The Lichenologist 46(4): 535–552 (2014) 6 British Lichen Society, 2014

doi:10.1017/S0024282914000127

Palicella, a new genus of lichenized fungi and its phylogenetic

position within Lecanoraceae

Pamela RODRIGUEZ FLAKUS and Christian PRINTZEN

Abstract: Non-saxicolous lecideoid lichens form a diverse and polyphyletic group of species with a
worldwide distribution. The phylogenetic relationships of the taxa and major clades are still largely
unresolved. Here we introduce a new genus Palicella Rodr. Flakus & Printzen for Lecidea glaucopa
Hook. f. & Tayl. and two closely related species. Our molecular phylogenetic results, based on
maximum likelihood and Bayesian phylogenetic trees, and a combined dataset of five gene loci
(ITS, nrLSU, mrSSU, RPB1 and RPB2), indicate that Palicella forms a monophyletic clade sister
to Lecanora symmicta and Pyrrhospora quernea. The new genus is characterized by biatorine apothecia,
which are often darkened (by the pigment Cinereorufa-green), an exciple of radiating, narrow hyphae
with strongly elongate lumina, a hymenium with branched and sparsely anastomosed paraphyses, an
ascus with a broad axial body surrounded by a distinct darker staining layer, and narrowly ellipsoid
ascospores. Atranorin is present in all, usnic acid in two and pannarin in one species of the genus.
Three new combinations are proposed: Palicella filamentosa (Stirt.) Rodr. Flakus & Printzen, P.
glaucopa (Hook. f. & Taylor) Rodr. Flakus & Printzen, and P. schizochromatica (Pérez-Ortega, T.
Sprib. & Printzen) Rodr. Flakus & Printzen. The molecular phylogenetic analysis confirms that
Lecidea hercynica Hauck & Schmull is a synonym of P. filamentosa. An identification key to the species
is provided.
Key words: combined analysis, Lecanora, Lecanorales, Lecidea, lecideoid lichens, taxonomy
Accepted for publication 29 January 2014

Introduction 1991; Printzen 1995; Printzen & Tønsberg

1999; Printzen & Kantvilas 2004; Czarnota
The genus Lecidea Ach. in its broad sense
2007; Kalb et al. 2011; Kantvilas & Elix
consists of species with hyaline, non-septate
2013).
ascospores, lecideine apothecia, a chlorococ-
As a result of recent phylogenetic studies,
coid photobiont, and crustose thalli (e.g.
it is known that most non-saxicolous leci-
Zahlbruckner 1925, 1940; Rambold 1989;
deoid species are only distantly related to
Hertel & Printzen 2004). It is the largest
the saxicolous genus Lecidea s. str. and repre-
group of lichens with a worldwide distri-
sent a polyphyletic assemblage (Schmitt et al.
bution and diversity hot spots in temperate
2003; Buschbom & Mueller 2004; Andersen
and polar regions of both hemispheres (e.g. & Ekman 2005; Pérez-Ortega et al. 2010;
Hertel 1997). The corticolous members of
Schmull et al. 2011). Because only a few spe-
the group have received less attention than
cies have been so far included in phyloge-
the saxicolous ones (e.g. Hertel 1967, 1984, netic analyses, the systematic position of the
1995, 2006; Hafellner 1984; Rambold 1989),
majority of non-saxicolous genera and spe-
and there are only a few, mostly regional,
cies described under the name Lecidea is still
taxonomic revisions of non-saxicolous spe-
unclear and requires further study.
cies (e.g. Coppins 1983; Brako 1991; Timdal
While studying the phylogeny and sys-
tematics of lecideoid non-saxicolous lichens
P. Rodriguez Flakus and C. Printzen: Department of from southern South America, we also ex-
Botany and Molecular Evolution, Senckenberg Research amined material originally described as Leci-
Institute, Senckenberganlage 25, D-60325, Frankfurt am
Main, Germany.
dea glaucopa Hook. f. & Taylor (Hooker &
Email: pamela.rodriguez@senckenberg.de Taylor 1844). This species was later trans-
536 THE LICHENOLOGIST Vol. 46

ferred to Biatora (Räsänen 1932) and finally Zeiss HYRAX KS 34, mounted in water or lacto-
to Lecidella (Kalb 1984). However, its nar- phenol cotton blue for permanent slides and examined
with a Zeiss stereomicroscope Stemi 2000 and Leitz
rowly elongate, apically widened excipular DIAPLAN microscope. Microscopic measurements were
hyphae did not fit those of any known Biatora carried out according to Printzen (1995) and Printzen &
species, and the narrow ascospores and con- May (2002). Spore measurements for P. glaucopa are
glutinate paraphyses distinguished it from presented in the following way: (arithmetic mean – stan-
dard deviation) arithmetic mean (arithmetic mean +
typical representatives of Lecidella. The ascus standard deviation), flanked by the minimal and maxi-
type also did not match exactly with that of mal measurements in parentheses, followed by the
both genera. number of measurements (n). The length/breadth ratio
The highly variable colour of the apothe- (l/b) and apothecial diameter are presented in the same
cia, ranging from light beige to black, often way. Insoluble lichen pigments were tested according to
Meyer & Printzen (2000). Secondary metabolites were
on the same apothecium, makes L. glaucopa identified using thin-layer chromatography (TLC) in
superficially similar to the North American solvents A, B and C, following the methods described
Lecanora schizochromatica Pérez-Ortega et al. by Culberson & Kristinsson (1970) and Orange et al.
This species has been assigned to the usnic (2001). For confirmation of results and identification of
xanthones, HPLC was used according to Feige et al.
acid containing members of Lecanora s. lat., (1993).
the so-called Lecanora varia group (Choisy
1929; Śliwa & Wetmore 2000; Printzen DNA extraction, polymerase chain reaction (PCR)
2001). Recent studies have shown that this and DNA sequencing
group is also not monophyletic and com-
DNA was isolated from two to three apothecia free
prises several smaller clades distinguished by from lichenicolous fungi with the DNeasyTM Plant
anatomical characters (Arup & Grube 1998; Mini Kit (QIAGEN) following the manufacturer’s in-
Printzen 2001; Pérez-Ortega et al. 2010). structions. We used the genomic DNA extract in PCR
Based on DNA sequence data from the inter- reactions of the mycobiont genes coding for: 1) the inter-
nal transcribed spacer of the nuclear ribosomal RNA re-
nal transcribed spacer of the nuclear riboso- peat (ITS), and parts of the 2) mitochondrial ribosomal
mal DNA repeat, Pérez-Ortega et al. (2010) large subunit DNA (mrLSU), 3) nuclear ribosomal
found evidence of a close relationship be- large subunit (nrLSU), 4) RNA Polymerase II largest
tween L. schizochromatica and L. filamentosa subunit (RPB1), and 5) RNA Polymerase II second
(Stirt.) Elix & Palice. largest subunit (RPB2). Primers used for amplifications
were: ITS4 (White et al. 1990), ITS1F (Gardes & Bruns
Here we introduce a new genus, Palicella 1993), LR7 (Vilgalys & Hester 1990), LR0R (Rehner &
Rodr. Flakus & Printzen, and investigate its Samuels 1994), mSSU1 (Zoller et al. 1999), MSU7
phylogenetic affinities. We propose three (Zhou & Stanosz 2001), RPB1cr (Stiller & Hall 1997),
new combinations, P. filamentosa (Stirt.) Rodr. RPB1Af (Matheny et al. 2002), RPB2-5F and RPB2-
7cr (Liu et al. 1999). PCRs of ITS, mrSSU and nrLSU
Flakus & Printzen, P. glaucopa (Hook. f. & contained 5 ml of DNA extract, 1 ml of each primer (10
Taylor) Rodr. Flakus & Printzen and P. mM), 18 ml of distilled water and PCR PuReTaqTM
schizochromatica (Pérez-Ortega, T. Sprib. & Ready-To-GoTM Beads (GE Healthcare, USA). For
Printzen) Rodr. Flakus & Printzen, and con- RPB1 and RPB2 we used 15 ml of DNA extract and 5 ml
firm by molecular phylogenetic data that of each primer. Thermal cycling parameters were: initial
denaturation at 94 C (5 min), four cycles of 94 C (30 s),
Lecidea hercynica Hauck & Schmull is a syn- 54 C (30 s), 72 C (1 min), 32 cycles of 94 C (30 s),
onym of P. filamentosa. 48 C (30 s), 72 C (1 min) and a final extension of 10
min at 72 C for ITS, mrSSU and nrLSU; initial denatu-
ration at 95 C (5 min), followed by eight cycles of 95 C
(1 min), 58 C (1 min), 72 C (145 min), 34 cycles of
Material and Methods 95 C (1 min), 50 C (1 min), 72 C (145 min) and a
Taxon sampling, morphology and chemistry final extension of 10 min at 72 C for RPB1; and initial
denaturation at 94 C (2 min), followed by eight cycles
The description of P. glaucopa is mainly based on of 94 C (1 min), 59 C (1 min), 72 C (2 min), 33 cycles
recent collections from Argentina and Chile deposited of 94 C (30 s), 50 C (30 s), 72 C (2 min) and a final ex-
in FR. We studied herbarium material from BM, FH, tension of 10 min at 72 C for RPB2. PCR products were
G, H, H-Nyl., MICH, S, UPS and the private herbaria visualized on agarose gels under blue light, bands were
of K. Kalb (Neumarkt) and J. Etayo (Pamplona). Ana- cut out from the gels and purified using the QIAquick
tomical sections 15 mm thick were cut with a cryotome Gel Extration Kit (QIAGEN, Germany). Purified DNA
2014 Palicella—Rodriguez Flakus & Printzen 537

was labelled with the BigDyeTM Terminator v3.1 Cycle means of the exponential distributions. Two supported
Sequencing Kit (Applied Biosystems) and cycle se- conflicts were detected by us. On the ITS tree, Parmo-
quenced at 94 C for 30 s, and 29 cycles of 95 C for 15 trema tinctorum grouped with Hypotrachyna revoluta,
s, and 60 C for 4 min using the PCR primers and, for while mrSSU supported a relationship with Puntelia
nrLSU, the additional internal primers LR6 (Vilgalys borreri. In ITS, Protoparmelia badia appeared as sister to
& Hester 1990) and LR3R (Vilgalys’s laboratory, un- Mycoblastus sanguinarius but grouped with H. revoluta
published, http://www.botany.duke.edu/fungi/mycolab). and P. borreri in RPB1. Because both conflicts only
Sequences were determined on an ABI PRISM 3730 involved outgroup taxa, we decided to also analyze the
DNA Analysis System (Applied Biosystem), and assem- concatenated five-gene dataset.
bled and edited using Geneious Pro, version 5.0.4 (Bio- The inferred branch length prior for the MCMC
matters Ltd). analysis of this dataset followed an exponential distribu-
tion with mean 1/18. MrBayes was set to sample every
1000th tree from three independent runs, each with
Sequence alignment and phylogenetic four chains that were incrementally heated using a factor
reconstruction of 015. The average standard deviation of bipartition
A total of 32 sequences for ITS, 33 sequences for frequencies among runs was calculated every 1 000 000
mrSSU, 33 sequences for nrLSU, 24 sequences for generations to infer convergence of the Markov Chains,
RPB1 and 22 sequences for RPB2 were analyzed. discarding the first 50% of the trees sampled as burn-in
BLAST searches in GenBank were performed to ascer- and including only those bipartitions with a frequency of
tain that all sequences used in the phylogenetic analyses at least 10%. The analysis was stopped after 3 000 000
originated from the lichens and not from contaminating generations when the standard deviation had dropped
organisms such as parasymbiotic fungi. The outgroup below 001. Finally, we also calculated a ML bootstrap
sequences from families of Cladoniaceae, Stereocaulaceae tree (1000 replicates) for the concatenated dataset using
and Parmeliaceae were selected based on previous phylo- RaxML version 7.2.6 (Stamatakis 2006), with the inter-
genetic studies (Arup & Grube 1998, 2000; Mia˛dlikowska face RaxMLGUI version 0.9 beta 2 (Silvestro & Michalak
et al. 2006). The complete dataset including sequences 2012) and unlinked GTRGAMMAI models for the six
downloaded from GenBank is described in Table 1. Se- partitions inferred by PartitionFinder.
quences were aligned within Geneious Pro version 5.0.4
using the Muscle algorithm and default settings (Edgar
2004). We removed poorly or ambiguously aligned re-
gions using Gblocks version 0.91b (Castresana 2000) Results
with default settings but allowing gaps in 50% of the se-
quences. This resulted in five alignments of the follow- A total of 56 new DNA sequences were ob-
ing length: ITS-425 bp, mrSSU-709 bp, nrLSU-738 tained for this study (Table 1). The final
bp, RPB1-675 bp and RPB2-1059 bp. To select an opti- alignment contained 40 OTUs and 3606 un-
mal partitioning scheme and substitution models for the ambiguously aligned nucleotide positions.
partitions, we concatenated the single gene dataset and
used PartitionFinder version 1.0.1 (Lanfear et al. 2012) The inferred optimal partitioning scheme
with default settings using the Akaike Information Crite- and substitution models for the dataset are
rion for model selection and suggesting 11 data blocks listed in Table 2. The total length of the ML
(ITS1, 5.8S rDNA, ITS2, mrSSU, nrLSU, and three tree was 420, the mean total branch length
independent codon positions for RPB1 and RPB2). of the Bayesian consensus tree 313. Figure 1
Before analyzing the concatenated dataset, we calcu-
lated phylogenetic trees for the single gene dataset, in shows the maximum likelihood tree with ML
order to detect potential conflicts. Phylogenetic analyses bootstrap values and posterior probabilities.
were carried out with MrBayes v.3.2 (Huelsenbeck & The Southern Hemisphere specimens of
Ronquist 2003), using the substitution models inferred Lecidea glaucopa Hook. f. & Taylor were
with PartitionFinder and default settings with the fol-
lowing exceptions. The ITS dataset was partitioned into grouped in a clade with high support values
two data partitions (ITS1+ITS2, 5.8S), and codon- (BP ¼ 100 and PP ¼ 10), and are also ana-
specific partitions were used for RPB1and RPB2. All tomically and chemically differentiated from
parameters of the substitution model were unlinked the other taxa in the dataset. A close relation-
between partitions. Gamma distributed rates across sites ship between L. glaucopa, Lecanora filamen-
were modelled as six discrete categories and with a mean
1 exponential prior, and an unconstrained, exponential tosa and Lecanora schizochromatica is also
branch length prior was used. The mean of this prior supported with BP ¼ 100 and PP ¼ 10.
was inferred by calculating ML trees for the datasets Lecidea hercynica M. Hauck & Schmull
using RaxMLGUI version 0.9 beta 2 (Stamatakis 2006; groups with the two accessions of Palicella
Silvestro & Michalak 2012) and applying either an un-
partitioned GTRGAMMAI model (mrSSU, nrLSU) or
filamentosa (BP ¼ 100, PP ¼ 10). The short
a partitioned GTRGAMMAI (the rest) and 20 runs. branches within this clade and anatomical
The mean branch lengths of the ML trees were used as similarities between the two taxa indicate
 Table 1. GenBank accession numbers and voucher information for taxa and sequences used in this paper.

538
GenBank Accession Numbers

Taxon Source – Specimen ITS mrSSU nuLSU RPB1 RPB2

Adelolecia pilati GenBank – Austria, Ekman 3373 (BG) n/a AY567713 n/a AY756379 n/a
Cladia aggregata GenBank – Australia, Kantvilas s.n. (HO) n/a GQ500936 GQ500972 n/a n/a
Cladonia rangiferina Genbank – Sweden, Wedin 50811 (BM); n/a AY3008811 AY3008321 DQ9155952 n/a
Sweden, Wedin 69352 (UPS)
Frutidella caesioatra GenBank – Norway, Andersen 91 (BG) n/a AY567765 AY756349 AY756383 n/a
Haematomma flexuosum GenBank – Costa Rica, Lücking 15097h1; Costa EU0755601 EU0755512 EU0755552 n/a n/a
Rica, Lücking 17626x32 (F)

THE LICHENOLOGIST
Hypotrachyna revoluta GenBank – Spain, Noya & Olea 60471 (MAF); AY6110751 DQ2878312 AY6077871 EF0921122 n/a
China, Crespo et al. 103942 (MAF)
Lecanora dispersa GenBank – Korea, Hur H06038 EU266081 n/a n/a n/a n/a
L. polytropa GenBank – AFTOL-ID 1798 (DUKE) HQ650643 DQ986807 DQ986792 n/a DQ992418
Lecanora symmicta Germany, Printzen 9999a (FR) GU480113 KJ152466 KJ152453 KJ152452 KJ152487
Lecidella carpathica GenBank – Austria, Arup L97005 (hb. Arup) AY541274 n/a n/a n/a n/a
L. elaeochroma GenBank – Belgium, Ertz 7605 (DUKE) HQ650605 n/a DQ986747 DQ986818 DQ992429
L. euphorea GenBank – AFTOL-ID 1374 (DUKE) HQ650596 DQ986784 n/a DQ986857 DQ992479
L. greenii GenBank – Antarctica, de los Rı́os s.n. (MA) JX036133 n/a n/a n/a n/a
L. meiococca GenBank – Sweden, Ekman 3101 (BG) AF517929 AY567714 AY300842 n/a n/a
Lepraria lobificans GenBank – Costa Rica, AFTOL-ID 325 n/a DQ986887 DQ986768 DQ986837 DQ992454
(DUKE)
Miriquidica garovaglii GenBank – Austria, Hafellner 585711 (GZU); EU0755381 AY5677112 AY7563572 AY7564202 n/a
Norway, Ekman s.n.2 (BG)
Mycoblastus sanguinarius GenBank – Sweden, AFTOL-ID 1961 (DUKE); DQ7828421 n/a AY7563412 AY7563933 DQ7828671
Wedin 69322 (UPS); Norway, Ekman s.n.3 (BG)
Palicella filamentosa 99 GenBank – Germany, Cezanne & Eichler 6761 GU480099 KJ152468 n/a n/a n/a
(hb. Cezanne-Eichler)
P. filamentosa 100 GenBank – Czech Republic, Printzen s.n. (FR) GU480100 n/a n/a n/a n/a

Vol. 46
P. filamentosa GenBank – Hauck s.n. (FH) HQ650663 HQ660568 n/a n/a n/a
(‘‘Lecidea hercynica’’ )
P. glaucopa 206 Chile, Rodriguez Flakus 2101 & Flakus (FR) KJ152484 KJ152473 KJ152456 KJ152443 KJ152490
 2014
Table 1. Continued

GenBank Accession Numbers

Taxon Source – Specimen ITS mrSSU nuLSU RPB1 RPB2

Palicella glaucopa 286 Argentina, Rodriguez Flakus 2447 & Flakus (FR) KJ152480 n/a KJ152464 KJ152449 KJ152494
P. glaucopa 288 Argentina, Rodriguez Flakus 2504d & Flakus (FR) n/a KJ152469 KJ152459 KJ152445 KJ152498
P. glaucopa 289 Argentina, Rodriguez Flakus 2504c & Flakus (FR) KJ152482 KJ152476 KJ152455 KJ152447 KJ152493
P. glaucopa 290 Argentina, Rodriguez Flakus 2504b & Flakus (FR) KJ152485 KJ152472 n/a KJ152444 KJ152496

Palicella—Rodriguez Flakus & Printzen

P. glaucopa 309 Argentina, Rodriguez Flakus 2540 & Flakus (FR) KJ152483 KJ152474 KJ152461 KJ152448 KJ152492
P. glaucopa 310 Argentina, Rodriguez Flakus 3174 & Flakus (FR) KJ152479 KJ152475 KJ152463 KJ152450 KJ152497
P. glaucopa 311 Chile, Rodriguez Flakus 2672 & Flakus (FR) KJ152481 KJ152477 KJ152462 KJ152446 KJ152491
P. glaucopa 324 Argentina, Rodriguez Flakus 2539 & Flakus (FR) KJ152486 KJ152471 KJ152460 KJ152451 KJ152495
P. glaucopa 348 Argentina, Rodriguez Flakus 2504a & Flakus (FR) KJ152478 KJ152470 KJ152458 n/a KJ152499
P. schizochromatica 1073 GenBank – Canada, Spribille 17016 GU480109 KJ152467 KJ152454 n/a KJ152488
(hb. Spribille)
P. schizochromatica 1070 GenBank – Canada, Spribille 16850A GU480110 KJ152465 KJ152457 n/a KJ152489
(hb. Spribille)
P. schizochromatica GenBank – USA, Hauck s.n. (hb. Hauck) HQ650652 HQ660555 HQ660532 n/a n/a
(‘‘Lecanora fuscescens’’ )
Parmotrema tinctorum GenBank – USA, AFTOL-ID 7 (DUKE) HQ650684 AY584627 AY584635 n/a AY584687
Protoparmelia badia GenBank – Austria, Hafellner 590431 (GZU); EU0755401 EU0755261 DQ8710112 DQ8709872 n/a
Spain, Lumbsch s.n.2 (F)
Punctelia borreri GenBank – Canary Island, Crespo102461; Spain, AY6134001 AY6134212 AY5789543 EF0921494 n/a
Divakar 102542; Portugal, Crespo 99193 & Crespo
et al. s.n.4 (MAF)
Pyrrhospora quernea GenBank – Sweden, Ekman 3019 (BG) AF517930 AY567712 AY300858 n/a n/a
Ramboldia stuartii GenBank – Australia, Tasmania, Elix 28664 (F) EU075549 EU075535 EU075522 n/a n/a
Stereocaulon tomentosum GenBank – Norway, Tønsberg 273351 (BG); n/a AY7563781 AY3405692 AY7564271 n/a
Sweden, Wedin 50892 (UPS)
Xanthoparmelia conspersa GenBank – USA, AFTOL-ID 4 (DUKE) n/a AY584633 AY584641 n/a AY584691

539
540 THE LICHENOLOGIST Vol. 46

Table 2. Optimal partitioning scheme and substitution models for each data partition

Subset Best Model Partitions Subset Sites

1 GTR+I+G ITS1, ITS2 1–154, 306–425

2 K80 5.8S 155–305
3 HKY+I+G mrSSU, RPB1 codon 3 426–1134, 1875–2547/3
4 GTR+I+G nrLSU, RPB1 and RPB2 codon 2 1135–1872, 1874–2547/3, 2549–3606/3
5 SYM+I+G RPB1 and RPB2 codon 1 1873–2547/3, 2548–3606/3
6 K80+G RPB2 codon 3 2550–3606/3

that they are conspecific. Likewise, GenBank Lecidella-type). Ascospores narrowly to rarely broadly
sequences of Lecanora fuscescens form a well- ellipsoid, colourless, simple or rarely 1-septate. Major
substances: atranorin.
supported group (BP ¼ 99, PP ¼ 10) with Generic type: Palicella glaucopa (Hook. f. & Taylor)
two collections of P. schizochromatica. Rodr. Flakus & Printzen.
Palicella appears as sister to a clade formed
by Lecanora symmicta and Pyrrhospora quernea Etymology. The new genus is named in
and is hence assigned to the family Lecanora- honour of the prominent Czech licheno-
ceae, although this relationship is only sup- logist, Dr Zdeněk Palice (Průhonice), for
ported by the MCMC analysis (BP ¼ 66, his substantial contributions to lichenology,
PP ¼ 099). The distinction between Leca- especially regarding lecideoid members of
noraceae and Parmeliaceae is not supported Lecanora.
because the backbone of the tree lacks statis-
tical support. Palicella filamentosa (Stirt.) Rodr.
Flakus & Printzen comb. nov.
MycoBank No.: MB807638
Taxonomic Treatment
Lecidea filamentosa Stirt., Scottish Naturalist 5: 218
Palicella Rodr. Flakus & Printzen [‘1879’] (1880).—Lecanora filamentosa (Stirt.) Elix &
Palice Mycol. Progress 9: 531 (2010); type: Scotland,
gen. nov. Glen Lochay, Killin, 21 July 1879, Stirton s. n. [GLAM-
NHB.1927.8.01010—lectotype, designated by Pérez-
MycoBank No.: MB807126 Ortega et al. (2010: 531); BM-731241—isolectotype].
Thallus crustose, rimose-areolate, partially continuous, For a full synonymy of the species see Palice et al.
initially endosubstratal; photobiont chlorococcoid. Apoth- (2011).
ecia very variable in colour from light yellow, pinkish
brown, bluish grey to greenish black, epruinose to prui- Remarks. Within the genus, the species is
nose, margin always present but frequently reduced characterized by pale ochre to charcoal grey,
when old. Exciple hyaline or brownish inside, often
with a pigmented outer layer (Cinereorufa-green and flat to convex apothecia with a biatorine mar-
unknown brown pigment), composed of strongly gelati- gin that, in young apothecia, is sometimes
nized branched and anastomosing, sometimes apically surrounded by a thalline amphithecium, as-
widened hyphae, often inspersed with crystalline gran- cospores (100–)124–135(–160)  (35–)
ules or oil droplets; young apothecia rarely surrounded
by an additional thalline margin. Subhymenium and
42–45(–50) mm, and the production of
hypothecium colourless. Epihymenium with hyaline to small amounts of usnic acid in addition to
pale ochre granules, and sometimes greenish black pig- atranorin (Printzen & May 2002; Palice et
mented (Cinereorufa-green). Hymenium colourless or al. 2011).
pale bluish green or ochre above, composed of gelati-
nized, weakly branched and anastomosing paraphyses,
with not or slightly expanded apices. Asci 8-spored, tho-
Distribution and ecology. Palicella filamen-
lus I+ blue with a wide axial body that is mostly sur- tosa is a rather widespread corticolous and
rounded by a distinct darker staining layer (Lecanora/ lignicolous species known from Europe and
 Table 3. Characters distinguishing Palicella from similar genera.

2014
Character Palicella Lecanora symmicta Cliostomum Lecidella Pyrrhospora quernea Ramboldia elabens
group

Exciple Radial hyphae Radial hyphae with Radial hyphae with Radial hyphae with Radial hyphae with Radial hyphae with
(sometimes irregular narrowly cylindric wide, cylindrical, thin-walled cylindri- thick-walled, cylin- thin-walled, cylindri-
inside) with narrowly lumina, apically ellipsoid to globose cal to ellipsoid drical lumina, api- cal lumina, usually
cylindric lumina, weakly swollen, lumina, apically lumina, apically cally weakly swollen, inspersed by red
apically weakly to strongly gelatinized weakly swollen, weakly swollen inspersed by red granules
strongly swollen, and outer part with strongly gelatinazed, granules
strongly gelatinized, granules usually with granules
sometimes with oil

Palicella—Rodriguez Flakus & Printzen

droplets
Pigments Cinereorufa-green Pale yellowish brown Pale to dark brown Cinereorufa-green in Unknown black to Unknown black and
and unknown brown pigments in epithe- pigments in epithe- epithecium and outer orange-brown pig- orange-brown pig-
pigment in epithe- cium (K--) cium (K--); black layer of exciple (K+ ments; epithecium, ments in epithecium
cium and outer layer pigment in pycnidia green); sometimes interspersed with and outer layer of
of exciple (K+ purple) brown pigments in granules, K+ dissolv- exciple; K+ pale
(K+ green) inner part of exciple ing, purplish (7- greyish brown
and hypothecium chloroemodin)
Paraphyses Branched, sparsely Weakly branched Simple or sparingly Simple, apically Simple to branched Sparsely branched
anastomosed, and anastomosed, branched and widened towards apices, and anastomosed,
apically unwidened apically unwidened anastomosed in apically widened mostly apically
upper part, apically unwidened
widened
Ascus type Lecanora-Lecidella Biatora Biatora Lecidella Lecanora Lecanora
Ascospores 0- rarely 1-septate, 0- rarely 1-septate, 1–3-septate, Simple, broadly Simple, broadly Simple, narrowly
narrowly ellipsoid narrowly ellipsoid narrowly ellipsoid, ellipsoid ellipsoid ellipsoid to bacilli-
oblong or bacilliform form
Chemistry
+atranorin major – major major trace e
+usnic acid e major – – – –
+xanthones e + – e major –
+terpenoids, e e e e + e
anthraquinones

541
542 THE LICHENOLOGIST Vol. 46

76 / 0.94 Cladonia rangiferina

99 / 1.0 Cladia aggregata

85 / 1.0 Stereocaulon tomentosum

Lepraria lobificans
Miriquidica garovaglii
Ramboldia stuartii
60 / 1.0
Mycoblastus sanguinarius
99 / 1.0 Protoparmelia badia
Hypotrachyna revoluta

100 / 1.0 Xanthoparmelia conspersa

71 / 0.97
Punctelia borreri
Parmotrema tinctorum
Haematomma flexuosum
Lecanora dispersa

99 / 0.92 Lecidella carpathica

Lecidella greenii
Lecidella euphorea
99 / 1.0
71 / 0.77 Lecidella elaeochroma
98 / 1.0
Lecidella meiococca
Lecanora polytropa
Frutidella caesioatra
Adelolecia pilati

80 / 1.0 Pyrrhospora quernea

Lecanora symmicta
“Lecidea hercynica”
100 / 1.0
Palicella filamentosa 100
66 / 0.99
100 / 1.0 Palicella filamentosa 99
Palicella schizochromatica 1070
99 / 1.0 Palicella schizochromatica 1073
100 / 1.0 “Lecanora fuscescens”
100 / 1.0 Palicella glaucopa 348
Palicella glaucopa 288
100 / 1.0 Palicella glaucopa 290
Palicella glaucopa 310
99 / 1.0
Palicella glaucopa 206
96 / 1.0
Palicella glaucopa 311
Palicella glaucopa 324
Palicella glaucopa 286
76 / 1.0
Palicella glaucopa 289
87 / 0.99 Palicella glaucopa 309

0.06

Fig. 1. Maximum likelihood consensus phylogram obtained from 40 species and five gene loci (ITS, nrLSU,
mrSSU, RPB1, RPB2) of the Lecanoraceae clade. The families Cladoniaceae, Parmeliaceae and Stereocaulaceae were
used as outgroups. Support values (in bold) are given above the branches. ‘‘Palicella’’ as a new genus with a value of
(PP ¼ 1) for MCMC and ML bootstrap (BP ¼ 100).
2014 Palicella—Rodriguez Flakus & Printzen 543

eastern North America (Printzen & May white, matt or shiny, flat to moderately con-
2002; Palice et al. 2011). vex (in old apothecia), greyish pruinose (es-
pecially young apothecia) or epruinose; mar-
gin prominent but often reduced when older,
Palicella glaucopa (Hook. f. & Taylor) often becoming flexuose, shiny black; exciple
Rodr. Flakus & Printzen comb. nov. laterally 40–160 mm wide, basally 70–220
mm wide, colourless within but strongly in-
MycoBank No.: MB807639
spersed by oil droplets and small crystals
Lecidea glaucopa Hook. f. & Taylor in Hook., London (POL+), I--, outer layer with dark pigmenta-
J. Bot. 3: 637 (1844).—Biatora glaucopa (Hook. f. & tion (Cinereorufa-green; Meyer & Printzen
Taylor) Räsänen, Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo
2: 58 (1932).—Lecidella glaucopa (Hook. f. & Taylor) 2000, and unknown brown pigment), K+
Kalb, Lichenes Neotropici, fascicle 8: 328 (1984); type: green, N+ purple-red, composed inside of
Chile, Cape Horn, [March–September 1842, J. D. loosely arranged, narrow hyphae, 08–20
Hooker] (BM-1104684!—lectotype, designated here; mm wide, that are apically widened to 5–8
FH-302837!, BM-1104685!—isolectotypes).
Lecidea conflectens Nyl., Lich. Fueg. Patag.: 15 (1888);
mm (lumina c. 3–5 mm); hypothecium colour-
type: [Argentina,] Fuegia, Ushuwaia, 1882, in Fago less to pale yellowish brown, 40–75 mm high,
antarctica, Spegazzini (H-NYL-20244!—lectotype, de- I+ blue; subhymenium 10–30 mm high, col-
signated here). ourless, I+ blue; hymenium 50–90 mm high,
Lecidea oculans Nyl., Lich. Fueg. Patag.: 13 (1888).— colourless, conglutinate, I+ blue; epihyme-
Lecidea glaucopa var. oculans (Nyl.) Zahlbr., Cat. Lich.
Univ. 3: 768 (1925).—Biatora glaucopa var. oculans nium 8–20 mm thick, light yellow to greenish
(Nyl.) Räsänen, Ann. Bot. Soc. Zool.-Bot. Fenn. Vanamo black, with Cinereorufa-green pigments and
2: 58 (1932); type: [Argentina,] Fuegia, Wallamatu, unidentified brown pigment, epipsamma of
1882, fagicola, Spegazzini (H-NYL-21305!—lectotype, hyaline, sometimes also yellow, granules
designated here).
Lecidea epichlorotica Müll. Arg., Miss. Sci. Cap Horn,
(POL+); paraphyses colourless, branched
Lich.: 164 (1889); type: [Chile], Cap Horn, Baie Orange, and anastomosed, conglutinate in water but
Ile Hoste, Terre de Feu, 1886, Fagus betuloides, Hariot S. easily separated in KOH, 10–15 mm, api-
52 pr.p. (G!—lectotype, designated here; G!—para- cally 15–25 mm wide; asci 8-spored, clavate,
types). 47–75  11–20 mm, with K/I+ blue tholus
Lecidea epichlorotica var. expallens Müll. Arg., Miss.
Sci. Cap Horn, Lich.: 165 (1889); type: [Chile], Ile resembling the Lecanora- or Lecidella-type,
L’Hermite, Baie San Martin, Terre de Feu, 1886, cortex usually with a small unstained region at the
Fagi betuloidis, Hariot S. 34. (G!—lectotype, designated top and a rather wide, apically closed axial
here; G!—paratype). body (occupying about 1/3 of the tholus
(Fig. 2) width) that may almost penetrate the tholus
and is surrounded by a strongly amyloid
Thallus corticolous, up to 15 mm diam., zone (less so in apical parts), ocular chamber
crustose, initially endosubstratal, later rimu- of paraboloid to almost cylindrical or slightly
lose-areolate, composed of dispersed to con- clavate shape; ascospores colourless, simple to
fluent, convex areoles up to 04 mm diam., rarely 1-septate, narrowly ellipsoid, variable
partially continuous, 40–150 mm thick; sur- in shape, sometimes slightly curved or irreg-
face matt, pale grey to white or yellowish ular, filled with large oil droplets, (90–)
white; prothallus sometimes present, white or 159–195–230(–320)  (48–)55–65–
black; pseudocortex very thin, indistinct, of 74(–95) mm, length-breadth index (16–)
loosely arranged gelatinized hyphae, 5–10 24–31–38(–56) mm (n ¼ 300), walls 05–
mm thick; medulla discontinuous, 30–60 mm 15 mm thick.
thick or partially lacking, composed of c. 2 Pycnidia rarely present, black, immersed
mm wide hyphae, with groups of chlorococ- in the thallus, marginal, walls paraplecten-
coid photobiont cells 9–19 mm diam. chymatous with brown wall, conidiophores
Apothecia rounded to irregular, single, unbranched, resembling type II–III of Vobis
never aggregated in groups, sessile with con- (1980); conidia straight to very slightly curved,
stricted base (03–)05–07–10(–14) mm filiform, borne apically on elongated coni-
(n ¼ 10); disc variable in colour, from black, diogenous cells, non-septate, 95–140 
pale grey, pale yellowish brown to almost 10–15 mm.
544 THE LICHENOLOGIST Vol. 46

Fig. 2. Palicella glaucopa, variation of apothecial characters; A, concave with strongly pruinose disc (Rodriguez Flakus
2973c); B, flat with evident margin and less pruinose disc (Rodriguez Flakus 2672b); C, convex with reduced margin
(Rodriguez Flakus 2973b); D, epruinose with non-pigmented disc (Rodriguez Flakus 3046); E, exciple composed
of pigmented cortical layer and hyaline interior inspersed by oil droplets and small crystals (Rodriguez Flakus 2499);
F, asci showing amyloid ascus apex (mounted in K/I; Imshaug 39305); G, ascospores (Rodr. Flakus 2672b);
H, conidia (Imshaug 39305). Scales: A ¼ 05 mm; B–D ¼ 1 mm; E ¼ 50 mm; F–H ¼ 10 mm.
2014 Palicella—Rodriguez Flakus & Printzen 545

Chemistry. Atranorin (major), chloroatra- orange or KC--, UV+ dull orange or UV--).
norin (minor), pannarin (major), isonorpan- We assume that the hyaline crystals which
narin (minor), zeorin (major), earthothelin are always observed in the epithecium repre-
(minor), ethiophanic acid (minor), eellagic sent pannarin (P+ rust-orange). The exciple
acid (minor), eseveral unknown terpenes contains droplets of probably undetermined
(minor to major). Thallus K+ yellow, P+ fatty acid (never observed on TLC plates).
yellow, C+ yellowish or orange or C--, KC+ We also observed an additional, ink purple,
yellowish to orange or KC--, UV--; apothecial granular pigment (K+ turquoise) in the exci-
disc K--, P+ rust orange, C+ orange or C--, ple of several specimens. Specimens with this
KC+ orange or KC--, UV+ dull orange or pigment were originally described as Lecidea
UV--. conflectens Nyl. (Nylander 1888). We could
not find any correlation between the occur-
Remarks. The epithet ‘‘glaucopa’’ introduced rence of the additional ink-purple pigment,
by Hook. f. & Taylor (1844) refers to the glau- the presence of xanthones and any morpho-
cous pruina that often covers apothecia (Fig. logical or anatomical characters, and there-
2A & B), but the apothecial colour of P. glau- fore assume that this represents intraspecific
copa is much more variable. Besides typical, variation.
black apothecia with blue-grey pruina, we Palicella glaucopa can hardly be confused
have also observed pure black, shiny and with other lecideoid species from the Leca-
epruinose, pale grey, pale yellowish brown nora varia group, such as L. symmicta (Ach.)
to almost white discs (with black pigment Ach., which is easily distinguished by its
located only in the margin). The shape of yellowish to orange-brown apothecial disc
apothecia and margin is also very variable. with an indistinct margin, an exciple of
Frequently on the same thallus, concave narrowly cylindrical, apically only slightly
apothecia with a thick elevated margin, plane widened hyphae, smaller ascospores and the
apothecia with an indistinct margin and production of usnic acid as a secondary sub-
strongly convex apothecia with a reduced stance (Printzen 2001). Superficially, Clios-
margin can be found. tomum griffithii (Sm.) Coppins resembles P.
Palicella glaucopa can be recognized by the glaucopa because of its similarly variable
characteristic exciple with an outer layer of apothecial colour ranging from pale pink,
strongly swollen and dark pigmented hyphae violaceous grey and brownish to pure black.
(Cinereorufa-green) and a more or less un- It clearly differs by its conspicuous, relatively
pigmented interior composed of narrow, large black (K+ purple) pycnidia, an exciple
loosely arranged hyphae, inspersed by oil composed of hyphae with thick gelatinized
droplets. Its characteristic type of ascus is walls and cylindrical to globose lumina, 1-
somewhat intermediate between the Lecanora- septate ascospores and Biatora-type asci
and Lecidella-type. Palicella glaucopa differs (Ekman 2004). Distinguishing characters of
chemically from P. filamentosa and P. schizo- P. filamentosa and P. schizochromatica are
chromatica by the lack of usnic acid, which is listed in the determination key below.
regularly observed in the other two species.
The distribution of secondary substances in Distribution and ecology. Palicella glaucopa
P. glaucopa is quite interesting. The thallus is known from many localities in southern
contains large quantities of atranorin (K+ Chile and Argentina (Fig. 3), where it is
yellow, P+ yellow) and varying concentra- common on tree trunks and branches in
tions of the xanthones arthothelin and thio- Nothofagus forests.
phanic acid (C+ yellowish to orange or C--, Selected specimens examined (of n ¼ 222). Argentina:
KC+ yellowish to orange or KC--), whereas Prov. Tierra del Fuego, Antártida e Islas del Atlantico Sur:
the apothecial discs always accumulate high Dept. Ushuahia, Tierra del Fuego National Park, near
Rayadito area, alt. 14 m, 54 50 0 26400 S, 68 33 0 22200 W,
concentrations of pannarin (P+ rust-orange) P. Rodriguez Flakus 2423 & A. Flakus (FR); Guanaco
and variable amounts of xanthones (C+ yel- Mountain, alt. 244 m, 54 49 0 11800 S, 68 33 0 29600 W,
lowish or orange or C--, KC+ yellowish to P. Rodriguez Flakus 2504a, 2504b, 2504c, 2504d, 2504e
 546
THE LICHENOLOGIST
Vol. 46
Fig. 3. Distribution map of Palicella species: P. filamentosa (squares), P. glaucopa (circles), P. schizochromatica (triangles). In colour online
2014 Palicella—Rodriguez Flakus & Printzen 547

& A. Flakus (FR); Roca Lake, alt. 2 m, 54 49 0 26800 S, Santesson 8418 (S); Prov. Malleco, Parque Nacional de
68 33 0 58800 W, P. Rodriguez Flakus 2530 & A. Flakus Nahuelbuta, M. Mahu 2646 (FH). Región de Magallanes
(FR); near Ushuahia, alt. 200 m, R. Santesson 452 (S); y de la Antártica Chilena: Prov. Magallanes, Penı́nsula
54 50 0 S, 68 28 0 W, H. Imshaug 55117 & K. Ohlsson de Brunswick, road to Cabo Froward, alt. 5 m,
(UPS); near Antuk Lake, 54 25 0 S, 67 25 0 W, R. Santesson 53 42 0 28700 S, 70 58 0 05600 W, P. Rodriguez Flakus
7070 (S); near Fagnano Lake, R. Santesson 7866 (S); 2617 & A. Flakus (FR); Punta Arenas, P. Dusén s. n.
near San Sebastian, P. Dusén 106 (S); Sierra Sorondo, (H); near Tres Brazos river, alt. 160 m, 53 17 0 S,
alt. 853 m, 41 20 0 24000 S, 71 35 0 12300 W, P. Rodriguez 71 03 0 W, H. Imshaug 49683 & K. Ohlsson (MSC);
Flakus 3098 & A. Flakus (FR); near Sierra Alvear, alt. Seno Otway, bahı́a Camden, H. Imshaug 39144 & R. C.
300 m, 54 43 0 S, 68 01 0 W, H. Imshaug 55217 & K. Harris (MSC); near Mina Rica, alt. 450 m, R. Santesson
Ohlsson (MSC-121067); Isla de los Estados, near Puerto 5273 (S); Fuerte Bulnes, alt. 100 m, K. Kalb 10804 (hb.
Vancouver, alt. 60 m, 54 47 0 S, 64 03 0 W, H. Imshaug K. Kalb); Canal Whiteside, Puerto Yartou, R. Santesson
52035 & K. Ohlsson (FH); near Puerto Hoppner, alt. 90 6851 (S); near Rio Rubens c. 50 km from Natales, R.
m, 54 45 0 S, 64 25 0 W, H. Imshaug 53640 & K. Ohlsson Santesson 5628 (S).
(FH). Prov. Rio Negro: Dept. Bariloche, Nahuel Huapi
National Park, near Cascada de los Duendes, alt. 819
m, 41 10 0 46400 S, 71 24 0 51100 W, P. Rodriguez Flakus Palicella schizochromatica
2976a & A. Flakus (FR); near Tronador Mountain, alt. (Pérez-Ortega et al.) Rodr. Flakus &
871 m, 54 15 0 15000 S, 71 41 0 28900 W, P. Rodriguez
Flakus 3071 & A. Flakus (FR); Llao Llao Municipal
Printzen comb. nov.
Park, alt. 816 m, 41 03 0 02000 S, 71 32 0 39300 W, P. MycoBank No.: MB807640
Rodriguez Flakus 2719 & A. Flakus (FR); Mascardi
Lake, alt. 841 m, 41 20 0 32700 S, 71 34 0 39200 W, P. Lecanora schizochromatica Pérez-Ortega et al., Mycol.
Rodriguez Flakus 3053 & A. Flakus (FR). Prov. Neuquen: Prog. 9: 531 (2010); type: Canada, British Columbia,
Dept. Huiliches, Lanin National Park, near Lanin Purcell Mountains, St. Mary’s Alpine Provincial Park,
Mountain, 39 34 0 45700 S, 71 27 0 14300 W, P. Rodriguez trail to Mortar Lake, 2006, T. Spribille 19965 & V.
Flakus 3277 & A. Flakus (FR); Aluminé Lake, alt. 1400 Wagner (CANL—holotype; FR— isotype).
m, A. Kalela 156f (H).—Chile: Región Aysén: Prov.
Coyhaique, Cerro Castillo National Reserve, road be-
tween El Blanco and Cerro Castillo village, alt. 870 m,
Remarks. Palicella schizochromatica is char-
45 59 0 08700 S, 71 54 0 43200 W, P. Rodriguez Flakus 1748 acterized by yellowish or pale brown to black,
& A. Flakus (FR); Queulat National Park, Sendero lecideine apothecia that are very variable in
Encantado, alt. 474 m, 44 37 0 32900 S, 72 27 0 04100 W, colour, small ascospores, 97-111  35-
P. Rodriguez Flakus 1526 & A. Flakus (FR). Region de 41 mm, and the production of atranorin
la Araucania: Prov. Cautı́n, Huerquehue National Park,
alt. 800 m, 39 09 0 09300 S, 71 42 0 5900 W, P. Rodriguez together with usnic acid and paraensic acids
Flakus 2002 & A. Flakus (FR); Conguillı́o National C and D (Pérez-Ortega et al. 2010).
Park, Captrén región, alt. 1180 m, 38 41 0 36600 S,
71 41 0 25300 W, P. Rodriguez Flakus 2052 & A. Flakus Distribution and ecology. The species occurs
(FR); Prov. Malleco, Malalcahuello National Reserve,
alt. 1359 m, 38 25 0 50500 S, 71 32 0 14600 W, P. Rodriguez on bark and wood of conifers from sea level
Flakus 2100 & A. Flakus (FR); Baie Orange, Hariot s. n., to the timberline (2400 m) in north-western
(UPS-L533353); near Coyhaique bajo, Baquedano, R. North America (Canada and USA).

Key to species of Palicella

1 Ascospores (average of 10) 160–230  55–75 mm; hymenium 50–90 mm tall,
apothecial disc P+ rust-orange (pannarin), thallus without usnic acid; Southern
Hemisphere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. glaucopa
Ascospores (average of 10) 95–145  35–45 mm; hymenium 40–60 mm tall,
apothecial disc P-, thallus usually with usnic acid as accessory substance; Northern
Hemisphere. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2(1) Ascospores on average 120–145  40–45 mm; apothecial margin concolorous or
slightly lighter than disc, young apothecia mostly with lecanorine margin; Europe
and eastern North America . . . . . . . . . . . . . . . . . . . . . . . . . . . P. filamentosa
Ascospores on average 95–115  35–45 mm; apothecial margin usually darker
than disc, often entirely black, biatorine from the beginning; western North America
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. schizochromatica
548 THE LICHENOLOGIST Vol. 46

Discussion An obvious similarity between these species is

the apothecial pigmentation, which may vary
strongly in all three species (Fig. 2; see also
Phylogenetic position and systematic
Printzen & May 2002, fig. 1; Pérez-Ortega
novelties
et al. 2010, fig. 3), especially due to varying
In their study on species of the ‘‘Lecanora concentrations of Cinereorufa-green. The
varia group’’, Pérez-Ortega et al. (2010) re- relatively high number of synonyms of P.
solved two of the three species treated in this filamentosa and P. glaucopa reflects this ex-
study as part of a monophyletic group that treme variability in apothecial colour.
they informally called ‘‘filamentosa group’’. Palicella appears as a sister clade to L.
In order to formalize this distinction, we de- symmicta and Pyrrhospora quernea, which how-
scribe here the new genus Palicella. The ever is only supported by the Bayesian analy-
genus comprises species that were formerly sis. Pérez-Ortega et al. (2010) already inferred
assigned to Lecidea and Lecanora. As tradi- a close relationship between P. filamentosa
tionally circumscribed (Zahlbruckner 1925, and L. symmicta in a phylogenetic analysis in-
1928), these two genera are distinguished volving ITS sequences from more Lecanora
by the presence (in Lecanora) or absence (in species than in this study. The analysis pub-
Lecidea) of a lecanorine amphithecium around lished by Kalb et al. (2008) confirmed a close
the apothecia. Lecanora symmicta and similar relationship between Pyrrhospora and several
species were always difficult to place in this lineages of Lecanora and Lecidella. At pres-
schematic concept, which is reflected by the ent, the phylogenetic position of Palicella
confusing nomenclatural history of the older within Lecanoraceae remains unclear, similar
species of the group. Originally, L. symmicta to that of Miriquidica and Ramboldia (Kalb et
was described as a variety of Lecanora varia al. 2008).
(Acharius 1810), but recombined into Lecidea In order to improve this study, future work
shortly afterwards (Acharius 1814). Zahl- should be based on a broader taxon sampling
bruckner (1928), however, kept it under Le- to clarify the relationships between these
canora. Palicella filamentosa and its numerous genera, as well as Adelolecia, Frutidella, Hae-
synonyms were alternately ascribed to Leca- matomma, Miriquidica, Ramboldia and the
nora (Zahlbruckner 1940; Palice et al. 2011) Tephromelataceae (Spribille et al. 2011). Like-
or Lecidea (Stirton 1879; Hedlund 1892; wise, the correlation between Palicella and
Schmull & Hauck 2005). On the other hand, lecideoid species groups within Lecanora,
P. glaucopa was always treated as a member such as L. fuscescens (Sommerf.) Nyl., L.
of Lecidea s. lat. Kalb (1984) had already ghisleri (Hepp) Arnold, L. boligera (Th. Fr.)
noted that it deviated from typical Lecidea Hedl. and L. anopta Nyl., remains unclear
and recombined it into Lecidella, with which and needs to be tested in a phylogenetic
it shares some similarities, such as a more or framework. Our results, however, indicate
less Lecidella-type ascus, paraphyses that are that even a five-gene dataset might not be
easily separated in KOH and the presence sufficient to robustly infer the backbone of
of Cinereorufa-green in the exciple. The such a phylogeny.
narrowly ellipsoid ascospores and strongly It has been previously suggested that Leci-
thickened excipular hyphae, however, indi- dea hercynica is conspecific with P. filamen-
cate that it does not belong there but should tosa (Palice et al. 2011) and our phylogenetic
rather be assigned to a new genus. tree confirms this proposal with maximal
Our phylogenetic analyses based on five support. A different case is probably that of
gene loci confirmed that P. glaucopa is dis- Lecanora fuscescens, GenBank sequences of
tinct from Lecidella and instead forms a well- which appear firmly embedded in a clade
supported lineage together with P. filamentosa of P. schizochromatica. Based on sequences
and P. schizochromatica within Lecanoraceae. from the same specimen, Schmull et al.
2014 Palicella—Rodriguez Flakus & Printzen 549

(2011) reported Lecanora fuscescens as closely that seems to be typical for Palicella (Fig.
related (sister clade) to ‘‘Lecidea hercynica’’. 2F). Our phylogenetic analysis did not re-
Because of anatomical and chemical differ- solve the relationship between the two gen-
ences between L. fuscescens and P. schizochro- era. Hence it is possible that they are in fact
matica, we believe that its odd position on more closely related to each other than sug-
our tree is due to a misidentification. In L. gested by the phylogeny in Figure 1.
fuscescens, excipular hyphae and paraphyses Morphologically, the Ramboldia elabens
have capitate, pigmented apical cells and group is also similar to Palicella, having apoth-
ascospores are broadly ellipsoid, while P. ecia with dark brownish, matt to glossy discs
schizochromatica has unthickened and unpig- and concolorous margins, as well as a biator-
mented excipular hyphae, mostly colourless ine exciple and narrowly ellipsoid ascospores
and narrow paraphysis tips, and narrowly (Elix 2009). The species differ, however,
ellipsoid ascospores. Furthermore, L. fusces- by typical Lecanora-type asci (tholus with a
cens produces fumarprotocetraric and lobaric strongly amyloid lateral part, and broad, api-
acids, while P. schizochromatica contains atra- cally diverging non-amyloid axial body), pale
norin, usnic and paraensic acids C and D orange pigments inside the exciple (K--), hy-
(Pérez-Ortega et al. 2010). We were unable pothecial granules (K+ purple) and a lack of
to re-examine the voucher on which Gen- atranorin (Hafellner 1993; Kalb et al. 2008;
Bank sequences were based, but according Elix 2009).
to Markus Hauck (pers. comm.) it origi- Cliostomum griffithii may resemble collec-
nated from Montana, where P. schizochroma- tions of P. glaucopa and P. schizochromatica
tica is one of the most common epiphytic on account of the very variable apothecial
lichens (T. Spribille, pers. comm.) and was pigmentation, but is readily distinguished by
collected before the latter species was de- two-celled ascospores, strongly gelatinized
scribed. excipular hyphae with rather wide and short
lumina, the frequent formation of conspicu-
Comparison of Palicella to similar ous black pycnidia reacting K+ purple, and
genera a typical Biatora-type ascus (Ekman 1997,
2004).
Species of Palicella share superficial simi- The phylogenetically closely related Pyr-
larities with species from Cliostomum, Leci- rhospora quernea (Fig. 1) does not resemble
della and Ramboldia, as well as Lecanora sym- any Palicella species and clearly differs by its
micta (Hertel 1983, 1984; Hafellner 1984, farinose to granular-sorediate thallus, dark
1993; Knoph 1990; Kantvilas & Elix 1994; reddish brown apothecia with a reduced
Printzen 2001; Ekman 2004). In particular, margin, the presence of the anthraquinone
P. filamentosa and L. symmicta can be very 7-chloroemodin (K+ reddish purple in sec-
similar in the field. Lecanora symmicta is best tion), and broadly ellipsoid ascospores (Kalb
distinguished by the lack of atranorin, et al. 2008).
slightly shorter ascospores (70–135 mm vs.
10-16 mm in P. filamentosa) and longer coni- The authors would like to thank the curators of the her-
dia (18–25 mm vs. 9–17 mm in P. filamentosa) baria cited above and Prof. Klaus Kalb for the loan of
specimens, Prof. Gernot Vobis (Bariloche) and Reinaldo
(Printzen & May 2002). On first view, the Vargas (Concepción) for their kind hospitality and help
genus Lecidella appears almost identical with with permits and organization of field trips. The first
Palicella glaucopa and P. schizochromatica. It author warmly thanks Dr Holger Thüs (London) for
differs, however, in important anatomical making specimens available for this study and for his
details. The paraphyses are slightly capitate kindness during her visit to BM, and Adam Flakus for
help with the photographic documentation, TLC and
and almost unbranched, and ascospores are fieldwork. Special thanks to Prof. Dr Imke Schmitt and
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Leuckert 2004). Typical Lecidella asci lack and the staff of the Grunelius Möllgaard laboratory,
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550 THE LICHENOLOGIST Vol. 46

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