Sninsky and White1990) with standard conditions. PCR products were sent to Eurofins Genomics (Germany) for sequencing.
The sequences were assembled and edited using Geneious Pro v. 6.1.8 (www.geneious.com), aligned with MAFFT v. 7.017 (Katoh et al. Reference Katoh, Misawa, Kuma and Miyata2002) and placed in context of the most recently published phylogenies of Ruprecht et al. (Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020) and Fayyaz et al. (Reference Fayyaz, Afshan, Niazi, Khalid and Ruprecht2022) based on the concept of Zhao et al. (Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015). The maximum likelihood analysis (ML) was performed using the IQ-TREE web server (Trifinopoulos et al. Reference Trifinopoulos, Nguyen, von Haeseler and Minh2016) with default settings (ultrafast bootstrap analyses (Hoang et al. Reference Hoang, Chernomor, von Haeseler, Minh and Vinh2017), 1000 BT alignments, 1000 max. iterations, min. correlation coefficient: 0.99, SH-aLRT branch test with 1000 replicates) and presented as a consensus tree. The best-fit model according to BIC, Tne + I + G4, was selected with the implemented model finder (Kalyaanamoorthy et al. Reference Kalyaanamoorthy, Minh, Wong, von Haeseler and Jermiin2017) of the program IQ-TREE. Bayesian phylogenies were inferred using the Markov chain Monte Carlo (MCMC) procedure as implemented in MrBayes v. 3.2. (Ronquist & Huelsenbeck Reference Ronquist and Huelsenbeck2003). The analysis was performed assuming the general time reversible model of nucleotide substitution including estimation of invariant sites and a discrete gamma distribution with six rate categories (GTR + I + Γ; Rodriguez et al. Reference Rodriguez, Oliver, Marin and Medina1990). Two runs with 2 million generations each starting with a random tree and employing four simultaneous chains were executed. Every 1000th tree was saved into a file. Subsequently, the first 25% of trees was deleted as the ‘burn-in’ of the chain. A consensus topology with posterior probabilities for each clade was calculated from the remaining 1501 trees. The phylogenies were visualized with the program FigTree v. 1.4.3 (Rambaut Reference Rambaut2014).
Results
Phylogenetic analyses
The final data matrix of this phylogeny contains 61 sequences of the ITS marker with a length of 538 characters, and was rooted with species of the genera Carbonea (Hertel) Hertel and Lecanora Ach. The phylogeny (Fig. 2) is divided into four main clades: L. stigmatea (Ach.) Hertel & Leuckert, L. enteroleucella (Nyl.) Hertel, L. elaeochroma (Ach.) M. Choisy (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015) and Lecidella sp. nov. (Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020). Five accessions from the maritime Antarctic were located in the very heterogeneous clade of L. stigmatea. One of these accessions (JR._0.303) can be assigned to L. siplei. Both new species described here are part of the L. elaeochroma clade and form two strongly supported distinct clades. The terricolous species Lecidella ayazii sp. nov., which occurs in maritime Antarctica, as well as in the alpine areas of the La Sal Mountains, Utah, USA and the Kivalliq Region, Nunavut in the north of Canada, is sister to accessions of L. elaeochroma, L. effugiens (Nilson) Knoph & Hertel and L. elaeochromoides (Nyl.) Knoph & Hertel. Lecidella drakensis sp. nov., restricted to southern Patagonia (sSA) and maritime Antarctica, is sister to a heterogeneous group consisting of accessions assigned to L. wulfenii (Hepp) Körb, L. elaeochroma, L. euphorea (Flörke) Hertel and L. flavosorediata (Vĕdza) Hertel & Leuckert. Lecidella meiococca (Nyl.) Leuckert & Hertel is basal to this highly supported group.
Figure 2. Phylogenetic analysis of ITS sequences of the genus Lecidella with the newly described species L. ayazii (turquoise) and L. drakensis (pink), and five other accessions (bold) integrated in the species concepts of Zhao et al. (Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015) and Ruprecht et al. (Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020). Maximum likelihood (ML) bootstrap values ≤ 95 were directly mapped on the Bayesian tree with posterior probability values ≤ 0.95 (branches in bold). Branch tips are labelled with voucher ID/Accession number_species_OTU. In colour online.
Taxonomy
Lecidella ayazii Halıcı & U. Rupr. sp. nov.
MycoBank No.: MB 851244
Differing from L. wulfenii by having a golden brown instead of a reddish or orange-brown hypothecium, growing on mosses over a siliceous instead of calcareous substratum and by a different chemistry, and from L. effugiens and L. elaeochromoides by growing on mosses instead of on rocks, and by having smaller apothecia and a different chemistry. It also differs from L. elaeochroma by growing on mosses instead of on bark and by having an inspersed hymenium.
Type: Antarctic Peninsula, James Ross Island, Berry Hill Mesa, 63°48′42.0″S, 57°50′5.4″W, 345 m a.s.l., 23 January 2017, M. G. Halıcı, (ERCH—holotype; ERCH—isotypes). GenBank Accession no.: OQ534855.
(Fig. 3)
Figure 3. Diagnostic characters of Lecidella ayazii. A, typical thallus and apothecia. B, cross-section of apothecium. C, Lecidella-type ascus and ascospores. Scales: A = 1 mm; B = 100 μm; C = 10 μm. In colour online.
Thallus crustose, granulose to rimose, up to 0.2 mm thick. Areoles distinct, almost squamule-like, irregular. Surface rough, pruinose. Colour greyish to chalky white.
Apothecia lecideine, black, abundant and evenly distributed, sessile, constricted at the base, up to 0.8 mm. Disc dull, black, epruinose, slightly convex, distinct margin. Margin thin, not raised, ±level with the disc, smooth. Exciple in section up to 50 μm thick, blue-green to blackish, I−, dark green (black) margin. Epihymenium dark green to bluish green, up to 20 μm, K−, N+ red (Cinereorufa-green). Hymenium hyaline, inspersed with oil droplets, up to 100 μm. Subhymenium light brown, up to 10 μm. Hypothecium golden brown, inspersed with crystals. Paraphyses hyaline, simple, slightly agglutinated, flexuous, vertical, partially inspersed. Apical cell slightly thickened up to 5.0 μm, dark green. Asci Lecidella-type, slightly clavate with dark amyloid tholus, eight unicellular spores, up to 40–50 μm. Ascospores simple, hyaline, broadly ellipsoid, (9–)10(–12) × (5–)6–7(–8) μm, l/w: 10.2/5.8 μm.
Conidiomata not observed.
Chemistry
Atranorin, thuringione and lichenxanthone by TLC. Spot test: thallus K+ yellow, C−, KC−.
Etymology
Named in honour of Çağan Ayaz Halıcı, dear son of the last author, who was born during his father's Antarctic expedition to James Ross Island in 2017.
Distribution and habitat
Lecidella ayazii is currently known from James Ross Island, located in the NE part of the Antarctic Peninsula, and Horseshoe Island in the SW; where it occurs on soil or sometimes on mosses from 5 to 345 m a.s.l., especially in humid habitats such as near streams. It is also known from the La Sal Mountains, Utah, USA and the Kivalliq Region, Nunavut, Canada, growing on mosses over siliceous substrata (Fig. 1).
Notes
Lecidella ayazii forms a distinct and highly supported clade of five accessions from maritime Antarctica as well as one accession from alpine areas of the La Sal Mountains, Utah, USA and two accessions from the Kivalliq Region, Nunavut in the north of Canada; it is placed in the L. elaeochroma clade (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015). According to Øvstedal & Smith (Reference Øvstedal and Smith2001), one of two muscicolous species of Lecidella known from Antarctica is L. wulfenii, which is also part of the L. elaeochroma clade, but the latter species has a different chemistry, an orange-brown (McCune Reference McCune2017) or reddish brown hypothecium and occurs over calcareous substrata (Knoph & Leuckert Reference Knoph, Leuckert, Nash, Ryan, Diederich, Gries and Bungartz2004; Wirth et al. Reference Wirth, Hauck and Schultz2013), in contrast to the newly described species which occurs over siliceous substrata. Furthermore, L. ayazii and L. wulfenii are phylogenetically placed in distinct clades and are not closely related. The second Antarctic species growing on mosses is L. siplei, which belongs to the L. stigmatea clade and is therefore clearly distinct from L. ayazii. Morphologically it differs with an often dark-pigmented (grey) thallus and a different chemistry (Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012; Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015). Two other phylogenetically related species, L. effugiens and L. elaeochromoides, grow on rocks, have a different chemistry and larger apothecia (Knoph & Mies Reference Knoph and Mies1995). The most closely related species is L. elaeochroma which shares a partially similar chemistry (atranorin, thuringione), but does not have an inspersed hymenium and typically grows only on bark (Wirth et al. Reference Wirth, Hauck and Schultz2013).
Additional specimens examined
Antarctica: Antarctic Peninsula: Horseshoe Island, 5 m a.s.l., 67°48′30″S, 67°17′39″W, M. G. Halıcı HS 0.009 (OQ534852; ERCH); James Ross Island, 260 m a.s.l., 63°49′80″S, 57°54′11″W, M. G. Halıcı JR 0.062 (OQ534850, ERCH); ibid., 292 m a.s.l., 63°49′46.2″S, 57°54′21.6″W, M. G. Halıcı JR 0.323 (OQ534851, ERCH); ibid., 142 m a.s.l., 63°48′24.9″S, 57°50′27.6″W, M. G. Halıcı JR 0.340 (OQ534853; ERCH).—Canada: Nunavut, Kivalliq Region, 9 m a.s.l., 62°52ʹ49.2″N, 92°08ʹ48.8″W, Fryday, A. M., McMullin, R. T, Allen, J., Sokoloff, P. Fryday_11820 (OR648670, MSU); ibid., 27 m a.s.l., 62°53ʹ24.1″N, 92°09ʹ24.1″W, Fryday, A. M., McMullin, R. T., Allen, J., Sokoloff, P. Fryday_11935 (OR648671, MSU) (Table 1).
Lecidella drakensis U. Rupr. & Halıcı sp. nov.
MycoBank No.: MB 851245
Differing from L. elaeochroma by having much smaller apothecia, from L. flavosorediata by the absence of soralia, from L. wulfenii by having more oblong ascospores, from L. euphorea by having a dark green to bluish green epihymenium, and from L. meiococca by having a thinner thallus. All the above-mentioned species also differ from L. drakensis in their chemistry.
Type: Chile, Región de Magallanes y de la Antártica Chilena, Patagonia chilena, Torres del Paine National Park, eastern slope of Co. Ferrer, 51°7′26″S, 73°8′33″W, on siliceous rock, 239 m a.s.l., 12 February 2018, U. Ruprecht (SZU—holotype; SZU, ERCH—isotypes). GenBank Accession no.: MK620158. Paratype (TLC): Antarctic Peninsula, James Ross Island. GenBank Accession no.: OQ534854 (Table 1).
(Fig. 4)
Figure 4. Diagnostic characters of Lecidella drakensis. A, typical thallus and apothecia. B, cross-section of apothecium. C, Lecidella-type ascus and ascospores. Scales: A = 1 mm; B = 100 μm; C = 10 μm. In colour online.
Thallus crustose, granulose to rimose, well developed, up to 0.2 mm thick. Areoles indistinct, irregular. Surface rough, pruinose. Colour whitish to beige.
Apothecia lecideine, mainly scattered, globular and sometimes merged, black, sessile, constricted at the base, up to 0.7 mm. Disc dull, black, pruinose, slightly convex when older and large, prominent and slightly rough margin. Exciple in section up to 80 μm thick, hyaline, I−, radiate hyphae, dark green (to black) margin, extending as a dark brown and wider margin along the base. Epihymenium dark green to bluish green, up to 10 μm, K−, N+ red. Hymenium hyaline, up to 70 μm, amyloid. Subhymenium light brown, 10 μm, I−. Hypothecium golden brown, inspersed with crystals, I−. Paraphyses hyaline, simple, flexuous, vertical, slightly agglutinated, 1–2 μm diam. Apical cell slightly thickened up to 4.0 μm, dark green. Asci Lecidella-type, slightly clavate with dark amyloid tholus, eight unicellular spores, 40–45 μm. Ascospores simple, hyaline, broadly ellipsoid, I−, (8–)10–11(–12) × (4–)5–6(–7) μm, l/w: 10.6/5.7 μm.
Conidiomata not observed.
Chemistry
Atranorin and thuringione by TLC. Spot test: K+ light yellow, C−, KC−.
Etymology
The term drakensis was chosen because the collected specimens occurred north (Chile, Región de Magallanes y de la Antártica Chilena) and south (maritime Antarctica) of the Drake Passage.
Distribution and habitat
Two specimens of Lecidella drakensis were found in the subantarctic areas of southern Patagonia (sSA, Morro Chico and Torres del Paine National Park) solely on siliceous rock and the other three in maritime Antarctica (James Ross Island) on siliceous rock and once on mosses. This species is currently recorded only from these areas.
Notes
The collections of L. drakensis form a distinct and highly supported clade containing accessions from southern Patagonia and maritime Antarctica, which is placed in the L. elaeochroma clade (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015). The species is clearly distinguished morphologically by the following characteristics: very small apothecia in contrast to L. elaeochroma (Wirth et al. Reference Wirth, Hauck and Schultz2013); lacking soralia and a different chemistry to L. flavosorediata (Wirth et al. Reference Wirth, Hauck and Schultz2013); more oblong spores and a different chemistry to L. wulfenii (Wirth et al. Reference Wirth, Hauck and Schultz2013); a dark green instead of violet-brown epihymenium and a different chemistry to L. euphorea (Zhao et al. Reference Zhao, Zhang, Zhao, Wang, Leavitt and Lumbsch2015); a thinner thallus and also a different chemistry to L. meiococca (Knoph & Leuckert Reference Knoph and Leuckert1994). Neither species described here is related to the Southern Hemispheric, not yet molecularly confirmed species L. sublapicida (C. Knight) Hertel. This species is distinguished by its different chemistry (arthothelin, isoarthothelin) and reddish brown hymenium (Knoph & Leuckert Reference Knoph and Leuckert1994). Kappen (Reference Kappen1985) mentioned a ‘L. antarctica’, which has not been formally described, that forms a greyish pulvinate crust on rock, from northern Victoria Land (H. Hertel, personal communication) and is clearly morphologically different from L. ayazii and L. drakensis.
Additional specimens examined
Antarctica: Antarctic Peninsula: James Ross Island, 142 m a.s.l., 63°48′24.9″S, 57°50′27.6″W, M. G. Halıcı JR 0.082 (OQ534854; ERCH); ibid., 2 m a.s.l., 63°52′39.0S″, 57°46′51.6″W, M. G. Halıcı JR 0.115 (OQ534857; ERCH); ibid., 292 m a.s.l., 63°49′46.2″S, 57°54′21.6″W, M. G. Halıcı JR 0.397 (OQ534856; ERCH) (Table 1).—Chile: Región de Magallanes y de la Antártica Chilena: Laguna Blanca, 440 m a.s.l., 52°3′31″S, 71°25′11″W, U. Ruprecht UR00086 (MK620140; SZU).
Discussion
Various lichen species are exclusively found in the Southern Hemisphere. They are often part of globally distributed genera but form well-distinguished groups located only in the southern polar areas. Two prominent examples are the common species Lecidea cancriformis C. W. Dodge & G. E. Baker and Usnea aurantiacoatra (Jacq.) Bory (Ruprecht et al. Reference Ruprecht, Fernández-Mendoza, Türk and Fryday2020; Lagostina et al. Reference Lagostina, Andreev, Dal Grande, Grewe, Lorenz, Lumbsch, Rozzi, Ruprecht, Sancho and Søchting2021). Both species are distributed not only in the subantarctic areas of southern South America but also in the maritime and/or continental Antarctic. Other species such as Usnea antarctcia Du Rietz are even more restricted to southern polar areas and occur only in the maritime Antarctic (Lagostina et al. Reference Lagostina, Andreev, Dal Grande, Grewe, Lorenz, Lumbsch, Rozzi, Ruprecht, Sancho and Søchting2021). Conversely, species such as Lecidea polypycnidophora (Hale et al. Reference Hale, Fisher, Keuler, Smith and Leavitt2019) and Lecidea andersonii (Hertel Reference Hertel2007; Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2010; Hale et al. Reference Hale, Fisher, Keuler, Smith and Leavitt2019) have an alpine and/or bipolar distribution which suggests that there are migration routes especially along the American continent (Garrido-Benavent & Pérez-Ortega Reference Garrido-Benavent and Pérez-Ortega2017; Hale et al. Reference Hale, Fisher, Keuler, Smith and Leavitt2019). The two newly described species, L. ayazii and L. drakensis, are part of the cosmopolitan genus Lecidella. They are both located in the L. elaeochroma clade, to which mostly Northern Hemisphere species have been assigned until now, and form clearly distinguished groups.
Lecidella ayazii, which was found by the last author only in the maritime Antarctic, was also found to cluster with a collection from alpine areas of the La Sal Mountains, Utah, USA (Leavitt et al. Reference Leavitt, Hollinger, Summerhays, Munger, Allen and Smith2021) and the Kivalliq Region, Nunavut, in the north of Canada (this study). The specimen from the La Sal Mountains was determined as Lecidella wulfenii because of its growth on moss, which is one of the most important traits for assigning this species. Unfortunately, there is another published sequence of L. wulfenii determined by Roman Türk (Ruprecht et al. Reference Ruprecht, Lumbsch, Brunauer, Green and Türk2012) which is not related and which is located in another clade (Fig. 2). This specimen was found on moss over calcareous rock, which is another important distinguishing characteristic (Wirth et al. Reference Wirth, Hauck and Schultz2013), in contrast to the specimens from the La Sal Mountains (Utah, USA) and the Kivalliq Region (Nunavut, northern Canada), where it can be assumed that the substratum is siliceous. The specimen of L. ayazii from the maritime Antarctic was also growing only over siliceous substrata. Lecidella wulfenii was described as Lichen muscorum by Wulfen (Jacquin Reference Jacquin1790), but because that name is illegitimate, Hepp (Reference Hepp1853) introduced the replacement name Biatora wulfenii Hepp, which was transferred to Lecidella by Körber (Reference Körber1861). Since Wulfen's species was described from Austria and the Türk collection mentioned above is also from Austria, we have no doubt that the epithet ‘wulfenii’ should be applied to the European species.
Thus far, Lecidella drakensis shows a similar pattern to that of Usnea aurantiacoatra (Lagostina et al. Reference Lagostina, Andreev, Dal Grande, Grewe, Lorenz, Lumbsch, Rozzi, Ruprecht, Sancho and Søchting2021). It occurs in southern South America as well as in maritime Antarctica. However, five other accessions were also recorded from the maritime Antarctic belonging to the cosmopolitan L. stigmatea clade, including one assigned to the bipolar species L. siplei.
Acknowledgements
We would like to give special thanks to Alan Fryday (Michigan State University) for his valuable advice and help as a reviewer, and for providing the two specimens from northern Canada. We are also grateful for the support of Sabine Agatha with microscopy, Anna Götz with the sequence analyses and Roman Türk with the morphological analyses (Paris Lodron University Salzburg). We would also like to thank Erciyes University for the financial support to carry out fieldwork on James Ross Island, Antarctica, ITU Polrec for its support and the J. G. Mendel Station for the use of infrastructure and facilities.
This study was conducted under the auspices of the Presidency of the Republic of Turkey, supported by the Ministry of Industry and Technology and coordinated by the TUBITAK MAM Polar Research Institute; it was funded by the grants TUBITAK 118Z587, 121Z771 and by the Turkish Academy of Sciences, as well as in whole or in part by the Austrian Sciences Fund (FWF) 10.55776/P26638 and 10.55776/P35512. For open access purposes, the author has applied a CC BY public copyright license to any author accepted manuscript version arising from this submission.
Author ORCIDs
Ulrike Ruprecht, 0000-0002-0898-7677; Feyza Nur Avci, 0000-0002-2010-3643; Mehmet Candan, 0000-0001-6496-4771; Mehmet Gökhan Halıcı, 0000-0003-4797-1157.
Competing Interests
The authors declare none.
Open access
