Fungal Diversity
Taxonomy, phylogeny and biogeography of the lichen genus Peltigera in Papua
New Guinea
Sérusiaux, E.1*, Goffinet, B.2, Miadlikowska, J.3 and Vitikainen, O.4
1
Plant Taxonomy and Conservation Biology Unit, University of Liège, Sart Tilman B22, B-4000 Liège, Belgium
Department of Ecology and Evolutionary Biology, University of Connecticut, 75 North Eagleville Road, Storrs CT
06269-3043 USA
3
Department of Biology, Duke University, Durham, NC 27708-0338, USA
4
Botanical Museum (Mycology), P.O. Box 7, FI-00014 University of Helsinki, Finland
2
Sérusiaux, E., Goffinet, B., Miadlikowska, J. and Vitikainen, O. (2009). Taxonomy, phylogeny and biogeography of the
the lichen genus Peltigera in Papua New Guinea. Fungal Diversity 38: 185-224.
The lichen genus Peltigera is represented in Papua New Guinea by 15 species, including 6 described as new for science:
P. cichoracea, P. didactyla, P. dolichorhiza, P. erioderma, P. extenuata, P. fimbriata sp. nov., P. granulosa sp. nov., P.
koponenii sp. nov., P. montis-wilhelmii sp. nov., P. nana, P. oceanica, P. papuana sp. nov., P. sumatrana, P. ulcerata,
and P. weberi sp. nov. Peltigera macra and P. tereziana var. philippinensis are reduced to synonymy with P. nana,
whereas P. melanocoma is maintained as a species distinct from P. nana pending further studies. The status of several
putative taxa referred to P. dolichorhiza s. lat. in the Sect. Polydactylon remains to be studied on a wider geographical
scale and in the context of P. dolichorhiza and P. neopolydactyla. The phylogenetic affinities of all but one regional
species (P. extenuata) are studied based on inferences from ITS (nrDNA) sequence data, in the context of a broad
taxonomic sampling within the genus. A key to all species is provided and available data on their ecology and general
distribution are discussed. The species P. canina, P. horizontalis, P. laciniata, P. malacea, P. polydactylon and P.
tomentosa are excluded from the lichen checklist of Papua New Guinea. All lichenicolous fungi observed on Peltigera
thalli in Papua New Guinea are listed.
Key words: lectotypification, lichenicolous fungi, The Philippines, Indonesia, Irian Jaya, systematics, ITS
Article Information
Received 16 September 2008
Accepted 16 February 2009
Published online 1 October 2009
*Corresponding author: Emmanuël Sérusiaux; e-mail: E.Serusiaux@ulg.ac.be
Introduction
The genus Peltigera Willd. (Peltigeraceae, lichenized Ascomycetes) is well represented in the mountains of Papua New Guinea,
where it grows on soil, gravel, mossy rocks and
tree trunks and branches. In his Catalogue of
the lichens of Papua New Guinea and Irian
Jaya (Indonesia), Streimann (1986: 96-98)
reports nine taxa for the genus Peltigera: P.
canina (L.) Willd. (with the mention that ‘some
collections may belong to a new species’), P.
dolichorhiza (Nyl.) Nyl., P. erioderma Vain., P.
horizontalis (Huds.) Baumg. (with the mention
that these reports are misidentifications for P.
erioderma), P. laciniata (G. Merr. ex Riddle)
Gyeln. (with the mention that ‘some collections
may belong to a new species’), P. cf. malacea
(Ach.) Funk (with the mention ‘record
doubtful’), P. polydactyla (Neck.) Hoffm.
(with the mention ‘record doubtful’), P. spuria
(Ach.) DC. (a species now named P. didactyla
(With.) J.R. Laundon) and P. tomentosa Vain.
Peltigera ulcerata Müll. Arg. has been added
to that list by Aptroot and Sipman (1991: 232).
Based on preliminary results of the present
study, Martínez et al. (2003) mentioned P.
cichoracea Jatta and P. oceanica Gyeln. from
Papua New Guinea and the Papuan Provinces,
respectively.
The taxonomy of the genus in Papua
New Guinea is poorly understood due mainly
to the application of taxonomic concepts
elaborated in other regions of the world. The
genus Peltigera has been revised for much of
the Northern Hemisphere (e.g. Vitikainen 1994;
185
Goward et al. 1995), and parts of the southern
hemisphere (e.g. Galloway, 2000) but tropical
taxa remain critically understudied. Swinscow
and Krog (1988) offered a treatment of East
African species and Vitikainen (1998) provided
a nomenclatural synopsis and preliminary key
for the Neotropics (Vitikainen, 1998). Few
comprehensive treatments of Asian taxa have
been proposed: Awasthi and Joshi (1982) for
India, and Stenroos et al. (1994) and Xi-Ling
(1986) for parts of China.
Numerous collections of Peltigera are
available from Papua New Guinea, based on
material collected mainly by T. Koponen (H)
and H. Streimann (CANB), and three expeditions organized in the country during the last
20 years (A. Aptroot and H. Sipman in 1987; A.
Aptroot, P. Diederich, E. Sérusiaux and H.
Sipman in 1992; A. Aptroot, P. Lambley, E.
Sérusiaux and H. Sipman in 1995). These
collections form the basis of the present
taxonomic account.
The genus Peltigera has a pioneer status
in modern lichen taxonomy as molecular
techniques were here first used to delimitate
species within a lichen genus: Goffinet and
Miadlikowska (1999) described the new P.
phyllidiosa from S-E United States, using ITS
sequences of rDNA to distinguish it from
related species (P. collina and P. neckeri);
Goward and Goffinet (2000) followed a similar
approach for the new P. chionophila, a
representative of the P. aphthosa-group; and
Goffinet et al. (2003) produced a robust
phylogeny of the P. didactyla-group through
analysis of nrDNA sequences. Furthermore, 38
described species and 8 putative undescribed
taxa were examined in great detail by
Miadlikowska and Lutzoni (2000), using
morphological and chemical characters and
LSU rDNA sequences. Their phylogenetic
analysis led to an infrageneric reorganization of
the whole genus. Finally, Miadlikowska et al.
(2003) examined patterns in morphological
variation in the P. canina-complex against
phylogenetic inferences in ITS and LSU rDNA
sequence including phylogenetic signal
extracted from regions of ambiguous alignment
using INAASE, a method developed by
Lutzoni et al. (2000).
The present study provides the first
taxonomic account of Peltigera in Papua New
186
Guinea, based on a critical study of morphological and chemical characters. We further
sequenced the ITS region (nrDNA) for all but
one regional species and complemented these
with sequences publically available to assess
the monophyletic nature and phylogenetic
affinities of the species from Papua New
Guinea. All taxa were placed and discussed in
the context of the currently accepted infrageneric classification of the genus.
The present study is a part of the
continuing effort towards a better knowledge of
the tremendous lichen biodiversity in tropical
mountains in S-E Asia (Aptroot et al., 2007,
Bjerke and Sipman, 2007, Jørgensen, 2007). A
representative set of the species of Peltigera
will be deposited at Papua New Guinea
National Herbarium in Lae.
Material and methods
This study is based on approximately 430
specimens, and integrates critical morphological, chemical and molecular characters with
field observations by the first author on habitat
and geographical distribution. Ascospores
measurements are given for apothecia sections
prepared in water and only for spores that
escaped the asci following gentle pressure on
such thin sections. For the descriptions and use
of terminology, we follow Vitikainen (1994: 517). Thin-layer-chromatography (TLC) has
been performed for all specimens studied,
following Orange et al. (2001) and using
solvents C and G; the main purpose of this
chemical analysis was to provide further
characters for species delimitation and thus no
effort was made to identify and name accessory
terpenoids.
For DNA extraction, lobes margins or
apothecia were sampled from herbarium
specimens of each species. DNA was extracted
using modified CTAB extraction by Doyle and
Doyle (1987) or the NucleoSpin®Plant kit
from Macherey Nagel (Düren, Germany)
following the manufacturer’s protocol. The
Internal Transcribed Spacer region (ITS1, 5.8S
and ITS2) of the nrDNA repeat was targeted
with the Polymerase Chain Reaction using
either the primers ITS1F (CTTGGTCATTTAG
AGGAASTAA; slightly modified from Gardes
and Bruns 1993 by Piercy-Normore et al. 2006)
Fungal Diversity
or ITS1 (5’TCCGTAGGTGAACCTGCGG3’)
with ITS4 (5’TCCTCCGCTTATTGATATG
C3’, White et al. 1990) directly, or via a nested
approach whereby the region was targeted first
using the primers BMBC-R (5’GTACACA
CCGCCCGTCG3’) and LS4R (5’TCAAGC
ACTCTTTGACTCTC3’, Shaw 2000), with the
product of this initial PCR serving as the
template for a second PCR using primers ITSF
or ITS1 and ITS4. The amplification reaction
was performed in a 25µL volume containing
0.75 units of HotMaster Polymerase Taq
(Eppendorf), 2.5 µL of its buffer, 1 µL of a
10µM solution of the primers, 2.5 mM of each
dNTP solution, and 1 µL of genomic DNA.
PCR products were screened on 1% agarose
gels stained with ethidium bromide, and
purified using NucleoSpin®Extract II kit from
Macherey Nagel (Düren, Germany) following
the manufacturer’s protocol. Sequencing
products were purified using Sephadex G-50
(Amersham) gel filters, and separated by
capillary electrophoresis using the ABI
Prism™ 3100 Genetic Analyzer. Nucleotide
sequences were edited using Sequencher 3.1
(Gene Codes Corporation), entered in
PAUP*version 4.0b10 for Macintosh-PPC
(Swofford, 2002), and manually aligned.
All newly obtained sequences were
submitted to GenBank (see Table 1 for
accession numbers). The beginning and end of
the ITS1 and the ITS2 spacers were determined
by comparison with sequences available from
GenBank. We excluded the 3’ end of the 18S
gene, and the 5’ end of the 26S gene from the
analyses. Sequences for specimens from Papua
New Guinea were complemented with
representtatives of allopatric taxa, for which
ITS sequences were available from GenBank;
their accession number is provided in the
figures (Figs 1-2). Peltigera tereziana and P.
polydactyloides were also targeted for their ITS
sequence (Table 1), to test their affinities to
species from Papua New Guinea.
Internal Transcribed Spacers could not be
readily aligned across all taxa; in fact, two
groups emerged from visual alignment.
Threfore, preliminary phylogenetic inferences
were derived from the 5.8S gene, aligned
across all sequences and analyzed using
maximum parsimony (MP) bootstrapping
(Felsenstein, 1985) with a heuristic search
algorithm on 200 pseudoreplicates each
analyzed 200 times by randomly adding
sequences; a limit of 1,000 trees saved per
pseudoreplicate was imposed. The lineages
recovered corresponded to the groups recognized based on visual alignment of the spacer
regions. Consequently, subsequent analyses
were restricted to sequences belonging to one
or the other group and inferences could then be
made from variation in the sequences of both
spacers.
All sequences were then assigned to
either of the two taxon sets corresponding to
the Sections Peltigera and Polydactylon
+Horizontales (Miadlikowska and Lutzoni,
2000). Phylogenetic relationships within these
groups were reconstructed using P. retifoveata
(Sect. Retifoveata) and members of the P.
neckeri group of Sect. Horizontales as
outgroups, respectively. Regions of ambiguous
alignment across sequences within each of
these two groups were excluded from
phylogenetic analyses. Separate unweighted
maximum parsimony analyses for the Sections
Peltigera and Polydactylon + Horizontales
were performed as follows in PAUP*4.0b10:
an initial run performed by using the “tree
bisection reconnection” (TBR) branch swapping algorithm, with the «steepest descent»
option turned on, and only 10 trees saved for
each of the 200 random addition replicates, was
followed by a second analysis whereby all
saved trees were swapped to completion with
no limit to the number of trees saved. All other
parameters were set to the default options (e.g.,
gaps were treated as missing data). Support for
the branches was estimated using the bootstrap
approach with a heuristic search algorithm on
300 pseudoreplicates each analyzed 2 times by
randomly adding sequences; a limit of 1000
trees saved per pseudoreplicate was imposed.
Bootstrap frequencies (MPBS) were obtained
by constructing a majority rule consensus tree
of all trees saved during the analysis.
For the second set of the MP analyses,
symmetric step matrices were created for
unambiguous portions of the alignments using
the STMatrix 2.1 (Francois Lutzoni and Stefan
Zoller, Department of Biology, Duke University), as outlined in Miadlikowska et al. (2003).
ITS1, ITS2 and 5.8S each were subjected to a
specific symmetric step matrix. Gaps from the
187
Fig. 1. Phylogenetic relationships of species of the Peltigera polydactylon s. lat.-clade (i.e., Peltigera Sect. Horizontales
and Polydactylon sensu Miadlikowska and Lutzoni 2000) occurring in Papua New Guinea, inferred from ITS sequences.
Single most likely tree (-ln = 926.58436). Alternative relationships supported via MP+INAASE analyses are shown to
the right. Support is indicated for branches characterized by posterior probabilities > 0.95, or bootstrap frequencies
exceeding 70% under maximum parsimony (MPB) or maximum likelihood (MLB). Bootstrap support derived from the
MP+INAASE (MPIB) is specified for branches not present in the most-likely tree.
unambiguous portions of the alignments were
treated as a fifth character state. Phylogenetic
signal from ambiguously aligned portions of
the alignments was integrated into maximum
parsimony analyses without violating positional homology, using the program INAASE
2.3b (Lutzoni et al., 2000), a method now
currently used in phylogenetic studies of
lichenized fungi (Amtoft et al., 2008, Gaya et
al., 2008, Reeb et al. 2007). All substitutions
were equally weighted (1.0). In addition, the
188
hypervariable region within the ITS1 (ITS1HR; Miadlikowska et al., 2003) for the Sect.
Peltigera was recoded into 23 characters using
the program arc v1.5 (Kauff et al., 2003;
Miadlikowska et al., 2003) with the nucleotide
option, as outlined in Reeb et al. (2004). Each
of the 23 characters obtained with arcnucleotide were subjected to a specific weight:
1.00 for character 1; 0.25 for characters 2–5;
0.10 for characters 6–15 and 0.50 for characters
16–23. Weighted MP analyses with recoded
Fungal Diversity
Fig. 2. Phylogenetic relationships of species of the Peltigera canina-clade (i.e., Peltigera Sect. Peltigera sensu
Miadlikowska and Lutzoni, 2000) occurring in Papua New Guinea inferred from ITS sequences. Single most likely tree
(-ln = 1538.2922). Alternative relationships supported via MP+INAASE analyses are shown to the right. Support is
indicated for branches characterized by posterior probabilities > 0.95, or bootstrap frequencies exceeding 70% under
maximum parsimony (MPB) or maximum likelihood (MLB). Bootstrap support derived from the MP+ INAASE (MPIB)
is specified for branches not present in the most-likely tree.
189
Table 1. Voucher information for specimens of Peltigera for which the ITS region was sequenced.
Sequences are identified by their GenBank accession numbers. All vouchers from Papua New
Guinea and deposited in LG unless otherwise indicated.
Taxon
Voucher
GenBank
P. cichoracea 1
P. cichoracea 2
P. cichoracea 3
P. didactyla
P. dolichorhiza s. lat. 1
P. dolichorhiza s. lat. 2
P. dolichorhiza s. lat. 3
P. dolichorhiza s. lat. 4
P. dolichorhiza s. lat. 5
P. dolichorhiza s. lat. 6
P. dolichorhiza s. lat. 7
P. dolichorhiza s. lat. 8
P. dolichorhiza s. lat. 9
P. dolichorhiza s. lat. 10
P. dolichorhiza s. lat. 11
P. dolichorhiza s. lat. 12
P. dolichorhiza s. lat. 13
P. dolichorhiza s. lat. 14
P. erioderma 1
P. erioderma 2
P. erioderma 3
P. fimbriata 1
P. fimbriata 2
P. fimbriata 3
P. granulosa 1
P. granulosa 2
P. granulosa 3
P. granulosa 4
P. granulosa 5
P. koponenii 1
P. koponenii 2
P. koponenii 3
P. montis-wilhelmii 1
P. montis-wilhelmii 2
P. montis-wilhelmii 3
P. nana 1
P. nana 2
P. oceanica 1
P. oceanica 2
P. oceanica 3
P. papuana 1
P. papuana 2
P. papuana 3
P. papuana 4
P. papuana 5
P. papuana 6
P. papuana 7
P. papuana 8
P. polydactyloides
P. sumatrana 1
P. sumatrana 2
P. sumatrana 3
P. sumatrana 4
P. sumatrana 5
P. tereziana 1
Huon peninsula, July 1992, E. Sérusiaux s. n.
Myola, E. Sérusiaux 15305
Mt Wilhelm, E. Sérusiaux 13983
Mt Sarawaket, T. Koponen 32182
Myola, H. Sipman 38337 (B)
Myola, October 1995, E. Sérusiaux s. n. (sample 1)
Myola, October 1995, E. Sérusiaux s. n. (sample 2)
Bundi gap, P. Diederich 11052 (hb Diederich)
Myola, October 1995, E. Sérusiaux s. n. (sample 3)
Gahavisuka, P. Diederich 10582 (hb Diederich)
Teptep, T. Koponen 34738 (H)
Teptep, 30 July 1992, E. Sérusiaux s. n.
Teptep, T. Koponen 34691 (H)
Gahavisuka, 5 November 1995, E. Sérusiaux s. n.
Myola, H. Sipman 38261 (B)
Gahavisuka, H. Sipman 22199 (B)
Myola, October 1995, E. Sérusiaux s. n. (sample 4)
Saruwaged Range, H. Sipman 24434 (B)
Myola, October 1995, E. Sérusiaux s. n. (sample 1)
Myola, October 1995, E. Sérusiaux s. n. (sample 2)
Mt Wilhelm area, E. Sérusiaux 14107
Mt Sarawaket, T. Koponen 32996 (isotype)
Teptep, T. Koponen 33912 (H)
Mt Wilhelm, A. Aptroot 33080 (hb Aptroot)
Bundi, November 1995, E. Sérusiaux s. n. (sample 1)
NE of Mt Sarawaket, T. Koponen 33267 (isotype)
Myola, E. Sérusiaux 15150
Bundi, H. Sipman 39313 (B)
Bundi, November 1995, E. Sérusiaux s. n. (sample 2)
Lake Wamba, T. Koponen 34327 (isotype)
Myola, October 1995, E. Sérusiaux s. n.
Oksapmin, L. Hoffman 90-103
Mt Wilhelm, E. Sérusiaux 13984 (holotype)
Mt Wilhelm, D. McVean 66236 (COLO)
Goroka-Gembolg, August 1992, E. Sérusiaux s. n.
Bundi, November 1995, E. Sérusiaux s. n.
Mt Wilhelm, E. Sérusiaux 13981
Myola, October 1995, E. Sérusiaux s. n.
Onim Station, A. Bellamy 1295 (B)
Mt Sarawaket, T. Koponen 31965
Teptep, E. Sérusiaux 13656 (holotype)
Mt Wilhelm area, E. Sérusiaux 14106
Mt Sarawaket, T. Koponen 31592 (H)
Mt Gahavisuka, P. Diederich 10581 (hb. Diederich)
Myola, H. Sipman 38336 (B) (sample 1)
Teptep, E. Sérusiaux 13655
Myola, H. Sipman 38336 (B) (sample 2)
Teptep, H. Sipman 35270 (B)
KENYA, Mt Kenya, September 2002, D. Killmann (KOBL)
Gahavisuka, P. Diederich 10585 (hb Diederich)
Mt Wilhelm, E. Sérusiaux 13985
Gahavisuka, November 1995, E. Sérusiaux s. n.
Bundi, E. Sérusiaux 16402
Saruwaged Range, H. Sipman 24513 (B)
AUSTRALIA, NSW, T. K. Overeem 11 (CANB)
FJ527278
FJ527277
FJ527276
FJ527258
FJ527295
FJ527301
FJ527302
FJ527294
FJ527292
FJ527293
FJ527296
FJ527297
FJ527298
FJ527299
FJ527300
FJ527303
FJ527279
FJ527280
FJ527263
FJ527264
FJ527265
FJ527272
FJ527273
FJ527274
FJ527253
FJ527254
FJ527255
FJ527256
FJ527257
FJ527269
FJ527270
FJ527271
FJ527260
FJ527261
FJ527262
FJ527282
FJ527286
FJ527283
FJ527284
FJ527285
FJ527246
FJ527247
FJ527245
FJ527249
FJ527248
FJ527252
FJ527251
FJ527250
FJ527275
FJ527288
FJ527289
FJ527290
FJ527291
FJ527287
FJ527266
190
Fungal Diversity
Table 1 (continued). Voucher information for specimens of Peltigera for which the ITS region was
sequenced. Sequences are identified by their GenBank accession numbers. All vouchers from Papua
New Guinea and deposited in LG unless otherwise indicated.
Taxon
Voucher
GenBank
P. tereziana 2
P. tereziana 3
P. ulcerata
P. weberi
AUSTRALIA, NSW, J. A. Elix 24690 (CANB)
AUSTRALIA, NSW, H. Streimann 63484 (CANB)
Gahavisuka, E. Sérusiaux 16200
Bundi, H. Sipman 39314 (B)
FJ527267
FJ527268
FJ527259
FJ527281
INAASE characters (for the Sect. Peltigera and
Polydactylon+Horizontales) and arc characters
(for the Sect. Peltigera only) were performed
as heuristic searches with 1000 randomaddition-sequence replicates, TBR branch
swapping, MulTrees option in effect, saving all
trees and collapsing branches with maximum
branch length equal to zero. Hereafter, these
weighted MP analyses will be referred to as
MP+INAASE. Branch support was assessed by
MP bootstrap analyses with full heuristic
searches, 1000 replicates using two random
addition sequence (RAS) per bootstrap replicate and by saving all trees. In both bootstrap
analyses, the same parameters as in the original
MP search were used, and constant sites were
excluded from all the analyses.
The optimal model of sequence evolution
for each matrix was identified using MrModeltest 2 (Nylander, 2004) based on the Akaike
Information Criterion (Posada and Buckley
2004). The model that was selected for the
Sections Polydactylon+Horizontales corresponds to the K80 (Kimura, 1980) + I model,
which was implemented as follows: transition/transversion ratio = 2.0140; base
frequencies are equal; proportion of invariable
sites = 0.5875. The model that was selected for
the Sect. Peltigera corresponds to the SYM
(Zharkikh, 1994) + G model. The substitution
rates were fixed to: 1.1668 (A-C), 4.5587 (AG), 1.6528 (A-T), 0.2951 (C-G) and 4.5772 (CT); the proportion of invariable sites was set to
zero and the gamma distribution shape
parameter to 0.2939. Maximum likelihood (ML)
search was performed using PAUP*4.0b10.
invoking the «steepest descent» option and
using the «TBR branch swapping» algorithm.
The search strategy implemented for the
bootstrap search was the same as that used in
unweighted MP: 300 pseudoreplicates, each
with two random additional replicates, TBR
and a maximum of 1000 trees saved per
replicate. Trees saved during the bootstrap
search were executed in PAUP and a 70%
majority-rule consensus tree built. Bootstrap
frequencies were considered significant if
higher than 70% (Hillis and Bull, 1993; Reeb
et al., 2004). Likelihood trees were sampled
from the tree space using a Bayesian approach
(MrBayes v.3.1.2; Ronquist and Huelsenbeck,
2003) using three heated and one cold chains.
The data were treated as a single partition. A
single tree was saved to a tree file every 50
generations for a total of 106 generations. Of
the 20,001 trees that were saved, the first 1001
(the “burnin”) were ignored for determining
posterior probabilities and confidence intervals
for model parameters. The Bayesian analyses
were repeated once to allow for the presence of
multiple local optima. The trees that were
sampled during both runs were ultimately
combined and parameter and posteriors were
estimated based on this set of 36,000 trees.
Posterior probabilities (PP) for bipartitions,
drawn from the 95% consensus tree were
considered statistically significant when the P ≥
0.95.
Both final datasets for the P. polydactylon s. lat.-group and the P. canina-group are
deposited in TreeBASE.
Results
Phylogenetic analysis of the ITS sequences
The ITS region was successfully
sequenced for 60 specimens including three
accessions of the Autralasian P. tereziana and
one accession of the African P. polydactyloides
(Table 1). DNA suitable for the amplification
of the ITS region could not be obtained for
Papua New Guinea samples of P. extenuata,
additional samples of P. erioderma, P. fimbriata, P. ulcerata, and P. weberi. Incomplete
191
sequences were obtained for P. cichoracea 3
and P. sumatrana 1 (each missing about 150
nucleotides [nts] in the ITS1), and P.
dolichorhiza s. lat. 5, for which the ITS1 is
missing the first 12 nts. For all three accessions
of P. fimbriata only the 5.8S and the ITS2
region could be sequenced.
Alignment of the sequences led to
multiple regions of ambiguous homology. An
analysis of the 5.8S gene segregated the
sequences into two groups, corresponding to
Sect. Peltigera (herefter named the P. caninaclade) and Sect. Horizontales+Polydactylon
(hereafter named the P. polydactylon s. lat.clade), respectively. Within these two clades
the spacer sequences aligned more readily
(although large regions of ambiguous alignment remained), and consequently analyses
were conducted on these groups separately.
Overall, sequence variation within the
ITS region permits to circumscribe the Sections
of Peltigera and various species complexes
within them, but fails to provide strong
evidence for the monophyly of various species
that seem well-defined on morphological and
chemical grounds. The ITS regions are characterized by significant local length variations.
The pattern is often congruent with species
delimitations and this putative phylogenetic
signal is indeed recovered when integrating
these regions in maximum parsimony analysis
as coded characters. Peltigera fimbriata and P.
montis-wilhelmii are well-defined by the ITS1HR region, which is unique for each putative
species. High intraspecific variation in the
ITS1-HR region within P. papuana may
indicate presence of multiple unrecognized
species. For example, P. papuana 2 & 3 share
an almost identical ITS1-HR pattern that
differs from the remaining specimens of P.
papuana. In the case of P. koponenii the
pattern in the ITS1-HR region is identical to
that observed in P. canina specimens included
in this study, but the species is defined by two
single nucleotide changes (one substitution and
one guanosine insertion). Phylogenetic signal
recovered from ambiguous regions of the
alignment thus generally permits to enhance
resolution within species complexes, and never
significantly contradicts phylogenetic relationships derived from the alignable portions of the
spacers, as already shown in previous phyloge192
netic reconstructions within Peltigera (Miadlikowska et al., 2003; Goffinet et al., 2003) or
other genera (Pseudocyphellaria: Miadlikowska et al., 2002; Teloschistaceae: Gaya et
al., 2003, 2008; Sticta: McDonald et al., 2003;
polysporous genera: Reeb et al., 2004).
Several morphologically well-defined
taxa failed to be resolved as monophyletic
entities based on ITS. Examples include P.
evansiana, an Eastern North American-Eastern
Asian taxon diagnosed from all its congeners
by the small granular laminal isidia or P.
elisabethae, a circumboreal species differing
from its closest relatives by the schizidia. With
regard to the Papua New Guinea Peltigera,
several morphotaxa lack support for their
monophyly, such as P. koponenii, P. montiswilhelmii, or P. granulosa. However, inferences from ITS fail to provide strong support
against the hypothesis of monophyly for these
taxa. This lack of a robust resolution despite
morphological differentiation may merely
reveal broad morphological amplitude or be
indicative of recent or active speciation events.
Recent cladogenic events can only be recovered within a phylogenetic scenario through
the extensive sampling of fast evolving loci
(Knowles and Carstens, 2007). At present such
loci have not yet been identified for Peltigera,
and ITS remains a main source of information
to address the phylogenetic significance of
morphological and chemical characters in this
genus.
The Peltigera polydactylon s. lat. clade (Fig. 1)
Within this clade, the ITS1 varied
between 185 in P. phyllidiosa and 237 nts in P.
dolichorhiza s. lat. 5, 6, 9, 10 & 11. The
alignment of the sequences resulted in a matrix
of 757 characters of which 439 ambiguously
aligned characters were excluded from
heuristic searches for unweighted MP and ML
analyses. Of the 318 characters included in the
analyses, 248 were constant, and 59 potentially
informative under the parsimony criterion.
Twenty-two regions of ambiguous alignment
were recoded using INAASE and included in
the weighted MP analyses.
The heuristic search based on the
nucleotide characters only yielded a set of six
equally parsimonious trees of length 89 (CI =
0.9101, CI-auto = 0.8947; RI = 0.9865, RC =
Fungal Diversity
0.8978; trees not shown). Inferences under ML
using PAUP converged on a single tree (-ln=
926.58436) that is topologically congruent with
those obtained under MP. The monophyly of
Sect. Polydactylon and that of its two main
clades is well supported. Extracting phylogenetic signal from regions of ambiguous
alignment (INAASE characters) provided more
resolution and support for many morphospecies
and phylogenetic relationships among them.
Peltigera cichoracea is placed within the
P. horizontalis group (MPBS=100%; MPIBS
=95% ; MLBS=100%; PP=1.00). Differentiation of the species based on ITS data is weak,
consisting primarily in short, mostly single
nucleotide indels. The monophyly of P. cichoracea and of P. elisabethae is confirmed when
INAASE characters were included in the MP
analysis (MIPBS=74% and 87%, respectively).
Within the well-supported Sect. Polydactylon, two main lineages are resolved. The first
clade (hereafter named Clade 1) is composed of
P. nana, P. oceanica, P. sumatrana and P.
weberi, and two specimens referred to P.
dolichorhiza s. lat. D. Validity of all sequences
from Clade 1 have been confirmed via new
extractions, amplifications and sequencing.
This combined lineage is well supported in
terms of bootstrap percentages (MPBS=84% ;
MLBS=88%) but fails to gather strong support
from the Bayesian (PP=0.92%) and MP+
INAASE analyses (MPIBS=52%). The two
P. dolichorhiza specimens share identical
sequences except of a dinucleotide duplication
in the ITS1 of P. dolichorhiza 13. They share
only indels as synapomorphies and hence they
form a well-supported monophyletic clade only
when indels are scored 1.00 (MPIBS=100%).
None of the analyses revealed P. sumatrana as
monophyletic. The five accessions share few if
any synapomorphies, unlike their relatives. The
monophyly of the clade combining P. nana, P.
oceanica, and P. weberi is well supported
(MPIBS=98% ; MLBS=78% ; PP=0.99) except
under unweighted MP when indels are not
scored (MPBS=66%). Within this group, only
P. oceanica is represented by several
specimens collected in different localities.
Monophyly of P. oceanica is well supported in
the MP+INAASE analysis (MPIBS=91%). The
relationships among P. nana, P. oceanica, P.
sumatrana and P. weberi are ambiguous. The
phylogenetic structure is congruent with
patterns of indel distribution: the clade 1 is
defined among others by lacking a 25 nts long
region in the ITS2; P. sumatrana differs from
its relatives by a single versus a double TAA
repeat, a GATA insertion and an 11 nts
deletion in the ITS2 compared to other
members of the clade.
The second clade (MPBS=98%; MLBS=
94%; MPIBS= 100%; PP= 1.00) comprises
mainly specimens that are morphologically
identified as P. dolichorhiza and two GenBank
accessions filed as P. dolichorhiza and the
bipolar P. neopolydactyla (both specimens
were collected in New Zealand; Thomas et al.,
2002). This clade is hereafter referred to as
Clade 2. The sequences within this group are
identical except for a three nucleotide duplication in the ITS1 shared by five accessions, and
an additonal T in a poly-T region in the ITS2.
The pattern of sequence variation among the
specimens from Papua New Guinea included in
Clade 2 matches that of the chemical variation.
Indeed, three chemotypes are recognized: one
is characterized by the production of peltidactylin and zeorin and is represented by a single
accession (P. dolichorhiza s. lat. 4); the second
one is represented by eight accessions (P.
dolichorhiza s. lat. 5-12), all with the unique
chemical pattern of four unknown terpenes
together with methyl gyrophorate, tenuiorin,
methyl lecanorate, methyl evernate, and methyl
orsellinate; the third group of three specimens
(P. dolichorhiza s. lat. 1-3) represents samples
from the same locality and is characterized by
the production of zeorin, and rather short,
fasciculate or densely branched rhizines (vs
long and simple for the other accessions of
Clade 2).
It is worth mentioning that both clades
are morphologically variable but that differenttiation between them is lacking. The only
diagnostic character seems to be the presence
or absence of dolichorrhizin: abundantly
produced in Clade 1, and never (or not clearly
detected) in Clade 2. However, this diagnostic
character vanishes when sequences from
specimens from Rwanda (Africa) morphologically identified as P. dolichorhiza are included
(Goffinet and Sérusiaux, unpubl. results): they
produce dolichorrhizin but unambiguously fall
within Clade 2.
193
The importance of chemistry within the
Sect. Polydactylon was first detected by
Miadlikowska and Lutzoni (2000), who even
provided a key to the chemotypes present.
They described a pattern (Fig. 10 in Miadlikowska and Lutzoni, 2000: 947) which is
present in the material from Papua New Guinea:
a chemotype with the three main terpenes
present (peltidactylin, dolichorrhizin and zeorin)
and others with one or two missing. A similar
variation was earlier described by HoltanHartwig (1993: 58) for P. neopolydactyla: two
chemotypes with the three main terpenes
present (the chemotypes being differentiated by
the occurrence of secondary ones), a
chemotype without dolichorrhizin and one with
zeorin only. Furthermore chemotypes were
correlated with morphological variation,
including the morphotype C with «thick, bushshaped to slightly branched rhizines »
associated with the chemotype producing only
zeorin (plus a secondary one). Interestingly,
P. dolichorhiza s. lat. 1-3 could match such a
description.
The delimitation between P. neopolydactyla and P. dolichorhiza remains ambiguous.
Peltigera dolichorhiza is traditionally considered to be restricted to the southern Hemisphere
with some extension into subtropical areas
North of the Equator (Ahti and Vitikainen,
1977: 93), and has on that basis been excluded
from temperate and boreal floras (ex.: Goffinet
and Hastings, 1994: 36-37). Peltigera neopolydactyla is by contrast thought to be primarily
(but not exclusively) circumboreal in distribution. Where the species are sympatric they
are distinguished on subtle tinge differences of
both sides of the thallus (Neotropics:
Vitikainen, 1998) or on the size and branching
of the rhizines (New Zealand: Galloway, 2000).
Peltigera neopolydactyla has been extensively
studied in Europe and Canada, where it is
morphologically and chemically variable
(Holtan-Hartwig, 1993: 57-62; Vitikainen,
1994: 67-69; Goward et al., 1995: 107-108).
The ITS was sequenced for both species
by Thomas et al. (2002), on material from New
Zealand and both samples are resolved within
the clade 2. We have not examined the
vouchers and it is possible that both are in fact
P. neopolydactyla or that they represent a
distinct taxon. Similarly, the whole Clade 2
194
could be interpreted as representing P. neopolydactyla or one or more distinct taxa. At
present this ambiguity cannot be resolved. This
putative complex of broadly lobed glabrous
Peltigera species is in need of a critical
taxonomic and phylogenetic revision. Pending
further research on Clades 1 and 2, we do not
give any status to the four chemotypes
recognized as P. dolichorhiza.
The Peltigera canina clade (Fig. 2)
Within this group, the ITS1 varied
between 162 in P. rufescens and 290 nts in P.
membranacea. For P. montis-wilhelmii 2 the
sequence is incomplete, and lacks about 20 nts
at the 3’ end. The alignment of the sequences
that vary in length between 506 and 646
resulted in a matrix of 768 characters. Of these
337 were included in the analysis. Of these,
236 are constant, and 82 are potentially
phylogenetically informative under parsimony.
The largest portion of excluded characters
belongs to the ITS1, and is composed primarily
of the hypervariable region sensu Miadlikowska et al. (2003). Fifteen regions of
ambiguous alignment were recoded using
INAASE and included in the weighted MP
analyses together with 23 are characters
derived from the ITS1-HR region.
The heuristic search under maximum
parsimony with nucleotide characters only
yielded 30 optimal trees of 184 steps (CI =
0.6793, CI-auto = 0.6424; RI = 0.8865, RC =
0.6023). The nodes of the backbone are
characterized by low support, and the
phylogenetic signal carried by the aligned
portions of the sequences is confined to the
monophyly of individual species or closely
related species. Analyses under the criterion of
maximum likelihood yielded a single tree (-ln
= 1538.2922). Bootstrap support (i.e.
frequencies >70%) under maximum likelihood
is congruent with maximum parsimony support.
Posterior probabilities equal or greater than
0.95 are also confined to the monophyly of
species or of closely related taxa (i.e. P. canina
s. lat.). The addition of recoded characters
(INAASE + arc) provided additional bootstrap
support for species or species complexes such
as P. granulosa, P. koponenii, P. montiswilhelmii. The only taxon for which the support
was always lacking is P. papuana.
Fungal Diversity
Peltigera
erioderma
(MPBS=83%;
MPIBS=93%; MLBS=80%; PP=1.00) is the
sister-group to a clade composed of three
accessions assigned to the newly described P.
montis-wilhelmii (MPIBS=86%). These two
taxa differ in their signature sequence in one
variable region, with P. montis-wilhelmii
exhibiting slight infraspecific variability. The
shared most recent common ancestry is well
supported
(MPBS=97%;
MPIBS=100%;
MLBS=95%; PP=1.00), as well as by two
unique insertions (2 and 1 nt) in the 5.8S gene.
The P. canina-complex comprises another set
of specimens from Papua New Guinea,
representing the morphologically distinct P.
koponenii, represented by sequences virtually
identical to those of P. canina (including the
ITS1-HR region) except for one substitution
and one single nucleotide insertion in
unambiguous portion of the alignment.
Peltigera koponenii was reconstructed as
monophyletic and highly supported (MPIBS
=94%) when phylogenetic signal from
ambiguous regions was incorporated into
phylogenetic analyses (MP with recoded
INAASE and arc characters).
The three accessions of P. fimbriata form
a well-supported monophyletic group that may
be closely related to the common ancestor to P.
ponojensis and P. monticola, but such affinity
remains ambiguous.
The remaining accessions from Papua
New Guinea belong to a clade related to the
cosmopolitan P. rufescens. The monophyly of
P. granulosa is well supported only by MP
with recoded INAASE and arc characters
(MPIBS=91%). The monophyly of P. papuana
lacks support.
Peltigera tereziana from Australia represents a well-supported monophyletic group,
closely related to P. degenii and P. membranacea, but of ambiguous affinities to either one
of these.
Key to species
1a
1b
2a
Soredia or soredioid masses present, at least in parts
of the thalli ............................................................ 2
Soredia or soredioid masses absent ....................... 7
Upper surface tomentose, at least in parts; soralia
mostly laminal, or submarginal on old thalli......... 3
2b
Upper surface glabrous; soralia laminal,
submarginal or marginal........................................ 4
3a
Rhizines abundant, densely branched to fibrillose;
young soralia C+ red ...........................P. extenuata
Rhizines sparse, simple to loosely branched; soralia
C-.......................................................... P. didactyla
3b
4a
4b
G enuine soralia only ............................................ 5
Genuine soralia present but always with soredioid
or isidioid masses, or with granules or phyllidia ... 6
5a
Soralia mostly laminal, or submarginal and
marginal; terpenoids absent................... P. ulcerata
Soralia strictly marginal; terpenoids present ..........
................................................................. P. weberi
5b
6a
6b
Margins disrupted into soralia, soredioid or isidioid
masses, or with granules or phyllidia; terpenoids
present with zeorin always abundantly produced;
thalli usually quite large, reaching 10 cm in diam...
..........................................................P. cichoracea
Margins with soralia or isidioid masses, when welldeveloped also present on the edge of the lower
surface; terpenoids absent; thalli usually quite small,
not exceeding 2-5 cm in diam. ........... P. granulosa
7a
7b
Marginal lobules or phyllidia present ................... 8
Marginal lobules or phyllidia absent .................... 9
8a
Upper surface smooth or faintly to distinctly
scabrous, sometimes with a whitish pruina near the
lobes margins resulting in a frosted appearance,
lobes carrying the apothecia usually tomentose;
marginal phyllidia usually abundant, but sometimes
few; lower surface with a conspicuous network of
usually dark, raised veins; rhizines mostly
threadlike or penicillate; apothecia typically
horizontal; terpenoids absent ............... P. papuana
Upper surface smooth, rarely with a whitish pruina
on lobes margins; marginal phyllidia rare (on lateral
lobes or on damaged parts); lower surface usually
lacking distinct veins; rhizines typically fasciculate
and densely branched, remaining isolate and
arranged in concentrical rows or forming dense
cushions under the thallus; apothecia typically
saddle-shaped; terpenoids present ...........................
...................................... P. sumatrana (rare forms)
8b
9a
9b
Upper surface tomentose, at least in parts;
terpenoids absent................................................. 10
Upper surface glabrous or pruinose (in parts or at
the extremities of young lobes); terpenoids present
or absent .............................................................. 14
10a Rhizines usually very abundant and forming dense
‘bushy’ masses, typically fibrillose to coralloid;
apothecia saddle-shaped..................... P. koponenii
10b Rhizines abundant or not, simple, threadlike,
penicillate or fasciculate, never fibrillose to
coralloid; apothecia horizontal or saddle-shaped ....
............................................................................ 11
195
11a Apothecia typically horizontal; pycnidia absent, or
rare and inconspicuous; tomentum covering large
parts of the thallus, or restricted to lobes carrying
the apothecia........................................................ 12
11b Apothecia typically saddle-shaped; pycnidia
usually present, at least on some lobes margins,
conspicuous (0.3-0.8 mm in diam.); tomentum of
the upper surface most usually well-developed, but
sometimes absent in parts of the thallus, with long
and conspicuous (to 1-1.4 mm long) hairs or not ....
............................................................................. 13
12a Thallus large, to 15 cm in diam., lobes typically
rounded at their extremities and 1-1.5 cm wide,
margin revolute; tomentum present on most of the
upper surface made of tiny whitish hairs (0.1-0.15
mm high) out of which longer ones (0.1-0.4 mm
long) may emerge...............................P. erioderma
12b Thallus forming attractive, small rounded rosettes
(3-6 cm in diam. in suitable conditions); lobes
imbricate, not exceeding 0.5-0.7 cm wide, with
raised and crenate (sometimes +/- crisped) margins;
tomentum usually restricted to lobes carrying
apothecia but sometimes covering large parts of the
thallus surface................... P. papuana (rare forms)
13a Tomentum of the upper surface sometimes almost
absent in parts of the thallus but always, and
especially near the lobes margins, with long (1-1.4
mm long) whitish to brownish hairs, sometimes
aggregated in tufts; lobes rounded; rhizines not
typically arranged in rows on the veins of young
lobes .................. P. fimbriata (typical populations)
13b Tomentum of the upper surface identical but with
shorter hairs, sometimes quite few and hardly
distinct from the tomentum, or even absent; lobes
rather elongated and narrow; rhizines typically
arranged in rows on the veins (especially on young
lobes) ................P. fimbriata (atypical populations)
14a Apothecia typically horizontal; terpenoids absent...
................................................. P. montis-wilhelmii
14b Apothecia saddle-shaped or absent; terpenoids
present ................................................................. 15
15a Tenuiorin and methylgyrophorate absent but
terpenoids present; upper surface smooth,
sometimes slightly pruinose at the margins or
incrusted; lobes with raised and crisped lateral
margins................................................. P. oceanica
15b Tenuiorin and methylgyrophorate always produced,
together with terpenoids ...................................... 16
16a Rhizines abundant, or rarely sparse, rather short,
fasciculate or densely branched........................... 17
16b Rhizines sparse or abundant, most usually long and
simple .................................................................. 19
17a Terpenoids present, mostly zeorin or peltidactylin
but dolichorrhizin not produced ..............................
...........................................P. dolichorhiza s. lat. B
17b Terpenoids present, dolichorrhizin abundantly
produced.............................................................. 18
196
18a Thallus rather fragile and thin, with a +/undulating surface ...................................................
...................... P. sumatrana (atypical populations)
18b Thallus robust, usually with large lobes; surface
never or hardly undulating ......................................
........................ P. sumatrana (typical populations)
19a Thallus robust, not distinctly undulating; lower
surface with a dense network of unraised, dark
veins with distinct rather small, elliptical, whitish
interstices; upper surface usually with pruina at the
lobes margins and sometimes with large and
conspicuous incrusted patches; dolichorrhizin as the
main terpenoid produced; terricolous......... P. nana
19b Thallus fragile, with a distinctly undulating surface
with shallow depressions, at least in well-developed
specimens; lower surface with a loose network of
raised, pale to dark veins with large, ellipitical,
whitish to pale orange interstices; terpenoids
produced in several chemotypes; mostly epiphytic,
rarely on rotten wood or on terricolous mosses... 20
20a Dolichorrhizin abundantly produced.......................
......................................... P. dolichorhiza s. lat. D
20b Dolichorrhizin not detected................................. 21
21a Terpenoids include zeorin and peldidactylin...........
......................................... P. dolichorhiza s. lat. A
21b Terpenoids unknown (four produced) including a
green and a red spot on TLC plates .........................
......................................... P. dolichorhiza s. lat. C
Phytogeography
Because of its wide range of wellpreserved habitats, the island of New Guinea
(Irian Jaya and Papua New Guinea) is expected
to be a hotspot for lichen biodiversity (Sipman
and Aptroot, 2006). We nevertheless consider
the total number of Peltigera species present in
Papua New Guinea as rather low (15 species
accepted in this survey), especially when
compared with the higher number of species in
closely related genera such as Pseudocyphellaria (25 species mentioned from Papua
New Guinea in Galloway, 1994), and with the
number of Peltigera species reported from
other parts of the world. For example 20
species of Peltigera are reported from Belgium
and Luxembourg (Sérusiaux et al., 2004), 29
from the European continent (Vitikainen, 1994),
and 28 species (plus several putative ones)
from British Columbia/Canada (Goward et al.,
1995, Miadlikowska and Lutzoni, 2000). We
could not find any collections of Peltigera
canina, P. horizontalis, P. laciniata, P. malacea, P. polydactylon and P. tomentosa, all
species reported from New Guinea by Strei-
Fungal Diversity
mann (1986: 96-98), and all are thus excluded
from the flora of the island.
Martínez et al. (2003) summarized
geographical distribution patterns of the genus
throughout the world and commented on the
paucity of records in S-E Asia and Australia. It
is thus quite difficult to assess the endemism
level reached by the genus on the whole island
of New Guinea. Almost no data are available
for nearby tropical mountains, especially in
Indonesia and the Philippines, but it can be
expected that several species newly described
here have a wider distribution in those parts of
the world. This is clearly suspected with the
resurrection of several epithets introduced by
Vainio or Gyelnik for species found in Papua
New Guinea and that were virtually unknown
beside their type collections: P. erioderma
Vain., P. nana Vain., P. oceanica Gyeln. (all
three described from Luzon/The Philippines)
and P. sumatrana Gyeln. (described from
Sumatra/Indonesia).
Our current understanding of the
taxonomy and distribution of species leads,
however, to the following observations: (1) no
species of Peltigera is strictly pantropical;
widespread species seem to be subcosmopolitan (e.g. P. didactyla) or if pantropical then
they are also present in some temperate areas
of either hemispheres (e.g. P. ulcerata); (2) no
species is disjunct between the Neotropics and
Papua New Guinea, whereas at least one
species (e.g. P. cichoracea) could be broadly
paleotropical as it is known in East Africa,
Taiwan and Papua New Guinea; (3) genuine
endemic species most probably exist, such as
the very typical and conspicuous P. fimbriata;
(4) putative endemic species (e.g. P. koponenii)
likely originated from rather recent cladogenic
events as suggested from the high ITS
sequence similarity with their closest relative.
The current data suggest that none of the
species of Peltigera on the island are relictual
species (i.e., paleoendemics).
The species
Peltigera cichoracea Jatta, Nuov. Giorn. Botan.
Ital. 14: 170, 1882.
(Figs 3 A-B)
Type: ‘Africa, [Ethiopia] Scioa, inter
muscos’ (? FI, type not found).
Thallus large, in suitable conditions
forming large rosettes of to 10 cm in diam.,
usually appressed to its substrate; lobes
imbricate and divided, 0.5-1.5 cm wide with an
ascending, undulate to crenate and crisped
margin. Upper surface orange to dark brown,
sometimes very dark brown, glabrous, smooth
and shiny, very rarely incrusted in patches,
with transversal, longitudinal or irregular
cracks in mature and old parts of the thallus,
exposing the medulla and with +/- raised edges
and margins occasionally with isidoid granules
in old thalli, but never forming schizidia.
Soralia or isidioid granules or small branched
phyllidia mixed up with thallus fragments
typically formed at the lobe margins, where the
cortex is typically broken and disrupted into
irregular fragments, sometimes present along
laminal cracks. Lower surface typically orange
near the lobes extremites, soon becoming
greenish brown, dark brown, sometimes almost
black, and without veins, interstices white
elliptical, sometimes numerous near lobes
margins but hardly present in old parts.
Rhizines few or abundant, fasciculate to
densely branched and bush-like, black except
at their tips which can be much paler.
Apothecia rare, developed on short lateral lobes,
with a dark reddish brown, circular or ellipsoid
(c. 0.3 mm in diam. or c. 0.3-0.5 × 0.4 mm, in
few cases reaching 0.6 x 0.5 mm), horizontal
disc and a scabrid to verrucose lower side.
Ascospores fusiform, with acute ends, mostly
straight, 3-septate, (34-)35-42(-52) × 6-8 μm.
Pycnidia not seen.
TLC: tenuiorin, methylgyrophorate, gyrophoric acid and zeorin mostly abundant
(other terpenoids present include peltidactylin
and dolichorrhizin but always in traces).
Nomenclature: The type collection of P.
cichoracea could not be located in FI and is
presumably lost; neotypification may be
needed. The epithet is used here following the
interpretation of Swinscow and Krog (1988:
200) for East Africa.
Ecology and distribution: Peltigera
cichoracea is a conspicuous feature of the
epiphytic flora of montane and subalpine
forests between 2300 and 3600 m and artificial
habitats like gardens hedges. It is also common
on trunks of treeferns in a treefern forest in a
197
Fig. 3. A-B. Peltigera cichoracea (PNG, Teptep village, 31 July 1992, E. Sérusiaux s. n., LG). A. General habit. B.
Details of the cripsed margin with granular soredia or isidioid granules. C, E. P. dolichorhiza s. lat. D (PNG, Mt
Gahavisuka, 5 November 1009, E. Sérusiaux s n., LG). C. General habit E. Long simple rhizines. P. dolichorhiza s. lat.
B (PNG, Myola, 16 October 1995, E. Sérusiaux s. n., LG). D. Foveolate upper surface. Scales: A, C = 1 cm, B, D-E = 2
mm.
deep valley in the mountains characterized by
rare but acute frosts and irregular fires. Its
highest locality is at 4100 m, on soil in the
alpine vegetation on Mt. Wilhelm. The species
is rare in the tropical mountains of East Africa
198
in Ethiopia, Kenya and Tanzania (Swinscow
and Krog, 1988: 200) but absent on the
mountains of Rwanda (Sérusiaux et al.,
unpublished results). It has been reported as "a
common species in Taiwan" (Aptroot et al.,
Fungal Diversity
2002: 287) and can be expected elsewhere in
the tropical mountains of S-E Asia. With the
data currently available, P. cichoracea appears
to be the vicariant of P. elisabethae for
montaneous zones in the Paleotropics. Based
on early results of the present study, P. cichoracea has been reported from Papua New
Guinea by Martínez et al. (2003: 307).
Selected specimens examined (out of 56
collections): Papua New Guinea: Madang prov., Huon
Peninsula, Finisterre range, Yupna valley, Teptep village,
trail in NNW and deep valley in N direction, 5°57’S
146°33’E, 2500 m, 30 July 1992, disturbed montane
forest, P. Diederich 10796 (hb Diederich) & E.
Sérusiaux s. n. (LG, 2 collections). Morobe prov.,
Cromwell Mts, 7 km SE of Indagen airstrip, 147°16’E
6°18’S, ca. 2450 m, 19 June 1981, open grassland
(‘kunai’) with scattered treeferns and sinkholes in
limestone, on trunk of treefern, T. Koponen 31066 (H).
Northern prov., Owen Stanley Range, Myola, c. 3 km
NE of guest house, 9°08’S 147°47’E, 2700 m, 16
October 1995, in treefern grassland in deep valley (frost
hollow), E. Sérusiaux 15305 (LG) & H. Sipman 38334
(B). Ibid., English Peaks, 3600 m, August 1988, P.
Lambley 17762 (BM). Simbu prov., Mt. Wilhelm,
Pindaunde valley, near the hut on the S-shore of lake
Piunde, 145°03’E 5° 47’S, 3600 m, 5 August 1992,
subalpine forest remnants on W-slope of valley, E.
Sérusiaux 13983 (LG). Ibid., along track to the summit,
145°03’E 5°47’S, 4100 m, 7 August 1992, on soil in
alpine vegetation, A. Aptroot 31560 (hb Aptroot).
Southern Highlands prov., Mt. Giluwe, 2 km E of the
summit, 143°53’E 6°05’S, 4000 m, 12 September 1982,
subalpine grasslands with scattered shrubs, on the
ground, H. Streimann 24310 (CANB).
Notes: ITS sequences reveal that P.
cichoracea belongs to Sect. Horizontales sensu
Miadlikowska and Lutzoni (2000; Fig. 1). This
paleotropical species is more closely related to
P. elisabethae than to P. horizontalis
(MPIBS=82%). Swinscow and Krog (1988:
200-201) suggested that P. cichoracea may be
considered the sorediate counterpart of P.
polydactyloides Nyl., a species so far known
only from mountains of tropical Africa. The
ITS sequence of a population recently collected
on the volcanoes of Rwanda confirms the
results of Miadlikowska and Lutzoni (2000)
that P. polydactyloides is a member of the P.
neckeri-clade within Sect. Horizontales. Peltigera cichoracea, P. elisabethae and P.
horizontalis differ primarily in their reproducetive mode. All develop apothecia, but P.
elisabethae also produces schizidia and P.
cichoracea is sorediate. Although it is possible
that reproductive strategies are poor species
descriptors as has been shown in other species
aggregates (Buschbom and Barker, 2006 for
Porpidia s. lato), and hence that these three
taxa should be considered conspecific in the
light of weak differentiation in their ITS
sequences, we reject this hypothesis. Indeed,
differences in vegetative morphology, primarily in the veination, seem to correlate with
reproductive characters (Swinscow and Krog,
1988; Vitikainen, 1994; Goward et al., 1995).
The lack of resolution in the ITS may be
indicative of lack of cladogenesis but is not
sufficient to reject our taxonomic hypothesis,
which must be tested further across a broad
geographic range and based on extensive
genetic character sampling.
Peltigera cichoracea is a large, usually
sterile species with a smooth and shiny upper
surface, easily recognized by its very much
disrupted, crisped margin with granular soredia,
isidioid granules or small phyllidia; cracks in
the upper cortex are frequent and can also help
for its identification. The closely related P.
elisabethae Gyeln. is distinguished by its
typically crisped or curled schizidia, formed on
the upper surface, usually near the margins
(schizidia never formed in P. cichoracea
although cracks occur frequently on the upper
surface). Peltigera horizontalis lacks specialized vegetative propagules, and further
differs from the sympatric P. elisabethae by the
network of typically distinct veins.
Peltigera didactyla (With.) J.R. Laundon,
Lichenologist 16: 217, 1984.
Type: Great Britain, "West Yorkshire,
Rombalds Moor" (see Laundon 1984: 217;
original description and material cited from
OXF not seen).
Thallus formed of small orbicular lobes,
0.5-0.8 mm in diam with slightly ascending
margins when young, becoming strongly ascending with almost vertical apothecia and then
forming large rosettes to 2 cm in diam. Upper
surface pale grey to greyish brown, with an
appressed tomentum made of tiny hairs usually
present at least on lobes margins, sometimes
almost absent in mature lobes bearing
apothecia. Soralia typically present on young
orbicular lobes, laminal and rounded, discrete
or confluent, with whitish to bluish soredia,
disapperaing or inconspicuous on mature lobes
199
bearing apothecia. Lower surface with distinct,
raised, pale orange to whitish, anastomosing
veins, leaving distinct interstices. Rhizines
usually few, simple to rarely branched and
bush-like, whitish to pale brown. Apothecia
present on mature lobes, almost vertical,
saddle-shaped, with a red brown disc, and a
denticulate margin. Ascospores fusiform, with
acute ends, almost straight, 3-7 septate, 55-70
(-77) × 4-5 µm. Pycnidia not seen.
TLC: no substances detected.
Nomenclature: The epithet introduced by
W. Withering has been studied in detail by
Laundon (1984), who synonymized the wellknown P. spuria (Ach.) DC. with P. didactyla
(With.) J.R. Laundon.
Ecology and distribution: A rather rare
species, found on humus and gravel, between
2380 and 4200 m elevation. The populations
discovered in Papua New Guinea show almost
the complete variation found in the Northern
hemisphere. Thalli with small, sterile orbicular
lobes with laminal soralia have been found at
high elevation (3600-4200 m), whereas specimens with exuberant, fertile erect lobes with
only scars of eroded soralia are found lower
down (2380-3250 m). The species was reported
from Papua New Guinea by Streimann (1986:
97), under P. spuria, but the corresponding
material has not been examined.
Specimens examined: Papua New Guinea:
Morobe prov., Mt. Sarawaket Southern Range, 4 km SE
of Lake Gwam, headwaters of Busu River, 147°09’E
6°21’E, 3250 m, 6 July 1981, dry stream bed in open
grassland, on humus, T. Koponen 32182 (H, LG). Simbu
prov., Mt. Wilhelm, 145°00’E 5°45’S, 4200 m, July
1967, on soil on summit track, D. McVean 6791 (CANB).
Ibid., Pindaunde valley, near the hut on the shore of lake
Piunde, 145°03’E 5°47’S, 3600 m, 5 August 1992,
subalpine forest remnants, P. Diederich 10198 (hb
Diederich). Southern Highlands prov., Mt. Giluwe,
143°50’E 6°05’S, 4110 m, 1967, wet heath on ridge, D.
McVean 67143 (CANB). Western Highlands prov.,
Minj-Nona Divide, Kubor Range S of Minj, 2380 m, 10
September 1963, advanced regrowth of lower montane
forest, on dead tree stump, R. Pullen 5403 (CANB).
Notes: Peltigera didactyla belongs to a
five-species aggregate studied by Goffinet et al.
(2003). It is characterized by its simple or
rarely branched and usually few rhizines, a
tomentose upper surface (at least at the margins
of young lobes) and the absence of terpenoids
and depsides. The ITS sequence of a
representative collection from Papua New
Guinea confirms its identification (Fig. 2). Two
200
other species of that group occur in Papua New
Guinea: P. extenuata that differs by its
abundant and branched rhizines, and P.
ulcerata that is characterized by its shiny,
etomentose and brown upper surface and its
laminal to marginal soralia.
Peltigera dolichorhiza (Nyl.) Nyl., Flora 57:
71, 1874.
(Figs 3, C-E)
Type: ‘Nova Granata’[Colombia], Bogotá,
2600-2700 m, 1860 (?), Lindig 2519 p. p. (HNYL. 33203—lectotype !; designated by
Galloway 1985: 364 and accepted by
Vitikainen 1988: 136).
Thallus large c. 5-7 cm across (rarely to
10 cm), sometimes rather elongate, with lobes
more or less separated from each other,
extremities usually rounded and 0.8-1.2 cm
wide, with flat or slightly revolute margins;
thalli rather fragile and easily broken apart
when dry. Upper surface glabrous, beige,
orange to dark brown, or bluish grey, smooth
and usually rather shiny, without tomentum or
pruina, usually distinctly undulating with
uneven but distinct depressions. Lower surface
pale orange to brownish near the margins,
becoming dark brown to blackish, with a
network of slightly raised, blackish veins, with
large, numerous, usually elliptical, whitish to
pale orange interstices, forming an attractive
design, with a thin, araneous, orange to
blackish layer of hyphae covering them,
especially near the margins. Rhizines abundant,
usually quite long (to 0.8-1 cm long), simple or
penicillate, exceptionally fasciculate, sometimes quite thick (c. 0.3 mm in diam.). Apothecia absent or rare, on rather narrow, +/- raised,
digitate and strongly revolute lobes, to 0.4-0.7
cm long (incl. the apothecial disc); disc dark
reddish brown, usually saddle-shaped; margins
incised. Ascospores (3-)7-septate, 68-102 ×
4.5-5 µm. Pycnidia not seen.
Notes: P. dolichorhiza is an epithet
usually applied to tropical populations with a
thin and rather fragile thallus, smooth upper
surface usually uneven and with distinct
depressions, raised and digitate apothecia, pale
orange lower surface near the margins, distinct
network of white elliptical interstices over a
dark background towards the center of the
thallus, simple and long rhizines and production of several terpenoids. Our ITS sequences
Fungal Diversity
demonstrate that specimens with such
characters (or slightly differing by their size
and rhizines branching pattern) belong to two
well supported clades forming a paraphyletic
entity (Fig. 1). No morphological or chemical
characters can differentiate the two lineages.
The type material of P. dolichorhiza contains
dolichorrhizin (large spot by TLC), and may be
identical to accessions P. dolichorhiza 13 & 14
in Clade 1, which both produce this terpenoid,
whereas all remaining accessions lack this
compound and are resolved within the wellsupported Clade 2. However several collections
from Rwanda (Africa) produce dolichorrhizin
in significant quantities but unambiguously fall
in Clade 2 (Goffinet & Sérusiaux, unpubl.
results). In the material from Papua New
Guinea, Clade 2 comprizes three distinct
chemotypes and at least one of them is
morphologically distinct (P. dolichorhiza s. lat.
B).
As discussed above (§ Phylogenetic
analysis of ITS sequences), further data must
be obtained before a final decision can be taken
on the taxonomical and nomenclatural status of
material here referred to P. dolichorhiza.
Several collections of P. dolichorhiza
were mentioned from Papua New Guinea by
Streimann (1986: 97) but none have been
examined by us.
Peltigera dolichorhiza s. lat. A
TLC: tenuiorin, methylgyrophorate,
gyrophoric acid and terpenoids present include
peltidactylin (sometimes absent) and zeorin.
Ecology and distribution: Peltigera
dolichorhiza s. lat. A occurs as an epiphyte in
montane forests within a narrow altitudinal
range between 2500-2900 m elev.
Specimens examined (out of 13 collections):
Papua New Guinea, Madang prov., Huon Peninsula,
Finisterre range, Yupna valley, Teptep village, trail in
NNW and deep valley, 5°57’ S 146°33’E, 2500 m, 30
July 1992, disturbed mountain forest, E. Sérusiaux
13622 (LG). Simbu prov., Mt. Wilhelm area, Bundi
Gap, on road Keglsugl-Bundi, 5°48’S 145°09’E, 2800 m,
4 August 1992, subalpine forest remnants, P. Diederich
11052 (hb Diederich).
Peltigera dolichorhiza s. lat. B
TLC: tenuiorin, methylgyrophorate,
gyrophoric acid and terpenoids: zeorin and at
least one unknown (in between dolichorhizin
and zeorin positions in G).
Ecology and distribution: Most of the
collections of P. dolichorhiza s. lat. B have
been found in a single locality, on treeferns
trunks in a treefern forest, in a deep valley in
the mountains characterized by rare but acute
frosts and irregular fires. The second locality
was also made on the trunk of a treefern, in the
Huon Peninsula. The altitudinal range is 27003250 m.
Specimens examined: Papua New Guinea:
Northern Prov., Owen Stanley Range, Myola, c. 3 km
NE of guest-house, 9°08’S 147°47’E, 2700 m, 16
October 1995, in treefern grassland in deep valley (frost
hollow), E. Sérusiaux (LG; 3 specimens collected on 3
different treefern “trunks”) & H. Sipman 38337 (B).
Morobe prov., Mt. Sarawaket Southern Range, 4 km SE
of Lake Gwam, headwaters of Busu R., 6° 21’S 147°09’
E, 3250 m, 6 July 1981, subalpine forest on steep SWfacing slope, on trunk of treefern, T. Koponen 32070 (H).
Notes: Peltigera dolichorhiza s. lat. B
differs from typical P. dolichorhiza by its
larger thalli (up to 15 cm across), with large
lobes, 1-1.8 cm wide, smooth and shiny upper
surface, rhizines few or sometimes abundant,
penicillate to fasciculate, and a unique chemistry (zeorin and at least one unidentified
terpenoid). Amongst other species in Papua
New Guinea, it is similar to P. sumatrana
which has the same morphological features
except for the more common, typically saddleshaped apothecia and a different chemistry
(dolichorrhizin always present and abundantly
produced). P. sumatrana is resolved into Clade
1, whereas three accessions of P. dolichorhiza
s. lat. B (all coming from a single locality)
form a well-supported clade nested within the
poorly resolved Clade 2.
A morphologically similar collection
(Koponen 32070) is tentatively assigned to this
species but it produces peltidactylin and zeorin.
Peltigera dolichorhiza s. lat. C
TLC and HPLC (made by J. A. Elix):
tenuiorin, methyl gyrophorate, methyl lecanorate, methyl evernate, methyl orsellinate, and
four unknown terpenoids (easily recognized on
TLC plates as they include a green and a red
spot).
Ecology and distribution: Peltigera
dolichorhiza s. lat. C exhibits a wide altitudinal
range in Papua New Guinea as it has been
primarily collected between 1850 and 3600 m
with one sample occurring at 4420 m on Mt.
Wilhelm. It mostly grows on mossy trees in
201
montane forests, but can also colonize humus
rich ground, dead stumps and ground mosses.
Selected specimens examined (out of 36
collections): Papua New Guinea, Eastern Higlands
prov., Mt. Gahavisuki Provincial Park, 11 km N of
Goroka, 145°25’E 6°01’S, 2300 m, 17 Mar 1987,
epiphyte in disturbed mossy mountain forest, H. Sipman
22199 (B). Ibid., 2400 m, 3 August 1992, little disturbed
mossy mountain forest, P. Diederich 10582 (hb
Diederich). Ibid., 2300 m, 5 November 1995, little
disturbed mossy mountain forest, dominated by
Castanopsis, E. Sérusiaux s. n. (LG). Madang prov.,
Huon Peninsula, Finisterre Range, Yupna valley, Teptep
village, towards Bambu airfield, 146°33’E 5°57’S, c.
2600 m, 30 July 1992, burnt forest with scattered old
trees and regrowth, E. Sérusiaux s. n. (LG), H. Sipman
35311 (B). Huon Peninsula, 2 km NW of Teptep Airstrip,
146°33’E 5°56.5’S, 2450-2700 m, 28 July 1981, heavily
cut montane rainforest on SE-sloping ridge, on trunk of
Pandanus, T. Koponen 34738 (H). Ibid., 4 km NW of
Teptep Airstrip, 146°32.5’E 5°36’S, 2800-2900 m, 28
July 1981, slightly disturbed montane rainforest on ridge,
on stump, T. Koponen 34691 (H). Morobe prov., Mt.
Kaindi Road, 4 km W of Wau, 146°40’E 7°21’S, 2100 m,
21 June 1979, Nothofagus-Phyllocladus dominated
forest, on rotting trunk, A. Kairo 363 (H). Northern
prov., Owen Stanley Range, Myola, 0-1 km along trail
from guesthouse towards NE, on the right bank of Iora
river, 147°46’E 9°09’S, 2100-2400 m, 15 October 1995,
primary mountain forest on mountain slope, E. Sérusiaux
s. n. (LG, 2 collections) H. Sipman 38261 (B). Simbu
prov., Mt. Wilhelm, 145°00’E 5°45’S, 4420 m, July
1967, on moss near summit, D. McVean 67111 (CANB).
Southern Highlands prov., Onim Forestry Station, 14
km NNW of Ialibu, 143°59’E 6°09’S, 2250 m, 14
September 1982, edge of disturbed montane forest and
grasslands, base of young Nothofagus, H. Streimann
24549 (H). Western Highlands prov., N slopes of
Sugarloaf complex (along Wapu River), 2790 m, 15 July
1960, montane cloud forest, epiphytic on moss, R. D.
Hoogland & R. Schodde 7085 (CANB).
Peltigera dolichorhiza s. lat. D
TLC: tenuiorin, methylgyrophorate,
gyrophoric acid and terpenoids present include
peltidactylin (sometimes absent), dolichorrhizin (always present and produced in large
quantities) and zeorin.
Ecology and distribution: P. dolichorhiza
s. lat. D is an epiphytic species found in
montane forests between 1800-2400 m elev.,
not detected among the plentiful material
collected at higher elevations.
Selected specimens examined (out of 18
collections): Papua New Guinea: Central prov., Owen
Stanley Range, trail from Myola to Naduri, 147°41’E
9°08’S, c. 1800 m, 20 October 1995, montain forest
remnants (with e. g. Lithocarpus), on tree, E. Sérusiaux s.
n. (LG). Eastern Highlands prov., 6 km SW of Lufa,
near Hogabi village, 145°16’E 6°21’E, 1850 m, 14 April
202
1982, montane forest ridge, Nothofagus dominated, on
treelet, H. Streimann 18671 (CANB). Morobe prov.,
Huon Peninsula, Saruwaged Range, Honzukngon village
S of Derim airstrip in Timbe valley, 147°06’ E 6°14’S,
2100 m, 7-8 March 1987, epiphyte in mossy mountain
forest above village, on thin stem, H. Sipman 24434 (B).
Mt. Kaindi, 4 km W of Wau, 146°41’S 7°21’S, 2350 m,
13 March 1982, dense Nothofagus regeneration on
summit, on small tree trunk, H. Streimann 17671 & A.
Bellamy (CANB).
Peltigera erioderma Vain., Philippine Journal
of Science, C. Botany, 8(2): 112, 1913.
(Figs 4 A-C)
Type: Philippines, Luzon, subprov.
Benguet, mons Pulog, ad truncos putridos
arborum et ad terram muscosam, s. d., "For.
Bur. 16362 Curran, Merrit & Zschokke, Bur.
Sci. 8936 McGregor" (TUR—Vainio 9837:
specimen originally numbered 16362 with
annotations by Vainio here selected as the
lectotype !; BM—isolectotype !)
Thalli typically large to 10-15 cm in
diam., with imbricate lobes, rounded at their
extremities and 1-1.5 cm wide, margin revolute.
Upper surface pale orange to brown but almost
white in the best developed specimens, with a
usually well developed and appressed tomentum made of tiny hairs (0,1-0,5 mm high) near
the margins and a +/- appressed, 40-110 μm
thick network of hyphae elsewhere and out of
which hairs 0.1-0.4 mm long, sometimes aggregated in small emerging tufts; in specimens
growing in less suitable conditions, tomentum
appressed (without erecting hairs) irregularly
scattered and mainly present on young lobes
and those carrying apothecia. Lower surface
whitish to pale brown, becoming almost black
towards the center in old specimens, usually
with the medulla hyphae easily seen under the
dissecting microscope, with a distinct and welldeveloped network of large (0.3-0.7 mm),
raised, and soon dark brown to black veins,
which are densely anastomosing; interstices
between the veins usually covered by an
araneous blackish layer near the margins.
Rhizines simple, threadlike to fasciculate and
rarely bushy, pale near the margins but soon
dark brown to black, sometimes very long,
usually not very abundant. Apothecia usually
present, typically horizontal, developed on
swollen lateral lobes; disc dark reddish brown,
remaining concave or becoming flat when
mature, ellipsoid, to 6.5 × 4 mm in diam.,
Fungal Diversity
Fig. 4. A-C. Peltigera erioderma (PNG, Myola, 16 October 1995, E. Sérusiaux s. n., LG). A. General Habit. B. Young
horizontal apothecia. C. Fasciculate rhizines near the edge of the lobes. D-F. P. fimbriata (PNG, T. Koponen 32996,
LG—isotype). D. General habit. E. Hairs over the upper surface. F. Rhizines. Scales: A, D = 1 cm, B-C, E-F = 2 mm.
203
margins slightly incised, rarely crenulate; lobes
carrying the disc distinctly tomentose.
Ascospores fusiform to acicular, straight or
slightly curved, 3(-5)-septate, with rather
rounded ends, (51-)60-75 × 4-5(-6) μm (one
abnormal spore with 11 septa and 90 × 5 μm
seen). Pycnidia absent or very rare, present on
lobes margins, sessile, black, globose, 0.1-0.3
mm in diam., with a central ostiole; conidia not
seen.
TLC: no substances detected.
Nomenclature: Two collections are mentioned in the protologue (n° 16362 and 8936)
and both are in TUR and BM. The specimen
originally numbered 16362 in TUR has hand
written annotations by Vainio and is here
selected a lectotype.
Ecology and distribution: Peltigera erioderma is a rare, epiphytic species growing on
mosses in the montane and subalpine forests
between 2700 and 3400 m; it is known only
from the Mt. Wilhelm area and a treefern forest
in a deep valley submitted to rare but acute
frosts and irregular fires in the Northern prov.
It was reported from New Guinea by Streimann
(1986: 97) and by Martínez et al. (2003: 307).
Selected specimens examined (out of 25
collections): Papua New Guinea: Northern prov.,
Owen Stanley Range, Myola, c. 3 km NE of guesthouse,
147°47’E 9°08’S, 2700 m, 16 October 1995, on treefern
trunk in treefern grassland in deep valley (frost hollow),
E. Sérusiaux s. n. (LG, 2 collections) & H. Sipman
38335 (B). Simbu prov., Mt. Wilhelm area, c. 11 km on
new road under construction from Gembogl to Goroka,
145°09’E 5°55’S, 2800 m, 9 August 1992, mossy
montane forest, A. Aptroot 32830 (hb Aptroot), P.
Diederich 11105 (hb Diederich), E. Sérusiaux 14107
(LG). Southern Highlands prov., Munia logging area,
14 km NW of Ialibu, 143°55’E 6°11’S, 2300 m, 8
September 1982, Nothofagus and Podocarpaceae
dominated forest, on treefern stem, H. Streimann 23224
(CANB, H). Mt. Giluwe, 143°50’E 06005’S, 3350 m, on
trees in subalpine forest, D. McVean 67134 (CANB).
Notes: Peltigera erioderma is readily
identified by its large thalli with a tomentose
upper surface and revolute margin, and its
horizontal apothecia. Peltigera fimbriata also
has a tomentose upper surface but it is
distinguished by its saddle-shaped apothecia
and much longer hairs on its upper surface
[0.1-0.4 mm in P. erioderma vs 1(-1.4) mm in
typical populations of P. fimbriata]. Peltigera
koponenii is another species forming large
thalli with a tomentose upper surface; it differs
204
by its saddle-shaped apothecia and its very
abundant, fibrillose to coralloid rhizines.
Peltigera montis-wilhelmii and P.
papuana are the only other species in the
Papuan flora with horizontal apothecia, the
former being readily identified by the complete
absence of tomentum on its shiny upper surface
and the latter by its much smaller size, smooth
to scabrose upper surface, sometimes with a
whitish pruina at the lobe margin and usually
by the production of marginal phyllidia.
Nevertheless, the morphological boundaries
between P. erioderma, P. montis-wilhelmii and
certains forms of P. papuana remain somewhat
ambiguous (see further comments under P.
papuana).
ITS sequences show that this species is
most closely related to the very similar P.
montis-wilhelmii that is easily distinguished by
its glabrous, shiny upper surface. The shared
ancestry of P. erioderma and P. montiswilhelmii is well-supported but their affinities
within the Sect. Peltigera are uncertain (Fig. 2).
Peltigera extenuata (Vain.) Lojka, Lichenoth.
Univ., fasc. V, n 222, 1886.
Type: Finland, Tavastia australis,
Asikkala, Kaitas 1863, Silén & Norrlin (H—
lectotype !).
Thallus medium-sized, of irregularly
divided lobes, 1-3 cm long and 0.5-1 cm wide,
with typically rounded and concave extremities
(forming attractive rounded ‘cupulae’ with a
slightly revolute margin). Upper surface pale
orange to brownish, in most parts with a dense,
appressed and whitish tomentum. Soralia
abundant, rounded, 0.1-0.2 mm in diam., flat or
slightly concave, but slightly excavate when all
soredia are gone, laminal, mostly present on
young lobes on which they are +/- regularly
scattered, with a distinct rim of cortex when
young; soredia granular, pale orange to dark
brown, usually abundant. Lower surface pale
with a network of whitish, slightly raised but
hardly delimited veins. Rhizines very abundant,
forming a dense cushion, present up to lobe
margins, pale orange or whitish, densely
branched to fibrillose. Apothecia and pycnidia
not seen.
TLC: methylgyrophorate and gyrophoric
acid (traces) in young sorediate lobes, the
soralia typically reacting C+ red.
Fungal Diversity
Nomenclature: The type material has
been studied by Goffinet and Hastings (1995),
and their interpretation is here followed.
Ecology and distribution: Known from
only one locality, on soil in alpine vegetation at
4200 m on Mt. Wilhelm. Peltigera extenuata
has a wide distribution in the Northern hemisphere, being found mainly in arctic tundras,
boreal and montane forests overgrowing
mosses on soil or on rocks, as well as in disued
quarries and other disturbed localities in the
temperate zone. In Asia, it is known from a
single locality in Northern China (for more
details, see Goffinet and Hastings, 1995: 4854). It is reported from New Zealand as a
“chemodeme” of P. didactyla by Galloway
(2000, 2007). It is new for Papua New Guinea.
Specimen examined: Papua New Guinea: Simbu
prov., Mt. Wilhelm, Pinaunde valley, along track to the
summit, 145°03’E 5°47’S, 4200 m, 7 August 1992, on
soil in alpine vegetation, A. Aptroot 33118 (hb Aptroot).
Notes: This taxon was resurrected by
Goffinet and Hastings (1995), and distinguished from the subcosmopolitan P. didactyla by its abundant and densely branched to
fibrillose rhizines, and the occurrence of
methylgyrophorate and gyrophoric acid in
young sorediate lobes and soredia. Recent
studies based on nrDNA sequences clearly
demonstrate its species status (Goffinet et al.,
2003). The single collection from Papua New
Guinea matches that description very well.
Peltigera fimbriata Vitik., Sérus., Goffinet &
Miądl. sp. nov.
(Fig. 4 D-F)
MycoBank: 513027
Etymology: This very spectacular species is
named after its most obvious character, the long hairs
that grow on its upper surface.
Ab aliis speciebus Peltigerae differt thalli superna
facie tomentosa et pilis numerosis et longis instructa.
Type: Papua New Guinea, Morobe prov.,
between Mt. Sarawaket Southern Range and
Iloko village, 2 km SW of Iloko, 147°10’E
6°06’S, 1800 m, 11 July 1981, along stream in
montane rainforest, on boulder, T. Koponen
32996 (H—holotype, LG—isotype).
Species forming large colonies to several
2
m in suitable localites, with large thalli to 2030 cm across, and lobes to 1-1.5 cm wide, with
a revolute margin. Upper surface pale grey or
orange brown to greyish, sometimes almost
white when tomentum and hairs are very abundant, with a thick and dense whitish tomentum
that usually covers it all but sometimes scarce
or even absent, rarely incrusted in patches, with
threadlike, whitish to pale orange, rarely pale
brown, hairs, to 1 (-1.4) mm long, that are
usually abundant and forming tufts or ‘bushes’
mainly near the margins and lobes extremities
(when rare, mostly present in such positions),
rarely absent in specimens with abundant
apothecia. Phyllidia very rare, produced at the
margins of regenerating old lobes. Lower
surface pale orange, with a network of strongly
raised veins which are pale orange near the
margins but soon become dark brown to black,
leaving large ellipsoid interstices. Rhizines
abundant to extremely abundant and then
forming cushions or fluffy masses under the
thallus, simple, penicillate or less frequently
fasciculate, often confluent, at first pale orange
but soon dark brown (at least in parts).
Apothecia absent or abundant, growing almost
vertically on digitated, raised and almost
completely revolute lobes (to 1.5 cm long, incl.
apothecial disc); disc dark reddish brown,
saddle-shaped and elongate, 3-5 mm long with
a slightly raised, usually smooth margin.
Ascospores 5-7-septate, acicular with +/rounded ends, straight, 65-80 × 3.5-4 μm.
Pycnidia almost always present (albeit
sometimes quite few) on lobes margins, sessile,
black, globose, sometimes pyriform, conspicuous because of their size (0.3-0.8 mm in
diam.), with a central, apical ostiole which is
wide open in old and empty ones; no conidia
seen (although c. 15 pycnidia examined).
TLC: no substances detected.
Ecology and distribution: Peltigera
fimbriata forms large and conspicuous thalli
that are primarily terricolous, growing on
gravel by rivers or streams, on bare earth in
recent landslides and on soil in grasslands. It is
locally abundant in the montane zone, but has
also been found in the subalpine and alpine
zones. It also grows on artificial habitats like
shaded road banks. In New Guinea P. fimbriata
exhibits the broadest altitudinal range (from
1300 to 4145 m elev.) among Peltigera species.
Selected specimens examined (out of 55
collections): Papua New Guinea: Enga prov., Mt.
Hagen-Wabag road, 18 km SE of Wapenamanda,
143°58’E 5°47’S, 2700 m, 27 June 1982, advanced
regrowth on slope, on the ground in semi-exposed road
cutting, H. Streimann 21268 (CANB). Madang prov.,
Huon Peninsula, Finisterre range, Yupna valley, Teptep
205
village, trail in NNW and deep valley in N direction,
5°57’S 146°33’E, 2500 m, in disturbed montane forest,
gravel slopes along stream, 30 July 1992, P. Diederich
10984 (hb Diederich). Morobe prov., Aseki-Mdamna
Track, 1 km SW of Aseki, 7°22’S 146°10’E, 1350 m, 22
January 1981, on shaded moist rock, rocky area besides
stream, advanced secondary vegetation besides large
stream in deep gorge, H. Streimann 12520 (B, CANB,
H). Saruwaged Range, Honzeukngon village S of Derim
in Timbe valley, 147°06’E 6°13’S, 1850 m, March 1987,
on soil in gardens, A. Aptroot 17784 & 17788 (hb
Aptroot). Aseki-Menyamya road, 146°06’E 7°16’S,
1910 m, 21 January 1981, cleared area through montane
forest, on moist and shaded road side, H. Streimann & E.
Tamba 12225 (CANB, H). Simbu prov., Mt. Wilhelm,
near the hut on the S-shore of lake Piunde, 145°03’E 5°
47’S, 3500 m, 12 March 1987, subalpine grassland and
shrubs on bottom of valley, H. Sipman 21967 (B).
Southern Highlands prov., Tari Gap, 29 km ESE of
Tari, 143°11’E 5°56’S, 2560 m, 17 December 1982, on
soil bank bordering grassland, J. A. Elix 13338 & H.
Streimann (CANB). Western Highlands prov., Al
River valley, NW of Nondugl, 2000 m, 3 April 1953, on
rock in broad creek, sunny, slightly above water level, R.
D. Hoogland 3216 & 3217 (CANB). Indonesia, Irian
Jaya: Carstensz Mts, 10 December 1971, G. Hope CGE
L6 (COLO). Ibid., Upper Meren Valley, 4145 m, 30
December 1971, terrestrial on fresh moraines, in moist
flat exposed area, G. Hope CGE L38 (CANB).
Notes: Phylogenetic inferences from 5.8S
and ITS2 sequence suggest that P. fimbriata
shares a common ancestor with the allopatric
and morphologically distinct P. ponojensis and
P. monticola (both northern temperate species)
but support for this hypothesis is lacking (Fig.
2).
Peltigera fimbriata is the most readily
recognized species in the Papuan flora as its
tomentose upper surface harbors long whitish
hairs, in particular along the lobes margins, that
furthermore almost invariably carry black
pycnidia. The species is especially common in
the Huon Peninsula at c. 2300-2700 m elev.
where it grows intermingled with P. koponenii.
The latter is easily distinguished by the absence
of whitish hairs and black pycnidia and by its
fibrillose to coralloid rhizines.
The neotropical P. laciniata (G. Merr. ex
Riddle) Gyeln. (Vitikainen, 1998) is
distinguished by its much smaller thallus and
narrower lobes (usually less than 0.5 mm wide
near the extremities), the absence of whitish
hairs, the pale or almost white upper surface
due to a dense tomentum, the upturned margins
of its lobes, and the production of zeorin. Its
rhizines are also very diagnostic as they are
206
quite long and squarrosely branched and
fasciculate.
Several populations are assigned to P.
fimbriata with some hesitation because the
typical whitish hairs on the upper surface tend
to be much shorter, in some cases are rather
sparse and virtually indistinct from the tomentum, or even completely absent. Such populations also have more elongated and narrower
lobes and their rhizines are usually quite
numerous, typically arranged in rows on the
veins (especially on young lobes) and form
compact fluffy masses. They are not clearly
different from the most typical P. fimbriata
populations (for example, the conspicuous,
marginal pycnidia are present) and several
intermediates occur, especially in the Huon
Peninsula. Our ITS sequences (P. fimbriata 2
and 3) do not support segregating these
specimens into another taxon distinct from
typical P. fimbriata (P. fimbriata 1 in Fig. 2),
but more detailed studies are needed to assess
the taxonomic value of the observed
phenotypic variation.
Representative specimens examined: Papua New
Guinea: Morobe prov., Lake Wamba 5 km S of Teptep
airstrip, Teptep-Wantuat trail 10 km S of Teptep,
146°33’E 6°2.5’S, 2550-2700 m, 24 July 1981, open
montane forest, on humus, T. Koponen 33912 (H).
Simbu prov., Mt. Wilhelm, Pindaunde valley, along
track to the summit, 145°03’E 5°47’S, 4000 m, on soil in
alpine vegetation, 7 August 1992, A. Aptroot 33080 (hb
Aptroot). Ibid., Lake Aunde, 6 km SE of Mt Wilhelm,
145°03’E 5°46’S, 3520 m, May 1966, in tussock
grassland, D. McVean 66137 (CANB). Southern
Highlands, Tari Gap, 25 km SE of Tari, 5°56’S
143°06’E, 2740 m, 13 September 1982, on a gentle slope
on semiexposed road cutting in montane forest,
Nothofagus dominated, H. Streimann 24511 (B, CANB,
H). Indonesia, Irian Jaya: Carstensz Mt, 10 December
1971, G. Hope CGE L6 p.p. (CANB).
Peltigera granulosa Sérus., Goffinet, Miądl. &
Vitik. sp. nov.
(Fig. 5 A-C)
MycoBank: 513028
Etymology: The species name refers to the
typically granulose margin.
Ab aliis speciebus Peltigerae differt thallo friabili
et sorediosa, granulosa vel isidiata margine.
Type: Papua New Guinea, Morobe prov.,
Kwama River valley NE of Mt. Sarawaket
Southern Range, 147°12’E 6°04’S, 1700 m, 12
July 1981, along trail in open grassland
between Gumum and Sape villages, on sand, T.
Koponen 33267 (H—holotype; LG—isotype).
Fungal Diversity
Thallus rather fragile, thin and usually
easily broken into pieces, formed of elongated
lobes with rather few divisions and with
rounded and imbricate lobes, usually with an
upturned, crisped margin; when growing on
mossy ground, most lobes erect. Upper surface
bluish grey, or brownish to dark brown, glabrous, smooth and rather shiny, sometimes
with a whitish delicate pruina near the lobes
margin, rarely with irregular cracks in old parts
of the thallus. Margins, especially at the
extremities of the lobes, usually dissolving into
soredioid granules, or isidioid fragments
(rarely with small, ill-looking phyllidia), with
many tending to accumulate on the edge of the
lower surface; tufts of small whitish hairs
sometimes present on the margin. Lower
surface whitish to almost pure white at the
margins, remaining so or becoming blackish
towards the center or in old specimens, with a
network of strongly raised, dark and anastomosing veins which can be almost invisible at
the margins. Rhizines abundant or not, rarely
reaching the margins, simple, very rarely
branched or fasciculate, long, blackish. Apothecia absent or few, developed at the extremities
of erect lobes (ca. 1 cm long) which are +/- flat
or slightly revolute, with a slightly convex dark
reddish brown disc to 4 × 1.5 mm. Ascospores
acicular, with rounded ends, straight or slightly
curved, 5-7 septate, 53-69 × 3-4 μm. Pycnidia
few, marginal, sessile, black, ovoid or slightly
elongate, 0.3-0.5 mm; conidia ovoid, 6-10 × 34 μm.
TLC: no substances detected by TLC.
Ecology and distribution: Peltigera
granulosa seems to be primarily a pioneer
species growing on bare ground, usually sand
or gravel but also on plant debris and peat,
either in natural or artificial habitats (e. g. on
road banks), and mossy soils in the montane
forest zone. It occurs also on mossy trunks
within forests where it usually develops smaller
and sometimes inconspicuous individual lobes.
Its localities are distributed between 1300 and
3660 m elev.
Selected specimens examined (out of 26
collections): Papua New Guinea: Eastern Highlands
prov., Mt. Gahavisuka Provincial Park, 11 km N of
Goroka, 145°25’E 6°01’S, 2300-2450 m, 11 August
1992, on soil in montane forest, A. Aptroot 32396 (hb
Aptroot). Madang prov., S side of Ramu valley, Bundi
village, along road to Bundi Gap, 5°44,9’S 145°14,1’E,
1300 m, 9 November 1995, on roadbank among
secundary vegetation, E. Sérusiaux s. n. (LG) & H.
Sipman 39313 & 39346 (B). Huon Peninsula, Finisterre
range, Yupna valley, Teptep village, trail in NNW and
deep valley in N direction, 5°57’S 146°33’E, 2500 m, 30
July 1992, disturbed montane forest, gravel slopes along
stream, P. Diederich 10875 (hb Diederich). Morobe
prov., Wau area, Edie Creek road, 1850 m, 5 August
1981, bank of road on grassy slope, H. Sipman 15605 &
15606 (U). Northern prov., Owen Stanley Range,
Myola, surroundings of guest-house, 9°09’S 147°46’E,
2100 m, 14-19 October 1995, gravel deposits along Iora
creek, E. Sérusiaux 15150 (LG). Simbu prov., Lake
Aunde, 6 km SE of Mt. Wilhelm, 145°03’E 5°46’S,
3660 m, May 1966, on mossy rock, D. McVean 66180
(CANB, COLO). Western Highlands prov., Kubor
Range, Nona River, 1910 m, epiphytic on tree-base
hanging over river, secondary growth, 4 September 1963,
W. Vink 16486 (hb Aptroot).
Notes: Peltigera granulosa belongs to
Sect. Peltigera, and most likely to the P.
rufescens-group (Fig. 2; Miadlikowska et al.
2003). Recoded INAASE and arc characters
helped to define it as a monophyletic lineage
(MPIBS = 91%), closely related to the sympatric P. papuana, and the allopatric
neotropical P. laciniata (MPIBS=82%). Two
monophyletic groups were detected but these
are indistinguishable morphologically.
Peltigera granulosa is easily recognized
by its rather fragile thallus with a smooth upper
surface, a soredioid, granulose to isidioid
margin, lack of chemistry and strongly raised
veins with long and simple rhizines (when
fully-developed). These features easily distinguish it from its closest relatives.
Peltigera koponenii Sérus., Goffinet, Miądl. &
Vitik. sp. nov.
(Figs 5 D-F)
MycoBank: 513029
Etymology: This new species is dedicated to Dr. T.
Koponen who made large and well processed collections
of Peltigera in Papua New Guinea, mainly from the
Huon Peninsula.
A Peltigera canina differt rhizinis numerosissimis, fibrillosis et coralloidibus.
Type: Papua New Guinea, Morobe prov.,
Lake Wamba 5 km S of Teptep airstrip,
146°34’E 6°0.5’S, 2400-2500 m, 26 July 1981,
open montane Nothofagus-Pandanus forest on
ridge, on trunk, T. Koponen 34327 (H—
holotype, LG—isotype).
Species forming large thalli up to 20 cm
in diam., with elongate lobes (to 6-7 cm long)
sparingly divided, with rounded lobes 0.5-0.8
mm wide at the extremities, and with a revolute
207
Fig. 5. A-C. Peltigera granulosa. A. General habit (PNG, Sarawaket, T. Koponen 33267, LG—isotype). C-B (PNG,
Bundi, H. Sipman 39346, B). B. Marginal soredioid granules (with tufts of hairs in this specimen). C. Lower surface. DF. P. koponenii (PNG, Lake Wamba, T. Koponen 34327, LG —isotype). D. General. E-F. Rhizines and lower surface.
Scales: A, D = 1 cm, B-C, E-F = 2 mm.
208
Fungal Diversity
margin; sometimes with shorter and more
imbricate lobes with their lateral margin
slightly raised to crenulate. Upper surface pale
grey, slightly orange or sometimes bluish-grey,
or with dark and rather large but not delimited
patches, especially towards the thallus center,
with an appressed, whithish to pale brown
tomentum over much of the surface, especially
near lobes margin but sometimes absent; wide
and longitudinal convex folds seen on all wellpreserved thalli. Lower surface whitish to beige
or pale brown, usually hardly darkened towards
the center, whitish interwoven hyphae of the
medulla easily seen under the dissecting
microscope, with a network of pale orange to
brownish, distinctly raised veins. Rhizines
abundant to extremely abundant and then
forming dense (to 5 mm thick) cushions under
the thallus, typically fibrillose and soon becoming coralloid because of further branching
of most lateral fibrils, or fibrillose to +/fasciculate and forming dense ‘bushy’ masses,
at first whitish to pale brown but soon
becoming dark brown to almost black, usually
long (several to 1 cm long). Apothecia absent
or abundant, growing almost vertically on
digitated, raised and revolute lobes (to 1.5 cm
long, incl. apothecial disc); disc dark reddish
brown, saddle-shaped and elongate, 4-5 mm
long, usually with an indistinct margin.
Ascospores 3-septate, acicular, straight, with
+/- rounded ends, 44-53 × 4-4.5 μm. Pycnidia
not seen.
TLC: no substances detected.
Ecology and distribution: Peltigera
koponenii is primarily an epiphytic species
growing on mossy trunks and branches in the
montane (incl. on treefern trunks in deep
valleys characterized by rare but acute frosts
and irregular fires) and subalpine forests. It is
also found overgrowing terrestrial mosses
cushions in the subalpine zone, on gravel by
rivers or streams and on artificial substrats, like
hedges or garden fences in the montane forest
zone. Its altitudinal range is 1850-4270 m.
Selected specimens examined (out of 46
collections): Papua New Guinea: Central prov.,
Kosipe Swamp, Kosipe, 2000 m, November 1992, in
swamp forest, P. Lambley 2028 (BM). Eastern
Higlands prov., Mt. Wilhelm slopes N of Lake Aunde,
11700 feet, June 1966, subalpine forest, terrestrial on
fallen logs and decaying wood, L. K. Wade 8025
(COLO). Madang prov., Huon Peninsula, Finisterre
range, Yupna valley, Teptep village, trail in NNW and
deep valley in N direction, 5°57’S 146°33’E, 2500 m, 30
July 1992, disturbed montane forest, gravel slopes along
stream, P. Diederich 10968 (hb Diederich). Northern
prov., Owen Stanley Range, Myola surroundings of
guest-house, 9°09’S 147°46’E, 2100 m, 14-19 October
1995, small shrubs in grassland, disturbed by clearance,
E. Sérusiaux s. n. (LG). Owen Stanley Range, Mt.
Scratchley, c. 3500 m, 22 December 1985, J. Ismay
16957 (BM). Southern Highlands prov., Mt. Giluwe,
6°05’S 143°50’E, 3200 m, on thickly covered mossy
ground in subalpine grasslands, 19 September 1984, A.
Bellamy 1622 (B, CANB, H). Simbu prov., Mt.
Wilhelm, Pindaunde valley, near lake Piunde, 145°03’E
5°47’S, 3600 m, 5-8 August 1992, on tree in subalpine
forest, A. Aptroot 31346 (hb Aptroot). West Sepik prov.,
S of Oksapmin, 5°14’S 142°12’E, 1850 m, 6 January
1990, epiphyte on a tree in montane forest, L. Hoffman
90-103 (LG, hb Aptroot).
Notes: Peltigera koponenii is nearly
identical with P. canina as it has a large thallus
with tomentose upper surface, saddle-shaped
apothecia growing on raised and rather large
revolute lobes. However, its rhizines are
typically fibrillose-coralloid (a feature sometimes seen in P. canina) and form dense
‘bushy’ masses that are never seen in P. canina.
The lower surface of P. koponenii is thus very
diagnostic. Peltigera koponenii clearly belongs
to the monophyletic P. canina-group together
with P. praetextata, P. evansiana and other
undescribed species (Miadlikowska et al.,
2003); the entire group requires further studies.
Quite interestingly, the ITS sequence of P.
koponenii is virtually identical with that of P.
canina, except for one substitution and one
single nucleotide insertion in the former. A
similar pattern characterizes P. evansiana, a
species that is easily recognized by its laminal
granular isidia, but is phylogenetically indistinguishable from P. canina based on ITS
sequences. A further complementary argument
to support the distinction of P. koponenii is that
the genuine P. canina was not found among the
numerous collections examined from Papua
New Guinea.
Peltigera koponenii is easily distinguished from P. fimbriata by the absence of whitish
hairs on the upper surface (this being a very
typical feature of most populations of P. fimbriata), the absence of conspicuous marginal
pycnidia, the fibrillose to coralloid rhizines and
the septation and size of ascospores (5-7septate and 65-80 × 3.5-4 μm in P. fimbriata vs
3-septate and 44-53 × 4-4.5 μm in P. kopo209
nenii). Peltigera erioderma is another related
species with a usually large thallus and a
tomentose upper surface and lack of secondary
compounds. This species is, however, readily
recognized by its horizontal apothecia, lack of
conspicuous pycnidia and its much shorter
hairs emerging from the tomentum.
Other species with fibrillose-coralloid or
densely squarrose rhizines are:
- P. fibrilloides (Gyeln.) Vitik., known
from the neotropical mountains (Vitikainen,
1998), may be a close relative of P. praetextata
(Flörke ex Sommerf.) Zopf (similar veins
patterns, frequent production of phyllidia, etc.)
from which it differs by its fibrillose rhizines
(rather long and slender, densely covered by
short outspread lateral branches). Although
somewhat reminiscent of those of P. koponenii,
the rhizines remain discrete and form a spongy
cushion.
- P. laciniata (G. Merr. ex Riddle) Gyeln.,
a widespread species in the neotropical
mountains (Vitikainen, 1998), is easily distinguished by its smaller thalli and lobes
(usually not exceeding 0.5 cm wide near the
extremities), the pale or almost white upper
surface due to a dense tomentum, the occurrence of conspicuous, black, marginal pycnidia,
and the production of zeorin. The rhizines are
rather long, squarrosely branched and
fasciculate and the veins tend to be covered by
a thick layer (ca 0.2 mm) of coralloid dark
hyphae; the lower surface of P. laciniata is
thus quite characteristic.
- P. retifoveata Vitik., a disjunct circumpolar species of NW Europe (where it is very
rare), N Asia and WN America (Goffinet, 1992;
Vitikainen, 1994), can be distinguished by its
large thallus, lower surface with a typical
reticulate and foveate veining pattern and the
production of depsides and terpenoids. Its
rhizines are densely squarrose but are rather
scattered and never form dense cushions as in
P. koponenii.
Peltigera montis-wilhelmii Sérus., Goffinet,
Miądl. & Vitik. sp. nov.
(Figs 6 A-C)
MycoBank: 513030
Etymology: This new species is named after the
mountain range where it was collected, the most famous
Mt. Wilhelm range, probably the mountain most
explored by botanists on the whole island.
A Peltigera erioderma differt thalli superna facie
etomentosa.
210
Type: Papua New Guinea: Simbu prov.,
Mt. Wilhelm, Pindaunde valley, near the hut on
the S-shore of lake Piunde, 145°03’E 5°47’S, c.
3600 m, 5 August 1992, subalpine forest
remnants on W-slope of valley, E. Sérusiaux
13984 (LG—holotype).
Thalli +/- circular, 5-7 cm across, with
imbricate rounded lobes to 4 mm wide, with a
revolute or slightly raised margin. Upper
surface smooth or rarely with some whitish
pruina near the lobes margins, or minutely
cracked near the margins, shiny, glabrous,
usually with a nice orange-chamois to brownish colour, or pale greyish brown. Lower
surface whitish to pale orange brown,
darkening to almost black towards the center or
in old parts, usually with the medulla hyphae
easily seen under the dissecting microscope,
with a distinct and well-developed network of
rather large (0.2-0.5 mm), raised and densely
anastomosing veins. Rhizines rather abundant,
typically fasciculate, or sometimes simple to
penicillate, pale when young but soon
becoming black. Apothecia few but present on
almost all thalli examined, typically horizontal,
developed on swollen lateral lobes, disc reddish brown, remaining rather concave, ellipsoid,
to 6.5 × 4 mm, margin typically incised crenate,
and outer exciple rugose to verrucose and
sometimes slightly tomentose or flocculose.
Ascospores fusiform to acicular, with rounded
ends, 3(-5)-septate, 50-61 × 4-5 μm. Pycnidia
not seen.
TLC: no substances detected.
Ecology and distribution: Peltigera
montis-wilhelmii seems to be rare and is
currently known only from the Mt. Wilhelm
area. It grows on branches in the upper
montane forest zone and on the ‘trunk’ of
treeferns in the subalpine forest, between 2800
and 3600 m elev.
Specimens examined: Papua New Guinea:
Simbu prov., same locality as the type, E. Sérusiaux
13987 (LG). Ibid., c. 11 km on new road under
construction from Gembogl to Goroka, 145°09’E 5°55’S,
c. 2800 m, 9 August 1992, mossy montane forest, E.
Sérusiaux s. n. (LG). Ibid., Lake Aunde, 6 km SE of Mt.
Wilhelm, 145°03’E 5°46’S, 3520 m, May 1966, on
treefern stem in tussock grassland near field station, D.
McVean 66236 (CANB, COLO).
Notes: Peltigera montis-wilhelmii is a
very attractive species, characterized by its
smooth, shiny and glabrous upper surface,
large, rounded lobes with usually revolute
Fungal Diversity
Fig. 6. A-C. Peltigera montis-wilhelmii (PNG, Mt Wilhelm, E. Sérusiaux 13984, LG—holotype). A. General habit. B.
Apothecia. C. Lower surface at the edge of the lobes. D-F. P. nana (PNG, Mt Wilhelm, H. Sipman 35729, B). D.
General habit. E. Incrustation on the upper surface. F. Lower surface. Scales: A, D = 1 cm, B-C, E-F = 2 mm.
margins, horizontal apothecia, mainly fasciculate rhizines growing on a distinct network of
dark veins and lack of chemistry. The veins
and interstices pattern of the lower surface and
the rhizines are strikingly similar to those of P.
papuana and populations of the latter with a
211
smooth upper surface without pruina and with
few, if any, marginal phyllidia are difficult to
distinguish from P. montis-wilhelmii. In such
cases the much larger lobes, the usually
revolute margins, and especially the shiny
upper surface are diagnostic. P. montiswilhelmii is yet another glabrous species nested
within the Sect. Peltigera, which is traditionally defined by the presence of laminal
tomentum.
ITS sequences suggest that this species is
most closely related to P. erioderma, a very
similar species that differs, however, by its
completely tomentose upper surface. Both
species form a well-supported clade with
ambiguous affinities within the Sect. Peltigera.
The monophyly of P. montis-wilhelmii (MPIBS
=100%; Fig. 2) is supported also by a unique
pattern of the ITS1-HR shared by all specimens
included in this study.
Peltigera nana Vain., Philippine Journal of
Science, C. Botany, 8(2): 114, 1913
(Figs 6 D-F)
Type: The Philippines, Luzon, subprov.
Lepanto, mons Malaya “F. R. Bona 156”, “ad
terram arenosam” (TUR-V 9850 !; lectotype
here selected).
= Peltigera nana var. philippina Gyeln., Ann.
Mus. Nat. Hungar. 30 (pars Bot.): 133, 1936. Nomen
superfl. Syn. nov.
Type: same as the lectotype.
= Peltigera tereziana var. philippinensis Gyeln.,
Ann. Mus. Nat. Hungar. 30 (pars Bot.): 133, 1936. Syn.
nov.
Type: The Philippines, Luzon, subprov.
Benguet, “Bur. Sci. 5878 Ramos”, “ad terram
arenosam” (TUR-V 9855 — holotype !).
= Peltigera macra Vain., Philippine Journal of
Science, C. Botany, 8(2): 114, 1913. Syn. nov.
Type: The Philippines, Luzon, prov.
Pangasinan, “Bur. Sci. 8298, Ramos”, “ad
terram calcaream et argillaceam et humosam”
(TUR-V 9854 !; lectotype here selected).
Thalli not forming large rosettes but
nevertheless quite robust, composed of
imbricate lobes, not exceeding 0.5 cm in width,
with a slightly upturned margin. Upper surface
orange brown to dark brown, or greenish
brown, almost blackish in old thalli, glabrous,
smooth and shiny, usually with a distinct
whitish pruina near the lobes margins and with
large, incrusted patches towards the centre of
the thalli, these patches appearing irregular and
212
slightly swollen when well developed. Lower
surface pale orange to brown at the margins,
soon becoming black (especially in old
specimens), without veins but with elliptical
and whitish to white interstices (in old specimens the white interstices are larger and hence
the veins form a conspicuous network), surface
hyphae of veins and especially interstices
typically very distinct. Rhizines sparse,
blackish, fasciculate or more rarely simple.
Apothecia present, rather abundant, developed
on digitated and raised lobes with revolute
margins, to 1.2-1.5 mm long (incl. the
apothecial disc); disc dark reddish brown,
elliptical, 0.6-0.7 × c. 0.4 cm, saddle-shaped
with a slightly incised or indistinct margin.
Ascospores fusiform to acicular, straight or
slightly curved 3(-5) septate, 62-82 × 5-6 μm.
Pycnidia not seen.
TLC: tenuiorin, methylgyrophorate,
gyrophoric acid, peltidactylin and dolichorrhizin (abundant), sometimes zeorin and other
unidentified terpenoids.
Ecology and distribution: Peltigera nana
has a wide ecological amplitude as it has been
found on roadbanks and rocky outcrops, in
rather open and secondary vegetation, at 13001800 m (lower montane forest zone), as well as
on Mt. Wilhelm and in the mountains of the
Huon Peninsula, on mossy and gravel soil, in
the subalpine zone at 3500-3700 m elev. We
have seen material of Peltigera nana from
Luzon Island in The Philippines and Papua
New Guinea; the species is mentioned from
India (Himalaya) by Awasthi and Joshi (1982:
55) and is liklely to be widespread in S-E Asia.
Selected specimens examined (out of 21
collections): Papua New Guinea: Madang prov., S side
of Ramu valley, Bundi village, along road to Bundi Gap,
5°44,9’S 145°14,1’E, 1300 m, 9 November 1995, on
roadbank among secundary vegetation, E. Sérusiaux s. n.
(LG). Morobe prov., Mt. Sarawaket Southern Range,
2.5 km S of Lake Gwam and E of Mt. Enggum,
147°07’E 6°21’S, 3500-3570 m, 9 July 1981, deep Ssloping valley with scattered scrub and spring, on basic
cliff, T. Koponen 32766 (H). Aseki-Menyamya Rd,
Spreader Divide, 12 km NW of Aseki, 146°06’ E 7°16’S,
1980 m, exposed, cleared area through montane forest,
21 January 1981, on shaded road cutting, H. Streimann
12057, 12098 & E. Tamba (CANB). Ogeranang, 1800 m,
28 June 1986, on roadside bank (partly sheltered), P. W.
Lambley 17685 (BM). Simbu prov., Mt. Wilhelm,
Pindaunde valley, near the hut on the S-shore of lake
Piunde, 145°03’E 5°47’S, c. 3600 m, subalpine forest
remnants on W-slope of valley, 6 August 1992, E.
Sérusiaux 13981 (LG). Western Highlands prov.,
Fungal Diversity
Baiyer River, upper Trauna valley, 1700 m, 18 April
1986, 1700 m, on rocky outcrops of ridgetop, P. W.
Lambley 16955 (BM).
Examined specimens of Peltigera melanocoma
(all in L): Indonesia, Java: n° 62 coll. Junegh., in m.
Pangerango 3-5000’ (lectotype — here designated); n°
60 coll. Jungh [two collections in different envelopes];
n° 61 coll. Jungh ; n° 63 coll. Jungh [two collections in
different envelopes]; sine numero coll. Jungh, in
cacumine m Sëndoro [+ illisible notes]; sine numero coll.
Jungh, April. Silva montis Pangerango 3-5000’. Prov.
Preanger, in decliv. austral. montis Panperango, prope
Tjibodas, in horto montano, regio nubium, alt. +/- 1420
m, 28 April 1894, V. Schiffner Iter Indicum 1893/94, n°
3290.
Notes: Peltigera nana is distinguished by
its usually robust thalli, albeit not forming large
colonies or rosettes, growing on the ground, its
upper surface usually with pruina at the lobes
margins and sometimes large and conspicuous
incrusted patches, its dark lower surface with
an attractive network of distinct elliptical,
whitish interstices, and surface hyphae usually
typically very distinct. It does produce terpenoids, with dolichorhizin in large quantities.
In our phylogenetic tree, P. nana falls
within the Sect. Polydactylon, forming a wellsupported clade with P. sumatrana, P. oceanica, and P. weberi. Peltigera nana is closely
related to P. oceanica (easily distinguished by
the absence of tenuiorin and methylgyrophorate) and P. weberi (easily distinguished by
the presence of marginal soredia) but its exact
position remains ambiguous. However, the two
specimens for which ITS sequences were
produced do not form a monophyletic group,
and more data are thus needed to assess the
variation of that species.
Besides its original description and
further comments by Gyelnik (1936), P. nana
has only been reported from New Zealand
[Murray, 1960, as P. dolichorhiza var. nana
(Vain.) Js. Murray, and Galloway, 2000]. The
morphological data provided for those populations (« broadly rounded, thin, papery lobes » ;
Galloway, 2000: 25, 2007, 2008) clearly point
to a different species.
Typification with further notes: Three
collections are mentioned in the protologue for
P. nana and all have been examined (TUR-V !);
as quoted by Gyelnik (1936: 132-133), they are
small and poorly developed. Their chemistry is
identical: tenuiorin, methylgyrophorate, gyrophoric acid, peltidactylin, dolichorrhizin (abun-
dant) and zeorin. The specimen TUR-V 9850
(«F. R. Bona 156») is the best developed and is
thus here selected as the lectotype for the
epithet nana. Gyelnik (1936: 133) considered
this collection to represent a new variety, e. g.
P. nana var. philippina Gyeln.; following our
lectotypification, this epithet becomes superfluous. The collection TUR-V 9855 (subprov.
Benguet, Bur. Sci. 5878 Ramos) was considered as a new variety of P. tereziana (var.
philippinensis) by Gyelnik (1936: 133).
Although the material is scanty, it matches very
well P. nana and we can thus reduce this
variety into synonymy with it.
As for Peltigera macra, we examined the
three collections that are mentioned in the
protologue (TUR-V!). Vainio claims that it
differs from P. nana (and from P. didactyla; as
P. spuria in the protologue) by its rhizines:
«nervis e rhizinis breviter crebreque tomentosis
dignota». We see no differentiation in rhizines
morphology. The chemistry of the syntypes of
P. macra is similar to that of P. nana but
differs in the presence of several further
tripernoids ; these chemical differences are not
sufficient to recognize a different taxon. The
specimen «Ramos 8298» bears annotations by
Vainio and is therefore here selected as the
lectotype.
The identity of P. melanocoma Mont. &
Bosch (Lichenes Javanici: 6, 1857; also in
Miquelon, Pl. Jungh. 4: 432, 1857) has also
been examined. Nine collections were received
from L. The original publication mentions
“Hab. ad terram et truncos ins. Javae (m.
Pangerango 3-5000’), Jungh.”; the specimen
labelled “n 62. Java coll. Jungh. in m.
Pangerango 3-5000’” is designated as the
lectotype. As all other specimens, it is terricolous, forms well-developed thalli with
subparallel lobes, crenate-crispy margins, scattered incrustations on the upper surface, dense
and rather long rhizines, reticulate dark veins
separated by ellipsoid, whitish and thus
forming a strong contrast interstices, digitate
and rather small apothecia, and it produces
tenuiorin, methylgyrophorate, gyrophoric acid,
peltidactylin and especially abundant dolichorrhizin. Peltigera melanocoma may be closely
related to P. nana and even be conspecific.
However, the thalli of P. melanocoma are
much larger and with subparallel lobes with a
213
crispy margin, and thereby reminiscent also of
P. oceanica. Pending further research on
material from Java, we decided to retain both
taxa as distinct.
Peltigera oceanica Gyeln., Fedde Repertor. 29:
9, 1931.
(Figs 7 A-C)
Type: Philippines, Luzon, Benguet prov.,
Baguio, March 1907, A. D. E. Elmer 8456 p.p.
(W—holotype !)
= Peltigera oceanica f. dealbata Gyeln., Fedde
Repertor. 29: 9, 1931. Syn. nov.
Type: Philippines, Luzon, Benguet prov.,
Baguio, March 1907, A. D. E. Elmer 8456 p.p.
(W—holotype !)
= Peltigera oceanica f. luzoni Gyeln., Fedde
Repertor. 29: 9, 1931. Syn. nov.
Type: Philippines, Luzon, Benguet prov.,
Baguio, March 1907, A. D. E. Elmer 8456 p.p.
(W—holotype !)
Thallus forming nice rosettes, to 10 cm
across, made of rather narrow lobes (2-3 x 0.40.6 cm) with a typically raised and crisped
lateral margin, usually showing the border of
the lower surface; margins sometimes +/regularly incised. Upper surface glabrous,
smooth and shiny in most parts but sometimes
slightly pruinose at lobes margins, pale
greenish brown to dark brown, rarely incrusted
in old parts, or distinctly foveolate (seen in
only one collection). Phyllidia or lobules
usually absent, rarely developed on lateral
margins (perhaps damaged and regenerating
parts). Lower surface white to pale orange with
numerous, raised, pale brown (near the margins)
to almost black (towards the centre) veins,
leaving a nice network of numerous, elliptical
and pale to whitish interstices of variable size.
Rhizines usually abundant, simple to fasciculate, rather long, dark brown to blackish,
usually with paler to almost whitish tips.
Apothecia present or absent, sometimes quite
abundant, typically saddle-shaped, developed
on strongly revolute lobes at the extremities
and usually raised, 0.4-0.7 cm long (incl.
apothecial disc); disc dark reddish brown with
a slightly raised and incised margin. Ascospores 5-septate, acicular, straight, with
rounded ends, 48-55 × 4-5 µm. Pycnidia not
seen.
TLC: only terpenoids (dolichorhizin
abundant, peltidactylin and zeorin usually
present in lower amounts, other unidentified
214
terpenoids sometimes present); tenuiorin and
methylgyrophorate always lacking.
Nomenclature: The type collection is
representative of the populations studied and,
although inappropriate (the distribution pattern
of this species has nothing to do with oceans),
the epithet can be used without hesitation.
Ecology and distribution: Peltigera
oceanica typically grows on gravel or sandy
soil, usually over rock boulders near streams; it
has never been found on humus, peat, plants
debris, or on trees. Its altitude range is between
1250 and 3300 m. Beside the type collections
(incl. forma’s) on Luzon Island in the
Philippines archipelago, it has been mentioned
from the “Papuan Provinces” by Martínez et al.
(2003: 307) on early results of the present
study. It is likely to be found elsewhere in S-E
Asia.
Selected specimens examined (out of 21 species):
Papua New Guinea: Morobe prov., Mt. Sarawaket
Southern Range, 4 km SE of Lake Gwam, 147°09’E
6°21’S, 3300 m, 5 July 1981, open grassland (‘kunai’)
with scattered treeferns, on basic cliff, T. Koponen
31965 (H, LG) & 31966 (H). Northern prov., Owen
Stanley Range, Myola, surroundings of guest-house,
9°09’S 147°46’E, 2100 m, 14-19 October 1995, on
boulders along Iora creek in open-field, E. Sérusiaux s. n.
(LG). Southern Highlands prov., Onim Forestry
Station, 14 km NNW of Ialibu, 143°59’E 6°09’S, 2280
m, 18 September 1983, montane forest beside river, on
rock near river bank, A. Bellamy 1295 (B, CANB).
Western Highlands prov., Al River Valley, NW of
Nondugl, 2000 m, 3 April 1953, on rocks slightly above
water level, R. D. Hoogland 3209 (CANB, H).
Notes: Only three species produce
terpenoids in the absence of tenuiorin and
methylgyrophorate (Vitikianen, 1986): P. dolichospora (Lu) Vitik., a member of Sect.
Polydactylon known from Nepal and China/
Sichuan, P. laciniata (G. Merr. ex Riddle)
Gyeln., a common neotropical tomentose species, and P. oceanica Gyeln., described from
The Philippines. The collections from Papua
New Guinea have been carefully compared
with the type material of the latter and they
appear conspecific. Besides the absence of
tenuiorin and methylgyrophorate, P. oceanica
is distinguished by its glabrous upper surface,
lobes with a raised and crisped (mainly
laterally) margins that are sometimes slightly
pruinose or incrusted, and a nice network of
conspicuous elliptical interstices on the lower
surface. Peltigera nana is quite similar, except
for the lateral margin that is very rarely raised
Fungal Diversity
Fig. 7. A-C. Peltigera oceanica (PNG, Onim, A. Bellamy 1295, B). A. General habit. B. Crisped margins. C. Lower
surface. D-F. P. papuana (PNG, Teptep, E. Sérusiaux 13656, LG—holotype). A. General habit. B. Apothecia and
marginal phyllidia. C. Lower surface. Scales: A, D = 1 cm, B-C, E-F = 2 mm.
215
and crisped, and the production of tenuiorin
and methylgyrophorate.
Peltigera oceanica falls within Sect.
Polydactylon, forming a well-supported clade
with P. nana and P. weberi (Fig. 1). Three
representative samples from Papua New
Guinea have identical ITS sequences that differ
clearly in their hypervariable region from P.
nana (MPIBS=91%). The relationships of these
species remain ambiguous.
Peltigera papuana Sérus., Goffinet, Miądl. &
Vitik. sp. nov.
(Figs 7 D-F)
MycoBank: 513031
Etymology: Together with P. fimbriata, this
species is the most distinctive taxon of the genus in
Papua New Guinea, and is thus named after this country.
Ab aliis speciebus Peltigerae differt thalli superna
facie laevigata vel scabrosa, marginibus phyllidiosis,
elevatis et crenatis, et apotheciis horizontalibus.
Type: Papua New Guinea: Madang prov.,
Huon Peninsula, Finisterre range, Yupna valley,
Teptep village, deep valley in N direction,
146°33’E 5°57’S, 2300-2750 m, 31 July 1992,
mossy montane forest, E. Sérusiaux 13656
(LG—holotype).
Thallus forming attractive rounded
rosettes to 10-12 cm in diam. in suitable
conditions, lobes imbricate, not exceeding 0.50.7 cm wide, with raised and crenate
(sometimes +/- crisped) margins (margins are
distinctly revolute and swollen when an
apothecium starts its development in them);
lobes extremities sometimes with tufts of tiny
whitish hairs. Upper surface orange brown,
beige brown to grey brown, rarely bluish grey,
smooth or faintly to distinctly scabrose with
tiny pellucid hairs developing on the tiny
‘verrucae’, especially near the lobes margins,
sometimes whitish-pruinose near young lobes
extremities which therefore have a frosted
appearence; some specimens with longitudinal
cracks. Lateral margins frequently becoming
incised-lacerate, with flattened, branched
lobules that are usually fragile and easily
removed; in well-developed specimens, lobules
developing into typical, rather brittle digitate
phyllidia, to 0.2 cm long, with their extremities
sometimes tomentose or pruinose and distinctly
enlarged. Lower surface whitish with the
interwoven hyphae of the medulla and the
bluish tinge of the photobiont easily seen under
the dissecting microscope, with an attractive
216
network of 0.2-0.3 mm large and at least
slightly raised veins, pale orange to brown near
margin, otherwise dark brown to blackish, and
separated by whitish elliptical interstitices,
especially near the margins. Rhizines abundant,
rather long, dark brown to black, thread-like to
penicillate, sometimes branched and rarely
confluent. Apothecia usually present, numerous,
typically horizontal, even at early stages,
developed on swollen lateral, and occasionally
tomentose lobes; disc dark reddish brown,
remaining concave and partly covered by
teared up and wooly remnants of vegetative
tissues for quite a long time (thus giving them a
crenate appearance), later becoming flat or
irregular, rounded when mature to ellipsoid,
reaching 6 × 4 mm but usually smaller;
margins typically incised-crenulate and lobes
carrying the disc usually distinctly tomentose.
Ascopores fusiform and rather narrow, with +/acute ends, 3-5-septate, (36-)38-51 × 4-5 μm.
Pycnidia very rare, as small swollen brownish
dots on the margins of lateral lobes; conidia not
seen.
TLC: no substances detected.
Ecology and distribution: Peltigera
papuana is a widespread species at mid
elevations, found mainly on trees in primary to
heavily disturbed montane forests; it is rather
ubiquitous as it also grows on mossy soil and
road banks. It has been found between 1700
and c. 2800 m, well below the subalpine zone.
This is true only if we do not include two
specimens of the “small form”, collected at
3400 and 3600 m.
Selected specimens examined (out of 40
collections): Papua New Guinea: Eastern Highlands
prov., along jeep road from junction of GorokaKundaiawa main road toward summit of ridge on way to
Marafunga, c. 5500 ft, 22 June 1968, forming dense pads
up to one foot diam. over fern debris on trunks of
Pandanus, W.A. Weber & D. McVean, Lichenes
exsiccati distributed by the University of Colorado
Museum, Boulder n° 319 (B, CANB, H, LG). Mt.
Gahavisuka Provincial Park, 145°25’ E 6°01’S, little
disturbed mossy mountain forest, 2400 m, 3 August
1992, P. Diederich 10581 (hb Diederich). Madang
prov., Huon Peninsula, Finisterre range, Yupna valley,
Teptep village, trail in NNW direction, towards Bambu
Airfield, 146°33’E 5°57’S, c. 2500 m, 30 July 1992, on
garden fence, H. Sipman 35270 (B). Ibid., trail in N
direction, 31 July 1992, on tree, E. Sérusiaux 13655
(LG). Morobe prov., Mt. Sarawaket Southern Range, 4
km NNE of Lake Gwam, 147°9.5’E 6°19’S, 2850 m, 4
July 1981, open grassland with scattered treeferns, T.
Koponen 31592 (H). Simbu prov., Mt. Wilhelm area, c.
Fungal Diversity
11 km on new road under construction from Gembogl to
Goroka, 145°09’E 5°55’S, c. 2800 m, 9 August 1992,
road bank in mossy montane forest, E. Sérusiaux 14106
(LG). Kombugomambuno, 8 km SE of Mt. Wilhelm,
145°03’E 5°47’S, 3320 m, May 1967, epiphytic in
subalpine tussock grassland with tree ferns, L. K.. Wade
(CANB; 2 collections). Southern Highlands prov.,
Kengaput, Mendi-Kauga Road, 6 km SSE of Mendi,
6°12’S 143°4’E, 1700 m, 10 September 1982, on
Pandanus stem in Dacrydium dominated swamp, H.
Streimann 23694 (B). Western Highlands prov.,
Nebilyer River, 28 km WNW of Mt. Hagen, 5°48’S
143°49’E, 2760 m, 23 June 1982, on upper branches of a
small Elaeocarpus in disturbed montane forest,
Nothofagus, Podocarpaceae and Cunoniaceae dominated,
H. Streimann 20585 (B, CANB, H).
Examined specimens for the “small” form akin P.
erioderma: Papua New Guinea: Northern Prov.,
Owen Stanley Range, Myola, c. 3 km NE of guest-house,
9°08’S 147°47’E, 2700 m, 16 October 1995, in treefern
grassland in deep valley (frost hollow), H. Sipman 38336
(B). Simbu prov., Mt. Wilhelm, Piunde valley, 3400 m,
13 April 1987, on treefern, P. W. Lambley 16965 (BM).
Western Highlands prov., N of summit of Mt. Giluwe,
c. 3600 m, 31 August 1987, P. W. Lambley 16963 (BM).
Selected specimens of P. tereziana examined (out
of 10 collections): Australia, New South Wales (NSW):
Southern Tablelands. Bendoura Range: the hearts of
‘Marble Arch’, past ‘The Big Hole’, 33 km (direct) SSW
of Braidwood. 35˚43’30S 149˚41’20E, 620m, limestone.
Very moist/wet area with mosses, fersn (Dicksonia sp.)
and elderberry buch (Sambucus sp.), foliose lichen on a
large rock that was exposed from within the creek –
downstream from the caves, January 1999, T.K.
Overeem 11 (CANB). NSW: Southern Tablelands,
Moodong Creek, Marble Arch, Deau, 34 km SW of
Braidwood. 35˚43’30”S 149˚41’24”E 580m, dry
sclerophyll on moderate slope to creek, on semi-shaded
boulder in seasonal stream. 11 January 1999, H.
Streimann 63484 (CANB). NSW: Poblue Swamp,
Barrington Topds State Forest, 50 km WNW of
Gloucester, 31˚58’S 151˚26’E 1450m, eucalyptus
pauciflora, Leptospermum and epicridaceae fringed
swampy area. On Eucalyptus, 25 April 1990, J. A. Elix
24690 (CANB). Victoria, Moranding, Kilmore,
144°57’E 37°18’S, April 1890, on rock by creek, F. R.
M. Wilson 1188 (VIC-isotype). New Zealand, North
Island, Hawke’s Bay Land District, Puketitiri, 176°31’E
39°17’S, on soil, coll. between 1975 and 1985, J. K.
Bartlett s. n. (CHR 449001). South Island, Otago Land
District, Moa Creek, on soil, 17 February 1979, I. Brown
s. n. (CHR 490548). Ibid., Lammermoor Range, on soil,
21 March 2007, D. J. Galloway s. n. (LG).
Notes: Peltigera papuana is easily
recognized by its smooth to scabrose upper
surface, usually with pruina near lobes margins,
small lobes with raised and crenate margins,
presence of marginal phyllidia, horizontal
apothecia developed on small lobes and
absence of chemical compounds. Some
specimens, though well-developed, almost lack
marginal phyllidia, and are morphologically
similar to P. montis-wilhelmii, which differs by
the larger and glabrous lobes with slightly
raised to revolute margins and a shiny upper
surface (see further comments under this
species).
ITS sequences of representative collections show that P. papuana likely belongs to
the P. rufescens-group (sensu Miadlikowska et
al., 2003) with close affinities to the newly
described P. granulosa, and the South American P. laciniata (Fig. 2). Support for the
monophyly of this taxon is lacking.
Whether these populations should be
assigned to P. tereziana Gyeln. (Öst. Bot. Z. 77:
220, 1928; = P. subhorizontalis Gyeln., Ann.
Cryptog. Exot. 5: 39, 1932; Vitikainen, 2004)
has been carefully assessed. Indeed, the type
material of the latter is described by Galloway
(1985: 365-366; 2000: 40-41) as forming ‘neat
rosettes’, with lobes ‘glossy in parts, rarely
delicately white-pruinose in parts imparting a
slightly frosted appearance to lobes, margins
incised-lacerate or minutely crenate-lobulate’,
with horizontal apothecia and lacking terpenoids or depsides. Examination of type
material and other specimens from Australia
and New Zealand (see list below) shows it is a
different species. The surface is never scabrose
and is usually smooth and glossy, with
distinctly pruinose and frosted patches, usually
at the margins but sometimes covering large
parts of the surface. The lobes are sometimes
incised and lobulate but never form the typical
brittle digitate phyllidia typical of the Papuan
specimens. Furthermore P. tereziana is primarily terricolous rather than epiphytic. A further,
more subtle difference pertains to the horizontal position of apothecia: in P. papuana, they
immediately develop in a horizontal position
with a slightly concave and crenate disc,
whereas in P. tereziana, they develop in an
erect position with a slightly saddle-shaped
disc, and they eventually get laterally larger
and adopt a horizontal position. Furthermore,
inferences from ITS sequences obtained from
typical collections of P. tereziana reveal that
this species shares a common ancestry with P.
membranacea and P. degenii (Fig. 2) and not
with members of the P. rufescens-group, which
217
includes P. papuana. Peltigera tereziana is
thus yet another species of Sect. Peltigera
lacking tomentum.
In Papua New Guinea, three specimens
were first assigned to a small form of P. erioderma as they share its diagnostic characters:
tomentose upper surface, horizontal apothecia
and lack of chemical compounds. However,
they form much smaller rosettes (to 3-6 cm in
diam.) with imbricate lobes c. 0.5-0.7 cm wide,
and are akin to P. papuana in general habit.
Indeed, ITS sequences from one of these
collections (H. Sipman 38336, P. papuana 7 in
Fig. 2) reveal affinities to specimens typical of
P. papuana even though they lack some
diagnostic characters of this species, especially
the smooth to scabrose upper surface and the
production of marginal phyllidia. However,
none of the characters of these specimens are a
priori incongruent with P. papuana. No further
sequences could be obtained and we thus
refrain from distinguishing another taxon on
that basis.
Peltigera sumatrana Gyeln., Rev. Bryol.
Lichénol. 5: 72, 1932.
(Figs 8 A-C)
Type: Indonesia, Sumatra, Korinchi Peak,
7300’, 28 April 1914, H. C. Robinson & C. B.
Kloss n° 139 (W—holotype !)
Species forming large (to 10 cm across)
and rather robust thalli, with large, rounded
lobes to 1 cm wide, or smaller ones with more
narrow or elongate lobes not exceeding 0.5-0.7
cm wide; margin revolute. Upper surface
glabrous, smooth and shiny in most parts but
sometimes distinctly pruinose at lobes margins
or on lateral phyllidia or lobules (when
developed), pale beige brown, bluish grey or
dark brown. Phyllidia or lobules usually absent,
sometimes developed on lateral lobes, margins
or rarely the surface of damaged and
regenerating parts of thalli, to 2-3 mm in length,
some typically pruinose. Lower surface pale
brown or orange with a reddish-brown to black
araneous cover near the margins, but soon
becoming jet black, usually without any veins
but with elliptical and whitish interstices
towards the center. Rhizines usually abundant,
typically fasciculate and densely branched,
sometimes forming a dense cushion on the
lower surface or remaining well separated from
each other and arranged in concentrical rows.
218
Apothecia present or absent, sometimes quite
abundant, developed on strongly revolute lobes
at the extremities which are not necessarily
raised, 0.5-0.8 cm long (incl. apothecial disc),
typically saddle-shaped; disc dark-reddish
brown with a slightly raised and incised margin.
Ascospores acicular to fusiform, straight or
slightly curved, 3-5-septate, with rounded ends,
58-78 x 4-5 μm.
TLC: tenuiorin, methylgyrophorate and
terpenoids; the full spectrum includes 9
terpenoids, but dolichorhizin always dominant
and peltidactylin and zeorin always present but
in smaller quantites.
Nomenclature: The holotype has been
examined, including by TLC: although quite
small and badly preserved, the specimen is
clearly conspecific with the numerous
populations from Papua New Guinea, that are
thus treated as P. sumatrana.
Ecology and distribution: Peltigera
sumatrana is typically a species of the montane
forest (1850-2900 m), where it can be quite
common on trees, over terricolous mosses and
even on peat (e. g. Gahavisuka Provincial Park,
Eastern Highlands prov.). A slightly different
population has been sampled at lower elevation
(1300-1600 m, see below). This species
reaches the alpine zone around 3600-4000 m
where it grows in rather protected niches, and
never on bare soil and gravel. It has also been
found on ‘artificial’ substrates in the montane
forest zone, like rocky road banks, soil in
gardens, or even on the roof of huts. Beside the
type collection from Sumatra, it has never been
mentioned in the literature. It is most likely
widespread in the mountains of S-E Asia, and
is new for New Guinea.
Selected specimens examined (out of 42
collections): Papua New Guinea: Eastern Hihglands
prov., Mt. Gahavisuka Provincial Park, 11 km N of
Goroka, along trail to lookout, 6°1’S 145°25’E, 2400 m,
3 August 1992, little disturbed mossy montane forest, P.
Diederich 10585 (hb Diederich). Ibid., 5 November 1995,
little disturbed mossy mountain forest, dominated by
Castanopsis, E. Sérusiaux s. n. (LG). Madang prov.,
Huon Peninsula, Finisterre range, Yupna valley, Teptep
village, trail in NNW direction towards Bambu Airfield,
146°33’E 5°57’S, c. 2300-2650 m, 30 July 1992,
disturbed montane forest, with scattered old trees and
regenerating shrub layer after fire, E. Sérusiaux 13609
(LG). Morobe prov., Mt. Kaindi, 5 km W of Wau,
7°19’S 146°44’E, 2300 m, 26 July 1982, on base of
small Evodiella in disturbed Nothofagus forest near
summit, H. Streimann 22425 (B, CANB). Saruwaged
Fungal Diversity
Fig. 8. A-C. Peltigera sumatrana (PNG, Gahavisuka, 5 November 1995, E. Sérusiaux s. n., LG). A. General habit. B.
Apothecia and margin of the lower surface. C. Lower surface. D-F. P. weberi. D, F (PNG, Goroka, W. A. Weber and D.
McVean L-50140, COLO—holotype). E. (PNG, Mt Kaindi, H. Streimann 33372, CANB). D. General habit. E.
Sorediate margins. F. Incrustations on the upper surface. Scales: A, D = 1 cm, B-C, E-F = 2 mm.
219
Range, Honzeukngon village S of Derim in Timbe valley,
147°06'E 6°13'S, 1850 m, 7-8 March 1987, on branches
and small trunks in clearing, H. Sipman 24513 (B).
Simbu prov., Mt. Wilhelm, Pindaunde valley, near the
hut on the S-shore of lake Piunde, 145°03’E 5°47’S,
3600 m, 5 August 1992, subalpine forest remnants on
W-slope of valley, E. Sérusiaux 13985 (LG). Southern
Highlands prov., Onim Forestry Station, 14 km NNW
of Ialibu, 6°09’S 143°59’E, 2250 m, 15 September 1982,
cleaned areas with grasses between road and the river, H.
Streimann 24783 (B, H, LG). Western Highlands prov.,
Jimi-Waghi Divide, 9 km N of Banz, on Tabibuga road,
5°43’S 144°38’E, 2400 m, 7 July 1982, on treelet stem,
H. Streimann 22295 (H). Indonesia, Irian Jaya:
Carstensz Mts, Carstensz meadow, 3540 m, 10
December 1971, open tussock grassland with Astelia
hummocks, on peat, G. Hope CGE L3 & L6 p.p.
(CANB).
Specimens examined: Papua New Guinea:
Madang prov., S side of Ramu valley, Bundi village, on
slope towards Mt. Pizetara, 5°44.9’ S 145° 14.1’E, 13001600 m, 8 November 1995, disturbed montane forest, E.
Sérusiaux 16401 & 16402 (LG), H. Sipman 39212 &
39213 (B).
Notes: Inferences from substitutions in
the ITS sequences reveal that this species is
related to the P. oceanica-complex and thus
falls into Clade 1 within the Sect. Polydactylon.
The accession P. sumatrana 5, altough
morphologically similar to the remaining specimens of P. sumatrana has a different ITS1-HR
pattern; although it is placed outside of the P.
sumatrana-clade (MPIBS=96%; Fig. 1), we
retain it within that species pending further
studies.
Peltigera sumatrana is easily distinguished by its usually large thalli with a smooth
upper surface, lower surface without veins,
most usually jet black but with elliptical and
whitish interstices, fasciculate and densely
branched rhizines, saddle-shaped apothecia and
production of terpenoids dominated by dolichorhizin. Populations growing on bare ground
by roads or in gardens can be atypical as they
are composed of smaller individuals with
smaller lobes; they can be easily identified by
their chemistry and by the typical lower surface,
especially the fasciculate and densely branched
rhizines.
Populations here referred to the same
species have been extensively sampled in
Bundi, in a lower montane forest (c. 1300-1600
m) on the southern side of the Ramu river.
They match P. sumatrana in all morphological
details (including the occurrence of pruina on
young lobes, and sometimes phyllidia on
lateral lobes, lower surface and rhizines, and
size and septation of ascospores) except for
two features: they form less robust and even
fragile, sometimes very thin thalli with a +/foveate surface — thus reminiscent of P.
Peltigera ulcerata Müll. Arg., Flora 63: 261,
1880.
Type: "Prope Apiahy Brasiliae merid.
crescit: Puiggari n. 1023 p. p." (G—holotype !;
W—isotype !).
Thallus usually quite small and inconspicuous, especially when growing amongst
healthy pleurocarpous mosses on trees, but
sometimes reaching 5-6 cm in diam. when
being a pioneer species on its substrate, formed
of adjacent or imbricate lobes, mostly
suborbicular but sometimes elongate, 0.5-1.0
cm wide, typically concave (especially when
young), with a +/- revolute margin. Upper
surface glabrous, smooth, rarely +/- scabrose,
dull or slightly shiny. Soralia always present,
orbicular to ellipsoid, c. 1-2 × 1 mm, laminal
but most usually near the lobes margins and
becoming +/- marginal in old specimens,
sometimes confluent; when young with a
distinct rim of remnants of cortex. Soredia
usually bluish-grey, farinose to granular,
abundant or almost absent. Lower surface pale
orange to greyish, with a network of slightly
raised, greyish dark veins that are inconspicuous near the margins. Rhizines simple to
fasciculate, usually abundant, dark brown
except near the margins where they are much
paler (especially at their base). Apothecia and
pycnidia not seen.
TLC: methylgyrophorate in soralia
(which thus react C+ red); gyrophoric acid
detected in some but not all collections.
Nomenclature: The use of this epithet has
been soundly established for a long period and
no problem was detected.
220
dolichorhiza — and most examined thalli (4
out of 5) fail to produce zeorin. Compared to
the usually quite robust specimens of typical P.
sumatrana that were all collected at higher
elevations, they look quite different and may
represent distinct populations evolving towards
such a status. The ITS sequence of a representative specimen (P. sumatrana 4, Fig. 1) is
identical to that of some more robust and hence
typical specimens P. sumatrana.
Fungal Diversity
Ecology and distribution: Peltigera
ulcerata is mainly a muscicolous species on
trees in montane forests, including in disturbed
localites, but it can also grow on dead logs on
the ground, on mosses over rocks and amongst
terricolous mosses in alpine vegetation. It has
not been collected on roadbanks or on ground
near streams or rivers. Its altitudinal range
extends from 2300 to 4200 m. This species is
widespread in the tropical mountains of the
three continents and in temperate areas of the
southern hemisphere (Swinscow and Krog,
1988: 203; Galloway, 2000: 41-42; Goffinet et
al., 2003), and occurs also in the Western
Himalayan Province (Martínez et al. 2003:
307). Aptroot and Sipman (1991: 232) reported
the first collections from Papua New Guinea.
Selected specimens examined (out of 13
collections): Papua New Guinea: Eastern Highlands
prov., Mt. Gahavisuka Provincial Park, 11 km N of
Goroka, 6°01’S 145°25’E, c. 2300 m, 5 November 1995,
little disturbed mossy montane forest dominated by
Castanopsis, 5 November 1995, E. Sérusiaux 16200
(LG). Madang prov., Huon Peninsula, Finisterre range,
Yupna valley, Teptep village, 146°33’E 5°57’S, 2300 m,
30-31 July 1992, on hedges of Cordyline between
gardens, A. Aptroot 32278 (hb Aptroot). Northern prov.,
Owen Stanley Range, Myola, surroundings of guest
house, 9°09’S 147°46’E, 2100 m, 14-19 October 1995,
in primary forest in valley, E. Sérusiaux s. n. (LG).
Southern Highlands prov., 30 km after Kaupena,
6°12’S 143°55’E, 2350 m, 11 October 1989, rainforest
clearfelling area, on dead wood, K. Van der Gucht 89695 (hb Aptroot). Simbu prov., Mt. Wilhelm, Pindaunde
valley, along track to the summit, 145°03’E 5°47’S,
4200 m, 7 August 1992, on soil in alpine vegetation, A.
Aptroot 31572 (hb Aptroot).
Notes: Peltigera ulcerata belongs to the
P. didactyla-complex (Goffinet et al., 2003). It
is diagnosed by its small, orbicular to elongate
lobes with a smooth, mostly shiny brown upper
surface and elliptical, laminal to submarginal
or marginal soralia producing rather farinose
and usually bluish soredia. Some small and
sterile specimens of P. didactyla may be
difficult to distinguish when their upper surface
has almost no tomentum; they can be
recognized by their mostly laminal soralia and
coarser, partly corticate soredia.
Based on its ITS sequence a representative collection from Papua New Guinea is
resolved as sister to a specimen of P. ulcerata
from Rwanda (Fig. 2), corroborating its identification, and hence the presence of this taxon in
Papua New Guinea.
Peltigera weberi Sérus., Goffinet, Miądl. &
Vitik. sp. nov.
(Figs 8 D-F)
MycoBank: 513032
Etymology: This new species is named after Prof.
W.A. Weber (Univertsity of Colorado) who was the first
lichenologist to collect lichens extensively in Papua New
Guinea; he brought splendid specimens from the upper
montane zone of Mt. Wilhelm and several became types
of new species, such as Calathaspis devexa I.M. Lamb &
W.A. Weber, Dimerella weberi Vězda and Pertusaria
gyalectoides Vězda.
Ab aliis speciebus Peltigerae differt thalli superna
facie etomentosa, marginis farinoso-sorediosis et
terpenoideas continente.
Type: Papua New Guinea: Eastern
Highlands prov., road just above Goroka on
way to power plant, 4000 feet, 22 June 1968,
on clay banks beside the road, W. A. Weber &
D. McVean L-50410 (COLO—holotype; LG—
isotype).
Thallus small and inconspicuous, made
of rounded, isolated or imbricated lobes,
mostly c. 0.5 cm large near the extremities,
pale olive brown to dark bluish green, flat or
slightly concave, with the margins typically
upturned when sorediate. Upper surface glabrous, smooth, rarely somewhat scabrose, and
rather dull. Soralia always present but not
developed on all lobes, typically marginal and
hardly spreading on the lower surface; soredia
usually bluish-grey, farinose to granular. Lower
surface orange to greyish, with a poorly
developed network of slightly raised, greyish
veins, which can be hardly visible at the
margins. Rhizines simple to fasciculate, not
abundant, pale brown. Apothecia and pycnidia
not seen.
TLC: tenuiorin, methylgyrophorate,
dolichorhizin and zeorin.
Ecology and distribution: Peltigera
weberi is a rare species and grows on earth and
rock debris, and on road banks, between 1200
and 1450 m.
Specimens examined: Papua New Guinea:
Madang prov., S side of Ramu valley, Bundi village,
along road to Bundi Gap, 5°44,9’S 145°14,1’E, 1300 m,
9 November 1995, on roadbank among secundary
vegetation, H. Sipman 39314 (B). Morobe prov., Mt.
Kaindi road, 5 km WNW of Wau, 146°41’E, 1450 m, 9
January 1983, montane forest on moderate slope, on the
ground and on rock, H. Streimann 33372 (CANB).
Notes: Peltigera weberi is easily
distinguished by the following combination of
characters: inconspicuous glabrous and smooth
thallus, soralia present and strictly marginal
221
and production of terpenoids. Peltigera cichoracea (Sect. Horizontales) produces soredia
and terpenoids but typically forms large thalli
(10 cm in diameter) on trees. All other
sorediose species found in Papua New Guinea
differ by their lack of terpenoids; moreover P.
didactyla, P. extenuata and P. ulcerata (Sect.
Peltigera) differ by their predominantly
laminal soralia, whereas P. granulosa (also
Sect. Peltigera) has a granulose margin that is
never genuinely sorediose. Peltigera didactyla
and P. extenuata differ further by their
tomentose upper surface. The allotatric P.
collina (Sect. Horizontales) has glabrous lobes
with marginal soralia, too, but these are
coarsely granular, partly corticate and fingerlike and thus easily distinguish this species
from P. weberi.
The ITS sequence of a single representative collection demonstrates that P. weberi
belongs to Sect. Polydactylon and is closely
related to P. oceanica and P. nana (Fig. 1).
Currently this is the only sorediate species in
the Sect. Polydactylon.
Lichenicolous fungi
Although lichenicolous fungi are not the
main purpose of this study, we report here
species growing on Peltigera thalli from Papua
New Guinea. We confirm the existing records
of the following peltigericolous taxa from
Papua New Guinea (Aptroot et al., 1997: 81,
97, 112 & 208): Leptosphaerulina peltigerae
(Fuckel) Riedl, Lichenopeltella peltigericola
(D. Hawksw.) R. Sant., Nectriopsis lecanodes
(Ces.) Diederich & Schroers and Vezdaea
dawsoniae Döbbeler. During this study, three
additional species were found (identifications
by Dr. P. Diederich): Corticifraga fuckelii
(Rehm) D. Hawksw. & R. Sant., on
unidentifiable necrotic material (Northern prov.,
Owen Stanley Range, Myola, 9°08’S 147°47’E,
2700 m, 16 October 1995, on tree fern, E.
Sérusiaux s.n., LG); Corticifraga peltigerae
(Fuckel) D. Hawksw. & R. Sant., on P.
koponenii (Simbu prov., Mt. Wilhelm, 4270 m,
26 August 1970, L. Stapf s. n., CANB); Scutula
epiblastematica (Wallr.) Rehm, on P.
koponenii (Central prov., Kosipe Swamp, 2000
m, November 1992, P. W. Lambley 2028, BM).
222
Acknowledgments
The authors are very grateful to the curators of
the following herbaria who placed interesting specimens
at their disposal: B, BM, CANB, CHR, COLO, L, U, and
to their colleagues and friends who also made large
collections available, especially Dr. André Aptroot, Paul
Diederich, Peter Lambley and Harrie Sipman. A set of
specimens from East Africa (Kenya and Rwanda) was
kindly made available to us by Prof. Dr. Eberhard
Fischer and Dr. Dorothee Killmann. A preliminary work
by Florin Crisan (Univ.’Babes-Bolyai’, Cluj-Napoca,
Roumania) on the material collected by E. Sérusiaux has
been helpful for the comprehension of the variation of
several widespread species. They also want to thank very
warmly Mr. I. Cremasco (University of Liège) for his
precious help in performing TLC analyses, Dr. P.
Diederich (National Museum of Natural History in
Luxemburg) for his help in the identification of
lichenicolous fungi, Prof. Jack A. Elix (Australian
National University in Canberra) for TLC and HPLC
examination of Peltigera dolichorhiza s. lat. C, Dr.
David J. Galloway and Dr. Simone Louwhoff for kindly
providing specimens of P. tereziana from New Zealand
and Australia, and Dr. Robynn Shannon for leading the
initial sequencing effort. The work by Emmanuël
Sérusiaux was made under several awards from the
Belgian "Fonds de la Recherche Fondamentale
Collective" that funded his expeditions to Papua New
Guinea and his visit to the University of Connecticut.
His stay in Helsinki was supported by the EU (European
Union) Large Scale Facility Program. Bernard Goffinet
acknowledges support from the University of
Connecticut.
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