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)
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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. Johnst., Mycotaxon 31: 504
(1988).
Newly obtained DNA sequences were generated in the
Pritzker Laboratory for Molecular Systematics and
Evolution at the Field Museum and at the Molecular
Laboratory, Department of Biology, Faculty of Pharmacy, Complutense University of Madrid. This study was
supported by a grant from the IDP/The Field Museum
Africa Training Fund and the Spanish Ministerio de
Ciencia e Innovación (CGL2013-42498-P).
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