Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tnzb20 Download by: [Mr Peter De Lange] Date: 23 August 2017, At: 18:31 New Zealand Journal of Botany ISSN: 0028-825X (Print) 1175-8643 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzb20 Lecanora kohu, a new species of Lecanora (lichenised Ascomycota: Lecanoraceae) from the Chatham Islands, New Zealand C. Printzen, D. J. Blanchon, A. M. Fryday, P. J. de Lange, D. M. Houston & J. R. Rolfe To cite this article: C. Printzen, D. J. Blanchon, A. M. Fryday, P. J. de Lange, D. M. Houston & J. R. Rolfe (2017): Lecanora kohu, a new species of Lecanora (lichenised Ascomycota: Lecanoraceae) from the Chatham Islands, New Zealand, New Zealand Journal of Botany, DOI: 10.1080/0028825X.2017.1364274 To link to this article: http://dx.doi.org/10.1080/0028825X.2017.1364274 Published online: 23 Aug 2017. Submit your article to this journal View related articles View Crossmark data
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Full Terms & Conditions of access and use can be found athttp://www.tandfonline.com/action/journalInformation?journalCode=tnzb20
Download by: [Mr Peter De Lange] Date: 23 August 2017, At: 18:31
Lecanora kohu, a new species of Lecanora(lichenised Ascomycota: Lecanoraceae) from theChatham Islands, New Zealand
C. Printzen, D. J. Blanchon, A. M. Fryday, P. J. de Lange, D. M. Houston & J. R.Rolfe
To cite this article: C. Printzen, D. J. Blanchon, A. M. Fryday, P. J. de Lange, D. M. Houston& J. R. Rolfe (2017): Lecanora kohu, a new species of Lecanora (lichenised Ascomycota:Lecanoraceae) from the Chatham Islands, New Zealand, New Zealand Journal of Botany, DOI:10.1080/0028825X.2017.1364274
To link to this article: http://dx.doi.org/10.1080/0028825X.2017.1364274
Lecanora kohu, a new species of Lecanora (lichenisedAscomycota: Lecanoraceae) from the Chatham Islands,New ZealandC. Printzena, D. J. Blanchonb, A. M. Frydayc, P. J. de Langed*, D. M. Houstond andJ. R. Rolfee
aSenckenberg Forschungsinstitut und Naturmuseum, Frankfurt, Germany; bDepartment of Natural Sciences,Unitec Institute of Technology, Biodiversity and Animal Welfare Research Group, Auckland, New Zealand;cHerbarium, Department of Plant Biology, Michigan State University, East Lansing, USA; dScience & PolicyGroup, Department of Conservation, Terrestrial Ecosystems Unit, Newton, New Zealand; eScience & PolicyGroup, Department of Conservation, Terrestrial Ecosystems Unit, National Office, Wellington, New Zealand
ABSTRACTLecanora kohu Printzen, Blanchon, Fryday et de Lange is describedas new to science from Rangatira (South East Island), ChathamIslands. It is morphologically similar to L. symmicta (Ach.) Ach.,from which it is distinguished by the continuous, areolate thallus,immersed apothecia with pale pink to pink-brown discs, and bythe presence of atranorin and psoromic acid rather than usnicacid, zeorin and xanthones in the thallus. The new species is so farknown only from Rangatira (South East Island), the southernmostof the three main islands of the Chatham Islands group where itwas collected twice in 2015 on the bark of Melicytus chathamicus(Violaceae) and on the bark of an undescribed species ofMuehlenbeckia (Polygonaceae), M. aff. australis. Using the NewZealand Threat Classification System, the new species has beenassessed as ‘Data Deficient’.
ARTICLE HISTORYReceived 27 June 2017Accepted 2 August 2017First published online 24August 2017
KEYWORDSChatham Islands; Lecanoraconfusa; Lecanora symmicta;Lecanora varia group; lichentaxonomy; New Zealandmycobiota
ASSOCIATE EDITORDr Leon Perrie
Introduction
Lecanora is a species-rich cosmopolitan genus of an estimated 600 species (Lumbsch & Elix2004; Ryan et al. 2004; Galloway 2007). Despite recent adjustments to the genus and thetransfer to other genera (e.g. Zhao et al. 2016) of a number of taxa previously assigned toit, the current circumscription still requires further adjustment (Printzen 2001; Lumbsch& Elix 2004; Galloway 2007). In the most recent treatment of the New Zealand species, Gal-loway (2007) accepted 47 taxa; even so, his comment ‘many nomenclatural problems in thisgenus await resolution, and in New Zealand many taxa cannot at present be named withcertainty. Much collecting and study of local populations is needed before Lecanora inNew Zealand is known at all adequately’ (Galloway 1985, p. 120) still applies.
Whilst Lecanora in New Zealand still needs critical assessment, the task of collecting thegenus from the less accessible parts of that country, as advocated by Galloway (1985), has
CONTACT P. J. de Lange [email protected]*Present addres: Department of Natural Sciences, Unitec Institute of Technology, Biodiversity and Animal Welfare ResearchGroup, Auckland, New Zealand
NEW ZEALAND JOURNAL OF BOTANY, 2017https://doi.org/10.1080/0028825X.2017.1364274
continued. Here we describe a new species of Lecanora collected from Rangatira (SouthEast Island), the southernmost of the main islands of the Chatham Islands group(Figure 1). The species was recognised from two collections made from that island atWhalers Bay and Western Landing in July 2015.
Materials and methods
Thallus and apothecial characters were examined using light microscopy on hand cut sec-tions mounted in water, Lugol’s iodine solution or lactophenol cotton blue. Spore
Figure 1. Chatham Islands group showing their relationship to New Zealand, the main islands of thegroup and the type locality of Lecanora kohu, Rangatira (South East Island) (in bold).
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measurements were made in Lugol’s solution and measurements of the apothecial charac-ters follow the methods of Printzen (2001) and Pérez-Ortega et al. (2010). Thin layerchromatography [TLC] was carried out using the methods of Culberson (1972) andWhite & James (1985) using solvents B and C.
DNA extraction and polymerase chain reaction (PCR) amplification were carried outusing the methods listed by Hayward et al (2014). Fungal ITS rDNA was amplifiedusing the primers ITS1F (Gardes & Bruns 1993) and ITS4 (White et al. 1990); nuLSUrDNA was amplified using the primers LR0R and LR5 (Vilgalys & Hester 1990);mtSSU rDNA was amplified using the primers mrSSU1 and mrSSU3R (Zoller et al.1999). DNA sequences are available on GenBank (accessions MF115999, MF116000,MF116001 and MF373839; see Table 1).
Single gene datasets containing the sequences listed in Table 1 were compiled and alignedusing the MAFFT algorithm (Katoh et al. 2005) as implemented on the GUIDANCE webserver (Penn et al. 2010a).GUIDANCEv2was also used to remove regions of uncertain align-ment (i.e. positions with less than 0.93 GUIDANCE score; Penn et al. 2010b). In addition,terminal residuals present in less than 66% of the Operational Taxonomic Units wereremoved from all single gene alignments, as well as a 498 bp insertion in the nuLSU sequenceof Lecanora achroa. The final alignments had the following dimensions: 92 sequences, 234 bpfor ITS; 91 sequences, 580 bp for mtSSU; and 56 sequences, 711 bp for nuLSU.
Datasets were concatenated to yield a final alignment of 98 sequences and 1525 bplength. A heuristic search for the maximum likelihood (ML) bootstrap tree with simul-taneous inference of the optimal partitioning scheme and substitution models for eachdata partition was performed using the online version of IQ-TREE (Nguyen et al. 2015;Chernomor et al. 2016; Kalyaanamoorthy et al. 2017) suggesting five initial partitions(ITS1, 5.8S rDNA, ITS2, mtSSU, nuLSU). Branch lengths were assumed to be equal forall partitions. Branch support was estimated with the ultrafast bootstrap algorithm(Minh et al. 2013) based on 1000 bootstrap replicates and using a maximum of 1000 iter-ations and a minimum correlation coefficient of 0.99 as a stopping rule. We used theMarkov Chain Monte Carlo approach implemented in MrBayes v3.2.6 (Ronquist et al.2012) to infer phylogenetic trees applying the partitioning scheme inferred with IQ-TREE and slightly simplified substitution models, because most of the models inferredby IQ-TREE are not implemented in MrBayes. See Table 2 for details on gene partitionsand substitution models. As mean of the exponentially distributed branch length prior weused the mean branch length of the ML tree (1/42.14). All parameters of the substitutionmodels were unlinked between data partitions. MrBayes was set to sample every 200th treefrom two independent runs, each with four chains that were incrementally ‘heated’ by0.15. The average standard deviation of bipartition frequencies among runs was calculatedevery 10000 generations to infer convergence of the Markov Chains, discarding the first50% of the sampled trees as burn-in and including only those bipartitions with a frequencyof at least 10%. The analysis was stopped after 26.2M generations when the standard devi-ation had dropped below 0.01.
Results
Maximum likelihood and Bayesian analysis yielded very similar phylogenetic trees. Resultsof both analyses are summarised in Figure 2, with bootstrap support values and posterior
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Table 1. Taxa and GenBank accession numbers of sequences used for phylogenetic inference.Sequences newly submitted to GenBank in bold.
branch probabilities projected on the ML tree. The phylogeny shows Lecanoraceae as awell-supported clade separate from Parmeliaceae. Within Lecanoraceae, few brancheswere supported in both analyses. Apart from Frutidella, Japewia and Pyrrhospora,which were each represented by a single species, only Protoparmeliopsis, Myriolecis, Leci-della and Palicella were reconstructed as monophyletic genera, the last two with poorsupport. Rhizoplaca chrysoleuca appears as sister to a well-supported clade comprisingProtoparmeliopsis and Myriolecis. On the ML tree Miriquidica leucophaea appearswithin a poorly supported Ramboldia, while on the Bayesian consensus tree it occupied
Table 2. Summary of data partitions and substitution models used for phylogenetic inference.ITS1 5.8S rRNA ITS2 mtSSU nuLSU
No. taxa 92 92 92 91 56Position 1–30 31–186 187–234 235–814 814–1525Substitution model (IQ-TREE) TIM2 + G4 TNe + I + G4 TIMe + G4 HKY + I + G4 TN + I + G4Substitution model (MrBayes) HKY + G4 K2P + I + G4 K2P + G4 HKY + I + G4 HKY + I + G4
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a position near Lecanora symmicta and L. flavopallida. The two collections of the unde-scribed Lecanora are not assigned to any of the segregate genera, but instead appear inan unsupported clade (BS 61%, PP 0.42) together with L. austrotropica, L. leproplaca,
Figure 2. Maximum likelihood phylogenetic tree of Lecanora and related genera and species withinLecanoraceae. Six species of Parmeliaceae were used as outgroup. Bold branches received ML boot-strap values ≥ 90% and Bayesian posterior probabilities ≥ 0.95. Support values for the other branchesare not displayed. Lecanora kohu in bold.
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L. subimmergens and L. toroyensis. These species are typical members of Lecanora s. str.(i.e. the so-called ‘L. subfusca group’) and bear no close morphological resemblance tothe new species. Further taxa attached to this clade (also with poor support) includeL. symmicta and Pyrrhospora quernea. Hence, only the assignment of the new speciesto Lecanoraceae is at present well-supported, while its correct position within thatfamily remains elusive. We therefore describe it as a species of Lecanora, although mor-phologically it differs strongly from typical members of Lecanora s. str., the new speciesresembles L. symmicta but is easily separated from it by its continuous, areolate thallus,immersed apothecia with pale pink to pink-brown discs, and by the presence of atranorinand psoromic acid rather than usnic acid, zeorin and xanthones in the thallus.
Discussion
Phylogenetic analyses based on ITS, nuLSU and mtSSU sequences did not convincinglysupport a specific phylogenetic position within Lecanoraceae for the new species. Themost similar sequences seemed to be those of Lecanora carpinea (L.) Vain., but a relation-ship between the two species was not supported in the phylogenetic analyses. Macroscopi-cally, the new species seems to belong to what was termed the ‘Lecanora varia group’ byGalloway (2007), for which he recognised three species from New Zealand: L. conizaeoidesNyl. ex Cromb., L. expallens Ach., and L. symmicta. However, molecular phylogeneticwork has shown that these species do not form a monophyletic group (Arup & Grube1998; Pérez-Ortega et al. 2010). Based on apothecial anatomy the new species has
Figure 3. Lecanora kohu holotype (de Lange CH2831, UNITEC 7497) on Muehlenbeckia aff. australis. A,Lecanora kohu (central thallus) on bark of Muehlenbeckia aff. australis. Scale bar = 5 mm; B, Lecanorakohu in dry state. Scale bar = 1 mm; C, Lecanora kohu in wet state. Scale bar = 1 mm.
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similarities with L. symmicta, although chemistry and other anatomical features (seeabove) readily distinguish the two. The gross morphology of the new species (Figure 3)suggested a closer relationship with L. confusa Almb., which on critical examinationwas also discounted (see Recognition below). Lecanora is one of the last remainingform genera of lichenised ascomycetes, and well known to be polyphyletic. Until suchtime as the wider generic issues of Lecanora are resolved, placement of the new specieswithin that genus seems for now a suitably cautious measure.
Taxonomy
Lecanora kohu Printzen, Blanchon, Fryday et de Lange, sp. nov.
Mycobank accession number: 822008
Holotype (Figure 3). NEW ZEALAND, Chatham Islands group, Rangatira (South EastIsland), Western Landing, P.J. de Lange CH2831, 28 July 2015 (UNITEC 7497, Isotype,AK, MSC)
Diagnosis. Lecanorae symmictae similis, thallo in vivo continuo areolato, chloro-albido, insicco cremeo vel pallide flavido, apotheciis immersis discis pallide roseis vel roseo-fuscisinstructis, ascosporis latioribus (10.0–12.5 × 5.0–6.5 µm), et thallo acido psoromicoinstructo differt.
Similar to Lecanora symmicta from which it is distinguished by the continuous, areo-late, greenish white (when fresh) thallus, drying cream to pale yellow; by the immersedapothecia with pale pink to pink-brown discs; the wider ascospores (9.0–12.5 × 5–7 µm); and by the presence of atranorin and psoromic acid in the thallus.
Description (Figures 3–5). Thallus crustose, areolate-cracked, greenish-white when fresh,drying to cream to pale-yellow, soredia and isidia absent. Areoles rounded to oblong with
Figure 4. Asci of Lecanora kohu from holotype (P.J. de Lange CH2831, UNITEC 7497) stained with iodine(Lugol’s iodine solution). Scale bar = 20 µm.
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irregular-crenate margins, 0.25–0.5 mm in length or diameter, plane to slightly concave,cream to yellow-grey in colour, smooth to slightly roughened between areoles. Photobionttrebouxioid, individual cells 5–10 µm in diameter. Apothecia 0.25–0.4 mm in diameter,immersed, flat to slightly concave, single to grouped, 1–3(–5) per areole, but severalareoles merge and apothecia become clustered, disc pale pink to pink-brown, matt, eprui-nose. Thalline margin generally not apparent, but discs are generally embedded in thallinetissue. Parathecium white to cream, scarcely evident in young apothecia, level with discand inconspicuous in older apothecia, in section 22–62.5 µm wide and 30–65 µm tall, con-sisting of gelatinised hyphae, colourless to pale straw-coloured, IKI–, HNO3−, POL+ crys-tals common, algae occasionally present at base. Hypothecium 37.5–87.5 µm tall,colourless or pale-straw-coloured, small oil droplets sometimes present. Hymenium 35–60 µm tall, colourless, IKI+ bright blue, HNO3−; epihymenium brown, granular, POL+,5–10(–25) µm thick; paraphyses colourless, simple or occasionally branched, withlumina 0.7–1.0 µm wide, apically not thickened; asci Lecanora-type, 32.5–45 × 9–15 µm(Figure 4); ascospores 8 per ascus, colourless, simple, ellipsoid, 9–12.5 × 5–7 µm; pycnidianot seen.
Additional specimens examined. Chatham Islands, Rangatira (South East Island),Whalers Bay Track, P.J. de Lange CH2832 & D.M. Houston, 27 Jul 2015 (UNITEC7499, Duplicates: AK, MSC).
Distribution. Chatham Islands, Rangatira (South East Island) (Figure 1). Although cur-rently known only from the Chatham Islands, it is likely that dedicated searching will
Figure 5. Cross section of apothecium of Lecanora kohu from holotype stained with lactophenol cottonblue (P.J. de Lange CH2831, UNITEC 7497) showing lack of algae in apothecial margins. Scale bar =50 μm.
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find Lecanora kohu elsewhere. This was the case with Caloplaca maculata D.J.Galloway,which when described (Galloway 2004) was known only from the Chatham Islands butwas subsequently discovered in the South Island of New Zealand (de Lange 2012).
Habitat. On Rangatira (South East Island) Lecanora kohu has been collected growing onthe exposed, mature branches of a widespread, common unnamed species of Muehlen-beckia (M. aff. australis) and from the exposed trunks of Chatham Island māhoe (Melicy-tus chathamicus). Associated lichens were typically sparse: on Muehlenbeckia, onlyOpegrapha agelaeoides was noted whilst on Melicytus, Lecanora kohu grew in associationwith Bacidia laurocerasi, Phlyctis sordida and P. uncinata.
Recognition. The presence of a conspicuous parathecium resembling a truly biatorineapothecial margin (Figure 5) and lacking algae suggests a similarity with L. symmicta(Printzen 2001), but chemistry, thallus structure and spore size differentiate the twospecies (see Results). Lecanora densa (Śliwa et Wetmore) Printzen contains psoromicacid in addition to usnic acid, and also differs in forming an areolate thallus, a corticateamphithecium with an algal layer and shorter ascospores of 8–9.5 µm on average. Leca-nora confusa Almb. can have immersed apothecia, but these have granular margins(Almborn 1955) and a different chemistry. Its separation from other members of theL. varia group (sensu Galloway 2007) are given in the identification key. Lecanora flavo-pallida is another similar species in the New Zealand biota but was not included in theL. varia group by Galloway (2007) because it lacks usnic acid. It is distinguished fromour new species by the presence of xanthones in the thallus that result in a C+ orange reac-tion and considerably larger ascospores of 14–19 × 9–12 µm (Guderley et al. 1998). TheTasmanian endemic species L. coppinsiarum is also remotely similar but the thallus ofthat species contains only atranorin and its apothecia are sessile and coarsely pruinose(Kantvilas 2012). A superficially similar, apparently undescribed Lecanora species withinnate apothecia is also known from the Falkland Islands, but that taxon has a granularthallus, an apothecial margin and a thallus containing usnic acid and zeorin. It is probablymore closely related to L. confusa. Species from the recently described genus Palicella(Rodriguez-Flakus & Printzen 2014) have apothecia with a distinct proper margin thatare often darkened by cinereorufa-green (Meyer & Printzen 2000), and have asci resem-bling the Lecidella-type.
Conservation status. Lecanora kohu is assessed as ‘Data Deficient’ using the New ZealandThreat Classification System (Townsend et al. 2008). Lecanora kohu is known from onlytwo localities on Rangatira (South East Island), Chatham Islands. Examination of themany collections of corticolous lichens from the Chatham Islands group held in AK,CHR and UNITEC did not reveal any further specimens of L. kohu. Nevertheless, wesuspect that it is more likely that L. kohu is overlooked than genuinely threatened, andthat its absence from other islands in the Chatham Islands group reflects that they haveyet to be collected by expert lichenologists.
Etymology. The epithet ‘kohu’ derives from Te Reo Māori for ‘mist’ (K. A. Raharaha,Ngati Kuri, pers. comm., 22 March 2017). The name alludes to the sea fog that oftenshrouds the location in which this species was found, Rangatira (South East Island), thesouthern-most of the main islands of the Chatham Islands group.
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Key to Lecanora kohu and morphologically similar New Zealand species of Lecanora(Lecanora varia group sensu Galloway 2007)
2. Thallus effuse, granular-scurfy, pale yellow-green when fresh, white when dry; apothe-cia when present crowded, disc yellow to yellow-fawn; Pd−, usnic acid, zeorin and thio-phanic acid present............................................................................................Lecanora expallensThallus coarsely granular, olive-green togrey-greenwhen fresh, greywhendry, apothecia sparseto absent, when present often obscured by mat of soredia, disc pale grey-green to grey-brown;Pd+ red, fumarprotocetraric acid present………………………… Lecanora conizaeoides3. Thallus effuse, uneven, or lacking, subgranular, cream-yellow when fresh, yellowishwhen dry; apothecia sessile, disc pale yellow to pale brown; Pd−, usnic acid and zeorinpresent, psoromic acid absent........................................................................Lecanora symmictaThallus continuous, areolate, greenish white when fresh, drying cream to pale yellow;apothecia immersed, disc pale pink to pink-brown; Pd+ orange, usnic acid and zeorinabsent, psoromic acid present................................................................................ Lecanora kohu
Acknowledgements
The authors would like to thank the Chatham Islands Area Staff, especially Connie Norgate (AreaManager 2014–2016), Amanda Baird, Bex Bell and James Launder for assistance in the field andtheir ongoing interest in the flora and mycobiota of Rangatira. We thank Ines Schönberger(Allan Herbarium, Landcare Research) for assistance with lichen literature. We would like toacknowledge Kamera Raharaha for suggesting the species epithet ‘kohu’. We thank NevilleWalsh, Royal Botanic Gardens, Melbourne, Victoria, Australia for providing the Latin diagnosis.Robert Lücking (Botanischer Garten und Botanisches Museum Berlin) and Jack Elix (AustralianNational University, Canberra) kindly reviewed a draft of the manuscript prior to submission.
Disclosure statement
No potential conflict of interest was reported by the authors.
References
Almborn, O. 1955. Lavvegetation och lavflora på Hallands Väderö. Kunglige SvenskaVetenskapsakademien Avhandlinger i Naturskuddsärenden. 11:1–92.
Arup, U, Grube, M. 1998. Molecular systematics of Lecanora subgenus Placodium. Lichenologist.30(4–5):415–425.
Chernomor O, von Haeseler A, Minh BQ. 2016. Terrace aware data structure for phylogenomicinference from supermatrices. Systematic Biology. 65:997–1008.
Culberson, CF. 1972. Improved conditions and new data for identification of lichen products bystandardized thin-layer chromatographic method. Journal of Chromatography A. 72:113–125.
de Lange PJ. 2012. Sole Chatham Islands endemic lichen discovered on south Otago Coastline.Chatham Island New Zealand website. [accessed 2017 May 17]. http://www.chathams.co.nz/index.php/naturalheritage/138-lichen-no-longer-endemic.
Galloway DJ. 1985. Flora of New Zealand. Lichens. Wellington: Government Printer. 662 p.Galloway DJ. 2004. New lichen taxa and names in the New Zealand mycobiota. New Zealand
Galloway DJ. 2007. Flora of New Zealand. Lichens. Vol. 1. Revised 2nd ed. Lincoln: ManaakiWhenua Press. 1006 p.
Gardes M, Bruns TD. 1993. ITS primers with enhanced specificity for basidiomycetes—applicationto the identification of mycorrhizae and rusts. Molecular Ecology. 2:113–118.
Guderley R, Lumbsch HT, Elix J. 1998. Lecanora flavopallida, a species of Lecanora sensu strictowith almost biatorine apothecia (Lecanorales). The Bryologist. 101:103–108.
Hayward GC, Blanchon DJ, Lumbsch HT. 2014. Molecular data support Ramalina ovalis as adistinct lineage (Ramalinaceae, Ascomycota). The Lichenologist. 46:553–561.
Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS. 2017. ModelFinder: fastmodel selection for accurate phylogenetic estimates. Nature Methods. 14:587–589.
Kantvilas, G. 2012. Lecanora coppinsiarum, a new Tasmanian lichen related to Lecanora symmicta.The Lichenologist. 44(2):247–251.
Katoh K, Kuma K, Toh H, Miyata T. 2005. MAFFT version 5: improvement in accuracy of multiplesequence alignment. Nucleic Acids Research. 33:511–518.
Lumbsch HT, Elix JA. 2004. Lecanora. Flora of Australia. 56A:12–62.Meyer B, Printzen C. 2000. Proposal for a standardized nomenclature and characterization of inso-
luble lichen pigments. The Lichenologist. 32:571–583.Minh BQ, Nguyen MAT, von Haeseler A. 2013. Ultrafast approximation for phylogenetic boot-
strap. Molecular Biology and Evolution. 30:1188–1195.Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ. 2015. IQ-TREE: a fast and effective stochastic
algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution.32:268–274.
Penn O, Privman E, Ashkenazy H, Landan G, Graur D, Pupko T. 2010a. GUIDANCE: a web serverfor assessing alignment confidence scores. Nucleic Acids Research. 38:W23–W28.
Penn O, Privman E, Landan G, Graur D, Pupko T. 2010b. An alignment confidence score capturingrobustness to guide tree uncertainty. Molecular Biology and Evolution. 27:1759–1767.
Pérez-Ortega S, Spribille T, Palice Z, Elix JA, Printzen C. 2010. A molecular phylogeny of theLecanora varia group, including a new species from western North America. MycologicalProgress. 9:523–535.
Printzen C. 2001. Corticolous and lignicolous species of Lecanora (Lecanoraceae, Lecanorales) withusnic or isousnic acid in the sonoran desert region. The Bryologist. 104:382–409.
Rodriguez-Flakus P, Printzen C. 2014. Palicella, a new genus of lichenized fungi and its phyloge-netic position within Lecanoraceae. The Lichenologist. 46:535–552.
Ronquist F, Teslenko M, van der Mark P, Ayres D, Darling A, Höhna S, Larget B, Liu L, SuchardMA, Huelsenbeck JP. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and modelchoice across a large model space. Systematic Biology. 61:539–542.
Ryan BD, Lumbsch HT, Messuti MI, Printzen C, Śliwa L, Nash III TH. 2004. Lecanora. In: Nash IIITH, Ryan BD, Diederich P, Gries C, Bungartz F, editors. Lichen flora of the greaterSonoran Desert region. Vol III, Tempe (AZ): Lichens unlimited, Arizona State University;p. 176–286.
Townsend AJ, de Lange PJ, Norton DA, Molloy J, Miskelly C, Duffy C. 2008. The New Zealandthreat classification system manual. Wellington: Department of Conservation. 30 p.
Vilgalys R, Hester M. 1990. Rapid genetic identification and mapping of enzymatically amplifiedribosomal DNA from several Cryptococcus species. Journal of Bacteriology. 172:4238–4246.
White FJ, James PW. 1985. A new guide to microchemical techniques for the identification of lichensubstances. British Lichen Society Bulletin. 57(suppl.):1–40.
White TJ, Bruns TD, Lee SB, Taylor JW. 1990. Amplification and direct sequencing of fungalribosomal RNA genes for phylogenetics. In: Innis MA, Gelfand DH, Sninsky JJ, White TJ,editors. PCR protocols: a guide to methods and applications. San Diego: Academic Press.p. 315–322.
Zhao X, Leavitt SD, Zhao ZT, Zhang LL, Arup U, Grube M, Pérez-Ortega S, Printzen C, Śliwa L,Kraichak E, Divakar PK, Crespo A, Lumbsch HT. 2016. Towards a revised generic classification
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of lecanoroid lichens (Lecanoraceae, Ascomycota) based on molecular, morphological andchemical evidence. Fungal Diversity. 78:293–304.
Zoller S, Scheidegger C, Sperisen C. 1999. PCR primers for the amplification of mitochondrial smallsubunit ribosomal DNA of lichen-forming ascomycetes. The Lichenologist. 31:511–516.