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Peltigera Hydrophila (Lecanoromycetes, Ascomycota), a New Semi-Aquatic Cyanolichen Species from Chile

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Peltigera Hydrophila (Lecanoromycetes, Ascomycota), a New Semi-Aquatic Cyanolichen Species from Chile Plant and Fungal Systematics 65(1): 210–218, 2020 ISSN 2544-7459 (print) DOI: https://doi.org/10.35535/pfsyst-2020-0016 ISSN 2657-5000 (online) Peltigera hydrophila (Lecanoromycetes, Ascomycota), a new semi-aquatic cyanolichen species from Chile Jolanta Miadlikowska1*, Nicolas Magain1,2, William R. Buck3, Reinaldo Vargas Castillo4, G. Thomas Barlow1, Carlos J. Pardo-De la Hoz1, Scott LaGreca1 & François Lutzoni1 Abstract. Peltigera hydrophila, a new species from Chile tentatively distinguished based Article info on phylogenetic evidence but not yet named, is formally described here. Morphological Received: 29 Jan. 2020 differences (e.g., non-tomentose thallus) and habitat preferences (semi-aquatic) corroborate Revision received: 23 Mar. 2020 molecular and phylogenetic distinctiveness of this early diverging lineage in section Pelti- Accepted: 24 Mar. 2020 gera. Due to overlapping ecological ranges, P. hydrophila shares some morphological traits Published: 2 Jun. 2020 with aquatic species from the phylogenetically unrelated section Hydrothyriae. Associate Editor Key words: cyanolichen, cyanobiont, Nostoc, mycobiont, symbiosis, taxonomy Damien Ertz Introduction The most recent multi-locus phylogenetic revision of sec- (clades 2a and 2b, respectively) within section Pelti- tions Peltigera and Retifoveatae of the genus Peltigera gera (Fig. 1B). Both species seem to be morphologically (Fig. 1) suggested the presence of 88 species, of which 50 distinct and geographically restricted, however multiple were new to science (Magain et al. 2018). Forty-nine of collections from different localities are available only the newly delimited species are part of section Peltigera, for P. sp. 14. Together with its sister species P. auber- which includes species with tomentose thalli (with some tii, known from the Holantarctic Kingdom (Martínez exceptions, e.g., P. frigida and P. degenii) and lacking et al. 2003), P. sp. 14, which is restricted to Chile, rep- secondary metabolites detectable by thin layer chroma- resents the first divergence event within sectionPeltigera tography (Miadlikowska & Lutzoni 2000; Magain et al. ( Fig. 1B). This species was recognized as potentially new 2018). Most of these newly delimited putative species are to science by Miadlikowska et al. (2014; corresponding restricted to a single biogeographic region, and hence few to ‘Peltigera sp. nov.’) in the phylogenetic context of the are widespread. In section Peltigera, specificity ofPeltigera Lecanoromycetes. Peltigera sp. 14 is one of the rare cases species (mycobionts) in their association with Nostoc in the genus Peltigera where both symbionts are spe- phylogroups (cyanobionts) ranges from strict specialists cialists and associate almost exclusively with each other (associating with only one Nostoc phylogroup) to broad (Magain et al. 2017a, b, 2018; Miadlikowska et al. 2018; generalists (up to eight Nostoc phylogroups), with wide- Pardo-De la Hoz et al. 2018; Chagnon et al. 2019). The spread Peltigera species recruiting a broader selection of mycobiont forms thalli with Nostoc phylogroup XXIII, Nostoc phylogroups than species with limited distributions which was also found in a single specimen of its closest (Magain et al. 2018). One potentially new species, P. sp. 13, relative, P. aubertii, a species also found in Chile that was found to belong to section Retifoveatae (Fig. 1B). otherwise associates with phylogroup XXII (Magain et al. Two species – Peltigera sp. 14 and P. sp. 16 – were 2018). Peltigera sp. 14 is the second water-associated found to be part of the two earliest diverged lineages lineage in the genus Peltigera. However, this species has a broader ecological amplitude than the distantly related North American species of section Hydrothyriae 1 Department of Biology, Duke University, Box 90338, Durham, NC (Fig. 1A), as it can also grow in moist terrestrial habitats 27708, USA (e.g., on mosses subjected to waterfall splashes) com- 2 Evolution and Conservation Biology, InBios Research Center, Institut de Botanique B22, Université de Liège, Chemin de la vallée 4, 4000 pared to the strictly aquatic P. aquatica, P. hydrothyria, Liège, Belgium and P. gowardii of section Hydrothyriae. Here, we for- 3 Institute of Systematic Botany, New York Botanical Garden, 2900 mally describe P. hydrophila to accommodate Peltigera Southern Boulevard, Bronx, NY 10458-5126, USA sp. 14, the non-tomentose, semi-aquatic new species in 4 Departamento de Biología, Universidad Metropolitana de Ciencias de la Educación, Avda. José Pedro Alessandri 774, Ñuñoa, Santiago, Chile section Peltigera. * Corresponding author e-mail: [email protected] © 2020 W. Szafer Institute of Botany, Polish Academy of Sciences. This is an Open Access article distributed under the terms of the Creative Commons Attribution License CC BY 4.0 (http://creativecommons.org/licenses/by/4.0/) J. Miadlikowska et al.: Peltigera hydrophila, a new semi-aquatic species from Chile 211 100 B PELTIDEA A OUTGROUP 100 POLYDACTYLON 100 Hydrothyriae* PeltigeraÊsp.Ê13 CladeÊ1 100 Phlebia RETIFOVEATAE 100 PeltigeraÊretifoveata 66 95 100 Chloropeltigera P2039 58 100 PeltigeraÊhydrophila 100 P2062 sp.Ênov.Ê 65 Peltidea CladeÊ2a 100 (PeltigeraÊsp.Ê14) P1534 99 Polydactylon 100 PeltigeraÊaubertii 100 Horizontales 100 PeltigeraÊsp.Ê16 PELTIGERA CladeÊ2b 100 100 100 100 Retifoveatae PeltigeraÊfrigida/patagonica 82 100 CladeÊ2c 99 PeltigeraÊkristinssonii Peltigera* 100 CladeÊ3 0.08 100 PeltigeraÊcontinentalis/isidiophora 94 CladeÊ8 PeltigeraÊdegeniiÊgroup 100 100 100 CladeÊ9 PeltigeraÊcaninaÊgroup 100 95 CladeÊ7 PeltigeraÊcinnamomeaÊgroup 100 76 CladeÊ4 100 PeltigeraÊdidactylaÊgroup 100 CladeÊ6 PeltigeraÊrufescensÊgroup 100 88 CladeÊ5 100 PeltigeraÊmonticola/ponojensisÊgroup 0.02 Figure 1. Phylogenetic placement of section Peltigera and the aquatic to semi-aquatic lineages (indicated by asterisks for section Hydrothyriae and P. hydrophila sp. nov. within section Peltigera) in the context of the genus Peltigera (A) and phylogenetic relationship of P. hydrophila sp. nov. within section Peltigera (B). Maximum Likelihood (ML) tree for the genus Peltigera (based on seven loci: ITS, nrLSU, β‐tubulin, RPB1, COR1b, COR3, COR16; panel A) is adapted from Figure 2 of Chagnon et al. (2019). Intrageneric classification (i.e., recognized sections) follows Miadlikowska & Lutzoni (2000). Numbers associated with internodes represent ML bootstrap support values (based on 1000 pseudoreplicates). The ML tree for sections Retifoveatae and Peltigera (based on five loci: ITS, β‐tubulin, COR1b, COR3, COR16; panel B) is adapted from Figure 1 of Magain et al. (2018). Clades were collapsed using FigTree v1.4.3 (Rambaut 2012). Clades containing more than two species are indicated as groups. The scales represent nucleotide substitutions per site. Materials and methods & Dudik 2008; Fick & Hijmans 2017). We chose to use MaxEnt because it has been shown to work well for modeling species Specimens were examined under a Leica MZ6 dissecting mi- with very few known records (Elith et al. 2006; Pearson et al. croscope. A total of 20 ascospores from three apothecia were 2006; de Siqueira et al. 2009). Locality data were thinned with measured using a Leica DMLB compound microscope. For the a 5 km radius rule to minimize spatial autocorrelation using the R descriptions and use of terminology we followed Vitikainen package spThin (Aiello-Lammens et al. 2015), and background (1994: 5–17). Thin layer chromatography (TLC) was performed extent was limited by a point buffer of 15 degrees. Model tuning on four specimens, using solvents A, B’ and C (Culberson & Am- included a variety of models reconstructed with different feature mann 1979; Culberson & Johnson 1982; solvent C is the same classes [i.e., linear (L), linear – quadratic (LQ), linear – quad- as solvent ‘TA’ of Holtan-Hartwig 1993). GenBank accession ratic – hinge (LQH), and hinge (H)] and different regularization numbers for sequences from specimens P2039, P2062 and P1534 multipliers (1, 2, 3, and 4) implemented in dismo (Hijmans are provided in Magain et al. (2018), whereas the accession et al. 2017) and ENMeval packages (Muscarella et al. 2014). numbers for the remaining specimens are part of the voucher Preparation runs were performed using the package Wallace information provided below. version 1.0.6.2 to optimize the run settings (Kass et al. 2018). To determine the potential distribution of P. hydrophila, The area under the ROC curve (AUC) for each model created we used ecological niche modelling (ENM) techniques for the was calculated to estimate the credibility of the analysis. AUC estimation of the habitat suitability for this species. The ENM values were calculated automatically and the higher AUC was was conducted using the maximum entropy method implemented selected as a reliable indicator of performance (Phillips et al. in MaxEnt version 3.4.1 (Phillips et al. 2017, 2020) based on all 2009; Peterson et al. 2011). Raster files were prepared in R known localities for P. hydrophila and all nineteen bioclimatic (R Core Team 2020) and all maps were edited in QGIS version variables available in Worldclim 2.1 at 30 arc seconds (Phillips 3.12.0 (QGIS Development Team 2020). 212 Plant and Fungal Systematics 65(1): 210–218, 2020 Taxonomy Peltigera hydrophila W. R. Buck, Miadl. & Magain, sp. nov. (Figs 2–4) MycoBank MB 834180 Diagnosis: Unique in its glabrous thallus upper surface, which becomes deep bluish-violet when wet; possessing somewhat phyllidiate and indented thallus margins; apothecia round, flat to convex,
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