The Lichenologist 49(3): 189–197 (2017) doi:10.1017/S0024282916000748 © British Lichen Society, 2017 The genus Relicinopsis is nested within Relicina (Parmeliaceae, Ascomycota) Paul M. KIRIKA, Pradeep K. DIVAKAR, Steven D. LEAVITT, Kawinnat BUARUANG, Ana CRESPO, George MUGAMBI, Grace W. GATHERI and H. Thorsten LUMBSCH Abstract: Macro-morphological features traditionally used to segregate genera in Parmeliaceae have been shown to be highly plastic, placing limits on their taxonomic value. Here we aim to elucidate the evolutionary relationships of the genera Relicina and Relicinopsis and reassess the phenotypic features traditionally used to separate these genera. To this end, we gathered ribosomal DNA sequences of ITS, nuLSU and mtSSU and analyzed them in a phylogenetic framework. Relicina was recovered as paraphyletic, with Relicinopsis nested within, and three different clades were identified within Relicina. Alternative hypothesis tests significantly rejected the monophyly of Relicina. Our results indicate that the presence or absence of bulbate cilia is of limited taxonomic value in this clade. Based on differences in conidia, however, we propose to accept Relicinopsis as a subgenus within Relicina as Relicina subgen. Relicinopsis (Elix & Verdon) Kirika, Divakar & Lumbsch. It is proposed that five new combinations of species previously classified in Relicinopsis be placed in Relicina. Key words: generic circumscription, integrative taxonomy, lichenized fungi, molecular systematics, parmelioid lichens Accepted for publication 8 September 2016 Introduction Phenotype-based circumscriptions of genera have repeatedly been challenged in different groups of lichenized fungi. In the hyperdiverse family Parmeliaceae, many genera that were chiefly separated based on vegetative traits have not been supported as monophyletic P. M. Kirika and G. W. Gatheri: Department of Plant Sciences, Kenyatta University, P.O. Box 43844-00100, Nairobi, Kenya. P. M. Kirika: Botany Department, National Museums of Kenya, P.O. Box 40658-00100, Nairobi, Kenya. P. K. Divakar and A. Crespo: Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, Madrid 28040, Spain. S. D. Leavitt and H. T. Lumbsch (corresponding author): Science & Education, The Field Museum, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA. Email: tlumbsch@fieldmuseum.org K. Buaruang: Department of Plant Pathology, Faculty of Agriculture, Kasetsart University, Bangkhen, Bangkok, 10900 Thailand. G. Mugambi: Department of Biological Sciences, School of Pure and Applied Sciences, Meru University of Science and Technology, P.O. Box 972-60200, Meru, Kenya. clades in molecular phylogenetic reconstructions (reviewed in Lumbsch 2007; Printzen 2010; Crespo et al. 2011; Thell et al. 2012; Divakar & Crespo 2015). The frequent incongruence between traditional circumscriptions of genera in Parmeliaceae and monophyletic evolutionary lineages highlights the necessity to carefully evaluate generic circumscriptions within an evolutionary context. Currently in Parmeliaceae c. 80 genera are accepted based on phenotypic features and analyses of multilocus sequence data (Thell et al. 2012; Divakar et al. 2015). The largest group within the family is the parmelioid core, to which the genera Relicina (Hale & Kurok.) Hale and Relicinopsis Elix & Verdon belong (Crespo et al. 2010; Divakar et al. 2015). The evolutionary relationships of these two genera have only been partially explored. The genus Relicinopsis was segregated from Pseudoparmelia Lynge based on morphological features, such as the presence of simple marginal cilia, fusiform conidia and usnic acid as a cortical extrolite (Elix et al. 1986). This genus includes a total of five 190 THE LICHENOLOGIST species, which are widely distributed in South-East Asia and Australasia (Hale 1976; Elix 1993, 1994; Divakar & Upreti 2005). The genus Relicina was segregated from Parmelia Ach. s. lat. (Hale 1974) based on having bulbate marginal cilia and bifusiform conidia, and containing usnic acid in the upper cortex. This genus includes c. 54 species (Thell et al. 2012) with a centre of distribution in South-East Asia and Australasia (Hale 1975; Elix 1993). In a recent study, Relicina and Relicinopsis formed a well-supported sister-group relationship, although the taxon sampling was limited and monophyly was not supported in an mtSSU single locus phylogeny (Buaruang et al. 2015). Moreover, the distinction of the two genera was supported in the ‘1GENE’ data analysis by Crespo et al. (2010). In the present study we used an extended taxon sampling to 1) examine the monophyly of Relicina and Relicinopsis and 2) evaluate the taxonomic significance of phenotypic features in these two genera. Materials and Methods Taxon sampling Data matrices of 36 samples including four of the five described species of Relicinopsis and six species of Relicina were analyzed, including eight new samples of these genera collected from East Africa. We assembled a multilocus DNA matrix comprised of nuLSU, ITS and mtSSU rDNA to infer evolutionary relationships. The multilocus data set included 74 sequences from a previous study (Buaruang et al. 2015) and 14 sequences generated for this study. Three species of Notoparmelia were used as the outgroup since the genus has been shown to be closely related to Relicina (Crespo et al. 2010; Buaruang et al. 2015). Information on material studied, including GenBank Accession numbers, is reported in Table 1. DNA extraction and PCR amplification Total genomic DNA was extracted from small pieces of thallus devoid of any visible damage or contamination using the USB PrepEase Genomic DNA Isolation Kit (USB, Cleveland, OH, USA) in accordance with the manufacturer’s instructions. We generated sequence data from nuclear ribosomal markers, the ITS region and a fragment of the nuLSU, in addition to a fragment of the mtSSU. Polymerase chain reaction (PCR) amplifications were performed using Ready-To-Go PCR Beads Vol. 49 (GE Healthcare, Pittsburgh, PA, USA) using dilutions of total DNA. Fungal ITS rDNA was amplified using primers ITS1F (Gardes & Bruns 1993), ITS4 and ITS4A (White et al. 1990; Larena et al. 1999); nuLSU rDNA was amplified using LR0R and LR5 (Vilgalys & Hester 1990); and mtSSU rDNA was amplified using the primers mrSSU1, mrSSU3R and mrSSU2R (Zoller et al. 1999). PCR products were visualized on 1% agarose gel and cleaned using ExoSAP-IT (USB, Cleveland, OH, USA). Cycle sequencing of complementary strands was performed using BigDye v3.1 (Applied Biosystems, Foster City, CA, USA) and the same primers used for PCR amplifications. Sequenced PCR products were run on an ABI 3730 automated sequencer (Applied Biosystems) at the Pritzker Laboratory for Molecular Systematics and Evolution at the Field Museum, Chicago, USA. Sequence editing and alignment New sequences were assembled and edited using Geneious v8.1.7 (Biomatters Ltd 2005–2015). Multiple sequence alignments for each locus were performed using the program MAFFT v7 (Katoh et al. 2005; Katoh & Toh 2008). For the ITS and nuLSU sequences we used the G-INS-i alignment algorithm and ‘20PAM/K = 2’ scoring matrix with an offset value of 0·3 and the remaining parameters set to default values. We used the E-INS-i alignment algorithm and ‘20PAM/ K = 2’ scoring matrix, with the remaining parameters set to default values, for the mtSSU sequences. The program Gblocks v0.91b (Talavera & Castresana 2007) was used to delimit and remove ambiguous alignment nucleotide positions from the final alignments using the online web server (http://molevol.cmima.csic.es/castresana/Gblocks_server. html), implementing the options for a less stringent selection of ambiguous nucleotide positions including “Allow smaller final blocks”, “Allow gap positions within the final blocks” and “Allow less strict flanking positions” options. Phylogenetic analyses Phylogenetic relationships were inferred using maximum likelihood (ML) and Bayesian inference (BI). Exploratory phylogenetic analyses of individual gene topologies showed no evidence of well-supported (≥70% bootstrap values) topological conflict, thus relationships were estimated from a concatenated, three-locus (ITS, nuLSU, mtSSU) data matrix using a total-evidence approach (Wiens 1998). We used the program RAxML v8.1.11 (Stamatakis 2006; Stamatakis et al. 2008) to reconstruct the concatenated ML gene tree using the CIPRES Science Gateway server (http://www.phylo.org/ portal2/). We implemented the ‘GTRGAMMA’ model, with locus-specific model partitions treating all loci as separate partitions, and evaluated nodal support using 1000 bootstrap pseudoreplicates. Exploratory analyses using alternative partitioning schemes resulted in identical topologies and highly similar bootstrap support values. We also reconstructed phylogenetic relationships from the concatenated multilocus data 2017 TABLE 1. Specimens used in this study together with location, reference collection detail and GenBank Accession numbers. Newly obtained sequences for this study are in bold and missing data are indicated with a dash (–). GenBank Accession numbers ITS nuLSU mtSSU Notoparmelia crambidiocarpa N. cunninghamii N. subtestacea Pseudoparmelia cyphellata (8609) P. floridensis (KS3) P. floridensis (KS11) P. floridensis (KS30) P. uleana (8706) Relicina abstrusa (37426) R. abstrusa (1085) R. abstrusa (1082) R. abstrusa (3194) R. abstrusa (3195) R. abstrusa (4505) R. abstrusa 9603) R. abstrusa (9608) R. abstrusa (9619) R. echinocarpa (9317) R. echinocarpa (9623) R. filsonii R. subabstrusa (3193) R. subnigra R. sydneyensis Relicinopsis intertexta (1083) R. intertexta (3177) R. malaccensis (9621) R. malaccensis (9635) R. malaccensis (628) R. malaccensis (1084) R. malaccensis (3172) R. malaccensis (3173) R. malaccensis (3174) R. rahengensis (3169) R. rahengensis (3170) R. rahengensis (3171) R. stevensiae (1073) New Zealand, Knight 60590 (OTA) New Zealand, Knight 60608 (OTA) New Zealand, Knight 60609 (OTA ) Mexico, Nash 46672 (ASU) USA, Scharnagl KS3 (F) USA, Scharnagl KS11 (F) USA, Scharnagl KS30 (F) USA, Seavey 1386 (LSU) Australia, Elix 37426 (CANB) Thailand, Lumbsch 19756g (F) Thailand, Lumbsch 19754f (F) Thailand, Buarang et al. 24368 (RAMK) Thailand, Buarang et al. 24369 (RAMK) Kenya, Kirika 4505 (EA, F, MAF) Kenya, Kirika 4506 (EA, F, MAF) Kenya, Kirika 4541 (EA, F, MAF) Kenya, Kirika & Lumbsch 4032 (EA, F, MAF) Kenya, Kirika & Mugambi 3567 (EA, F) Kenya, Kirika 4432 (F, MAF) Australia, Elix 37267 (CANB) Thailand, Buarang et al. 24370 (RAMK) Australia, Louwhoff et al. (MAF-Lich 10184) Australia, Lumbsch & Mangold 19179a (F) Thailand, Lumbsch 19756g (F) Thailand, Buarang et al. 24372 (RAMK) Kenya, Kirika 4499 (EA, F, MAF) Kenya, Kirika 4508 (EA, F, MAF) Australia, Elix 36972 (hb. Elix) Thailand, Lumbsch 19752a (F) Thailand, Buarang et al. 24373 (RAMK) Thailand, Buarang et al. 24374 (RAMK) Thailand, Buarang et al. 24375 (RAMK) Thailand, Buarang et al. 24376 (RAMK) Thailand, Buarang et al. 24377 (RAMK) Thailand, Buarang et al. 24378 (RAMK) Australia, Elix 37835 (CANB) GU994571 GU994572 GU994573 KM657272 KM657274 KM657273 KM657275 KM657276 GU994580 KM657278 KM657277 KM657279 KM657280 – KX434464 – KX434465 – – KM657281 KM657282 AY785274 GU994581 KM657283 – KX434466 – – KM657284 – – – KM657285 KM657286 KM657287 KM657288 KM657289 KM657290 GU994573 KM657291 KM657293 KM657292 KM657294 KM657295 GU994580 KM657297 KM657296 KM657298 KM657299 – KX434472 KX434473 KX434474 KX434471 KX434476 – KM657300 AY785267 GU994630 KM657301 KM657302 KX434475 KX434477 GU994631 KM657303 KM657304 KM657305 KM657306 KM657307 KM657308 KM657309 KM657310 GU994665 GU994666 GU994668 KM657311 KM657313 KM657312 KM657314 KM657315 – KM657317 KM657316 KM657318 KM657319 KX434467 – – KX434469 KX434468 KX434470 – KM657320 AY785281 GU994675 KM657323 KM657324 – – GU994677 KM657325 KM657326 KM657327 KM657328 – KM657329 KM657330 – 191 Location and collection information Relicinopsis is nested within Relicina—Kirika et al. Taxon (DNA sample number) 192 THE LICHENOLOGIST matrix under BI using the program BEAST v1.8.2 (Drummond & Rambaut 2007). We ran two independent Markov chain Monte Carlo (MCMC) chains for 20 million generations, implementing a relaxed lognormal clock, a birth-death speciation process prior. The most appropriate model of DNA sequence evolution was selected for each marker using the program PartitionFinder v1.1.1 (Lanfear et al. 2012) treating the ITS1, 5.8S, ITS2, nuLSU, and mtSSU as separate partitions. The first 2 million generations were discarded as burn-in. Chain mixing and convergence were evaluated in Tracer v1.5 (Rambaut & Drummond 2009) considering effective sample size (ESS) values >200 as a good indicator. Posterior trees from the two independent runs were combined using the program LogCombiner v1.8.0 (Drummond et al. 2012) and the final maximum clade credibility (MCC) tree was estimated from the combined posterior distribution of trees. Alternative hypothesis testing Since the results of the phylogenetic analyses did not support the monophyly of Relicina as currently circumscribed, we tested whether our data were sufficient to reject the monophyly of that genus. For the hypothesis testing two different methods were employed: 1) Shimodaira-Hasegawa (SH) test (Shimodaira & Hasegawa 1999) and 2) expected likelihood weight (ELW) test (Strimmer & Rambaut 2002). The SH and ELW tests were performed using TREE-PUZZLE 5.2 (Schmidt et al. 2002) with the combined data set on a sample of the best trees agreeing with the null hypotheses and the unconstrained ML tree. These trees were inferred in TREE-PUZZLE employing the GTR+I+G nucleotide substitution model. Morphological and chemical studies Morphological characters, including lobe shape, size and width, and cilia and rhizines were studied using a Leica Wild M8 dissecting microscope. Key morphological and chemical features used to segregate Relicina and Relicinopsis are listed in Table 2. Chemical constituents were identified by high performance thin-layer chromatography using standard methods (Arup et al. 1993; Lumbsch 2001) with a Camag horizontal developing chamber (Oleico Laboratory, Stockholm) using solvent system A. Vol. 49 Results and Discussion Molecular phylogeny and phenotypic features The aligned matrix contained 455 unambiguously aligned nucleotide positions in the ITS, 808 in the nuLSU and 735 in the mtSSU rDNA data sets. The final alignment of the concatenated data set was 1999 positions in length, with 548 variable characters. The ITS PCR product obtained ranged between 600–800 bp. Differences in size were due to the presence or absence of a group I intron of c. 200 bp at the 3' end of the 18S rDNA (Gutierrez et al. 2007). Introns from the ribosomal gene (18S) were removed from the analysis. GTR+I+G for ITS1, K80+I+G for 5.8S rDNA, TrN+G for ITS2, TrN+I+G for nuLSU rDNA and GTR+G for mtSSU rDNA were estimated as best fit models of evolution for each partition. All the newly generated sequences for this study were deposited in GenBank under Accession numbers KX434464–KX434477 (Table 1). Tests for topological incongruence showed no supported conflicts (results not shown). The partitioned ML analysis of the concatenated data matrix yielded an optimal tree with ln likelihood value = − 8048·95 (Fig. 1). In the Bayesian analysis, ESS values of all estimated parameters were well above 200 indicating that convergence among parallel runs was reached. ML and Bayesian topologies were largely similar and did not show well-supported conflict (e.g. PP ≥ 0·95 and ML bootstrap ≥70%) and thus the ML tree topology is shown here with the Bayesian posterior probabilities added (Fig. 1). TABLE 2. Main morphological and chemical features used to distinguish Relicina and Relicinopsis. Features Relicina Relicinopsis Ascospores (µm) Conidia (µm) Marginal cilia Rhizines Habitat Ellipsoid (6–8 × 3–5) to bicornute (10–12 × 3) Bifusiform (6–10 × 1) Bulbate Simple, furcate or agglutinate Tropical-subtropical to temperate Ellipsoid (5–8 × 3–5) Fusiform or cylindrical (5–7 × 1) Simple (without swollen base) Simple or agglutinate Tropical Relicinopsis is nested within Relicina—Kirika et al. 0.99/91 193 Relicinopsis malaccensis_9635 0.99/99 R. malaccensis_9621 R. malaccensis_628 –/73 1.00/100 R. intertexta_3177 R. intertexta_1083 1.00/100 R. malaccensis_3174 R. malaccensis_1084 1.00/99 1.00/100 Clade 1 R. malaccensis_3173 R. malaccensis_3172 0.99/95 R. rahengensis_3169 1.00/100 R. rahengensis_3171 1.00/100 Relicina subgen. Relicinopsis 2017 R. rahengensis_3170 R. stevensiae_1073 Relicina abstrusa_1085 –/67 R. abstrusa_37426 R. abstrusa_1082 R. abstrusa_3194 –/64 R. abstrusa_3195 Clade 2 R. subabstrusa_3193 0.91/85 1.00/100 R. abstrusa_9603 Relicina s. lat. –/97 R. abstrusa_9619 R. abstrusa_9608 –/97 R. abstrusa_4505 0.99/96 R. filsonii 1.00/100 R. sydneyensis Clade 3 R. subnigra 1.00/100 1.00/100 R. echinocarpa_9623 Clade 4 R. echinocarpa_9317 0.99/96 Pseudoparmelia floridensis_KS11 1.00/97 P. floridensis_KS3 1.00/100 1.00/100 P. floridensis_KS30 P. uleana_8706 P. cyphellata_8609 1.00/97 1.00/100 Notoparmelia crambidiocarpa N. subtestacea Out-group N. cunninghamii 0.02 Substitutions per site FIG.1. Phylogenetic relationships of the genera Relicina and Relicinopsis based on maximum likelihood (ML) and Bayesian analyses of a concatenated three locus data set (ITS, nuLSU & mtSSU rDNA.) The ML tree obtained with RAxML is shown here. Posterior probabilities ≥0·95 from the Bayesian analysis (before the slash) and ML bootstrap values ≥70% (after the slash) are given above branches. Three species of Notoparmelia (N. crambidiocarpa, N. cunninghamii and N. subtestacea) were used as the outgroup. 194 THE LICHENOLOGIST Results of the multilocus phylogeny showed that species of the genus Relicinopsis did not cluster with Pseudoparmelia in which they had previously been classified based on morphology (Hale 1975; Swinscow & Krog 1988). In agreement with previous molecular studies (Buaruang et al. 2015; Divakar et al. 2015), however, they grouped with Relicina species. Morphological similarities between Relicinopsis and Relicina species have been discussed previously (Elix et al. 1986; Elix 1993; Divakar & Upreti 2005). In this study, Relicina was recovered as paraphyletic with Relicinopsis nested within it (Fig. 1). Both the SH and ELW tests significantly rejected monophyly of Relicina as currently circumscribed (P ≤ 0·005). These data clearly indicate that the current phenotype-based generic circumscription (Hale 1975) does not reflect evolutionary relationships. Genus-level paraphyly has been found in other groups of parmelioid lichens, including Hypotrachyna (Vain.) Hale (Divakar et al. 2013) and Bulbothrix Hale (Divakar et al. 2010), and similar patterns have been found in other groups of lichen-forming fungi (reviewed in Lumbsch 2007; Printzen 2010). All species of Relicinopsis were recovered in a well-supported (PP = 1·00 and ML bootstrap = 100%) monophyletic clade (clade 1), nested within Relicina (Fig. 1). Clade 1 of Relicinopsis included four of the five species currently known in this genus, including the type species R. intertexta (Mont. & Bosch.) Elix & Verdon. Species of Relicina were grouped in three well-supported monophyletic clades (clades 2, 3 and 4). However, the sister-group relationship of clade 2 and clade 1 recovered in the ML tree lacked support (Fig 1). Furthermore, in the Bayesian tree, clade 2 formed a well-supported (PP = 0·95) sister-group relationship with clade 3 (data not shown). Clade 2 included samples of Relicina abstrusa (Vain.) Hale and R. subabstrusa (Gyeln.) Hale from Australia, Kenya and Thailand, whereas clade 3 consisted of three species, viz. R. filsonii Elix & J. Johnston, R. sydneyensis (Gyeln.) Hale and R. subnigra Elix & J. Johnston occurring in Australasia and South-East Asia. Clade 4 included two samples of R. echinocarpa Vol. 49 (Kurok.) Hale from Kenya. This relationship was strongly supported in the ML analysis (ML bootstrap = 85%) but received weak support in the Bayesian tree reconstruction (PP = 0·91). Our results showed that the relationships among these clades remain unresolved suggesting that additional sampling is necessary to better understand the evolutionary relationships among the clades within the Relicina-Relicinopsis clade. In fact, although all but one of the described Relicinopsis species were studied, only 15 samples collected from Africa, Australia and South-East Asia, representing only six of 54 described Relicina species, were sampled here. Relicina was initially thought to be closely related to Bulbothrix (Hale 1975, 1976; Elix 1993) since both genera are characterized by the presence of bulbate cilia. However, molecular data showed that the two genera were only distantly related with Bulbothrix belonging to the Parmelina clade and Relicina belonging to the Parmelia clade, closely related to Relicinopsis (Crespo et al. 2010; Divakar et al. 2015). The key phenotypic features used to delineate the genera Relicina and Relicinopsis are summarized in Table 2. Both genera differ in the morphology of the marginal cilia (simple in Relicinopsis vs. bulbate in Relicina) and the type of conidia (fusiform or cylindrical in Relicinopsis vs. bifusiform in Relicina). Other characters, such as ascospore form and size, and rhizine morphology overlap (Table 2). Different types of conidia can be found in a number of currently accepted genera in Parmeliaceae such as Hypotrachyna, Melanelixia O. Blanco et al., Melanohalea O. Blanco et al., Myelochroa (Asahina) Elix & Hale, Parmotrema A. Massal., Punctelia Krog and Xanthoparmelia (Vain.) Hale (Crespo et al. 2010); thus this feature can be variable within genera in this family. Consequently, Relicinopsis is reduced here to synonymy with Relicina. However, given that Relicinopsis species formed a well-supported monophyletic clade (clade 1) and are distinguished by conidium morphology, we propose recognising the clade at subgeneric rank. Consequently, the subgenus Relicina is paraphyletic with Relicinopsis nested within it. However, following 2017 Relicinopsis is nested within Relicina—Kirika et al. Divakar et al. (2013) we consider recognition of the monophyletic clade at the subgeneric level preferable since no paraphyletic taxa at generic level are produced (Hörandl & Stuessy 2010). While most of the traditionally circumscribed species in Relicina s. lat. sampled for this study were recovered in monophyletic clusters, a few species did not form monophyletic groups, such as Relicinopsis malaccensis (clade 1) and Relicina abstrusa (clade 2). Additional studies are necessary to evaluate species boundaries in these nominal taxa. Taxonomic Treatment Relicina subgen. Relicinopsis (Elix & Verdon) Kirika, Divakar & Lumbsch comb. et stat. nov. MycoBank No.: MB 817621 Relicinopsis Elix & Verdon, in Elix et al., Mycotaxon 27: 281 (1986); type species: Relicina intertexta (Mont. & Bosch) Kirika, Divakar & Lumbsch, Lichenologist XX: XX (2016). Parmelia intertexta Mont. & Bosch, in Miquel, Pl. Jungh. 4: 445 (1855).— Pseudoparmelia intertexta (Mont. & Bosch) Hale, Phytologia 29: 190 (1974).— Relicinopsis intertexta (Mont. & Bosch) Elix & Verdon, in Elix et al., Mycotaxon 27: 281 (1986). A subgenus in the genus Relicina, corresponding to clade 1 in Fig. 1, including all species currently placed in Relicinopsis (Elix et al. 1986; Elix 1993). New Combinations Relicina dahlii (Hale) Kirika, Divakar & Lumbsch comb. nov. MycoBank No.: MB 817622 Pseudoparmelia dahlii Hale, Smithson. Contr. Bot. 31: 28 (1976).—Relicinopsis dahlii (Hale) Elix & Verdon, in Elix et al., Mycotaxon 27: 281 (1986). Relicina intertexta (Mont. & Bosch) Kirika, Divakar & Lumbsch comb. nov. MycoBank No.: MB 817624 Parmelia intertexta Mont. & Bosch, in Miquel, Pl. Jungh. 4: 445 (1855).— Pseudoparmelia intertexta (Mont. & Bosch) Hale, Phytologia 29: 190 (1974).— Relicinopsis intertexta 195 (Mont. & Bosch) Elix & Verdon, in Elix et al., Mycotaxon 27: 281 (1986). Relicina malaccensis (Nyl.) Kirika, Divakar & Lumbsch comb. nov. MycoBank No.: MB 817623 Parmelia malaccensis Nyl., J. Linn. Soc., Bot. 20: 52 (1883).—Pseudoparmelia malaccensis (Nyl.) Hale, Phytologia 29: 190 (1974).—Relicinopsis malaccensis (Nyl.) Elix & Verdon, in Elix et al., Mycotaxon 27: 282 (1986). Relicina rahengensis (Vain.) Kirika, Divakar & Lumbsch comb. nov. MycoBank No.: MB 817625 Parmelia rahengensis Vain., Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo 1: 39 (1923).—Pseudoparmelia rahengensis (Vain.) Hale, Phytologia 29: 191 (1974).—Relicinopsis rahengensis (Vain.) Elix & Verdon, in Elix et al., Mycotaxon 27: 282 (1986). Relicina stevensiae (Elix & J. Johnst.) Kirika, Divakar & Lumbsch comb. nov. MycoBank No.: MB 817626 Relicinopsis stevensiae Elix & J. 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