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INTRODUCTION
The systematic placement of lichenized fungi in
which no sexual stage is known has traditionally been
based on thallus structure and form, supplemented by
secondary chemistry. Numerous sterile and often wide-
spread lichens have been placed in genera characterized
by a particular sexual stage using these types of charac-
ters (e.g., Coppins & James, 1979). In some cases, sub-
sequently discovered ascomata have led to placements
being confirmed, or found to be erroneous. As lichenized
fungi are wisely exempted from Art. 59, the option for a
dual nomenclature is closed. Molecular methods, howev-
er, mean that as in other fungi for which no sexual stage
is known, totally sterile lichenized species can be
unequivocally placed in families or genera whose mem-
bers have sexual stages (e.g., Arup & Grube, 1999;
Ekman & Tønsberg, 2002).
This paper addresses the nomenclature and place-
ment of a sterile crustose-placodioid lichen with a
whitish, pruinose and sorediate thallus. The lichen is
locally frequent on somewhat soft calcareous rocks in the
western Mediterranean, and is currently generally
referred to as Lecanora lisbonensis G. Samp. (Nimis,
1993; Llimona & Hladun, 2001; Fig. 1). The species was
placed in Lecanora Ach. (Lecanoraceae) because of
reported apothecia with simple colourless ascospores
described by Sampaio (1921), but has also been referred
to Placodium auct. non (Ach.) DC. (Lecanoraceae) as the
thallus was placodioid by Klement (1965), and to Buellia
De Not. (Physciaceae) by Werner (1976) who considered
he had fertile material with brown muriform ascospores.
In addition, the monotypic genus Coscinocladium occi-
dentale Kunze (Kunze, 1846a) and the species name
Variolaria gaditana Clemente (Clemente, 1807; Fig. 2)
have been suggested to belong to the same species,
notably by Tavares (1956, 1958). De Notaris was evi-
dently shown material by Kunze, and concluded that this
lichen belonged to an unpublished genus; Kunze accept-
ed De Notaris’ view and coined the generic name
Coscinocladium. De Notaris added: “Botanicis occiden-
Crespo & al. • Coscinocladium, an overlooked lichen genus53 (2) • May 2004: 405–41453 (2) • May 2004: 405–414
Coscinocladium, an overlooked endemic and monotypic Mediterranean
lichen genus of Physciaceae, reinstated by molecular phylogenetic analysis
Ana Crespo
1
, Oscar Blanco
1
, Xavier Llimona
2
, Zuzana Ferencová
3
& David L. Hawksworth
1*
1
Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, Plaza de Ramón y
Cajal, Ciudad Universitaria, Madrid 28040, Spain. acrespo@farm.ucm.es (author for correspondence),
oba@farm.ucm.es; davidh@terra.es
2
Departament de Biologia Vegetal (Botànica), Universitat de Barcelona, Diagonal 645, Barcelona 08028,
Spain. llimona@pothos.bio.ub.es
3
Institute of Biology and Ecology, University of Pavol Jozef Šafárik, Mánesová 23, Košice 04001, Slovakia.
zuzferenc@yahoo.com
The position of the sterile western Mediterranean crustose-placodioid lichen generally known as Lecanora lis-
bonenesis has been investigated using mitochondrial SSU rDNA sequence data. It proves to belong to an inde-
pendent genus of Physciaceae, for which the generic name Coscinocladium is available, and the earliest species
name is Variolaria gaditana. Previous reports of apothecia in the species prove to be a result of mixtures with
other lichen species. A lectotype is selected for L. lisbonensis, a neotype designated for V. gaditana, and the
new combination Coscinocladium gaditanum made. The species is described, illustrated, its ecology discussed,
and a distribution map provided.
KEYWORDS:
Ascomycota, biogeography, Buellia, Europe, Lecanora, Lecanorales, Mobergia.
Fig. 1. Coscinocladium gaditanum (MAF 9855 - neotype).
405
talis hauc plantam lichenosam, accuratus observandam
commendo”
1
(in Kunze, 1846a, b). However, these early
species names were not taken up by later workers
because of either uncertainties over their typification, or
because a combination of Kunze’s epithet into Lecanora
was precluded by the existence of the name L. occiden-
talis (Lynge) Lynge (Lynge, 1940).
We endeavoured to follow De Notaris’ admonish-
ment 157 years later through collecting and studying
fresh material of this taxon, examining its phylogenetic
position by molecular methods, and also investigating
and typifying the pertinent names suggested to belong to
the species. As a result, we show that the species repre-
sents a hitherto unrecognised genus of Physciaceae, we
fix the application of relevant species names by neo- and
lectotypification, and we determine that the binomial that
should be applied under the Code is Coscinocladium
gaditanum.
MATERIALS AND METHODS
Taxa sampled. —
For this study we used material
or sequences from collections named as Lecanora lisbo-
nensis (i.e., Coscinocladium gaditanum) and several
other samples from different taxonomic groups. 19 spec-
imens were collected specifically for DNA isolation,
amplification and sequencing for this study, while
sequences from 24 other taxa were downloaded from
GenBank. Full details of the specimens and GenBank
accession numbers are included in Appendix 1 and 2
(see online version of Taxon). Sequences from the
mtSSU were used for estimating the taxonomic position
of the target samples. Then, in order to identify the clos-
est relationships further, nuclear ITS rDNA was also
used to analyse relationships with the most similar
sequences resulting from a BLAST search.
DNA extraction and PCR amplification. —
Total DNA was extracted using the DNeasy Plant Mini
Kit (Qiagen), with minor modifications as described by
Crespo & al. (2001). Mitochondrial DNA amplification
was undertaken with the primers NMS1 and NMS2 (Li
& al., 1994) and SSU1 and SSU3R (Zoller & al., 1999),
and nuclear ITS rDNA with ITS1F (Gardes & Bruns,
1993) and ITS4 (White & al., 1990).
Each PCR reaction contained the following ingredi-
ents: 29.75 µl dH
2
O, 5 µl 10X PCR buffer where the
MgCl
2
2 mM was already included (Tris-HCl 75mM pH
9.0; KCl 50 mM; (NH
4
)
2
SO
4
20mM), 1 µl dNTP 10 mM,
2.5 µl of a 10 µM dilution of each of the primers and 1.25
µl DNA Polymerase (1 unit/µl, Biotools). This cocktail
was mixed with 8 µl of the DNA template. The PCR
amplification for the mitochondrial gene ran 35 cycles:
denaturation at 94ºC for 60 s, annealing at 57–58ºC for
60 s, and extension at 72ºC for 90 s. The reaction was
carried out in an automatic thermocycler (Hybaid
OmniGene). The PCR amplification of the ITS region
ran 30 cycles: denaturation at 94ºC for 60 s, annealing at
54ºC for 60 s, and extension at 72ºC for 90 s. This reac-
tion was carried out in an automatic thermocycler
(Techne Progene).
The PCR products were purified through a Biotools
Bioclean DNA Purification column kit, according to the
manufacturer’s specifications. Sequencing was per-
formed on both strands using the ABI PRISM
TM
Dye ter-
minator cycle Sequencing Ready Reaction kit (Applied
Biosystems), with the PCR primers. The following
cycling profile was used: denaturation for 3 min at 94ºC,
25 cycles of 10 s at 96ºC, 5 s at 50ºC and 4 min at 60ºC.
The products for sequencing were electrophoresed on an
ABI PRISM 377 DNA sequencer (Applied Biosystems).
Sequence analysis. —
Sequences were compared
with the assistance of Windows SeqMan (DNAStar) to
check for reading errors, and, when possible, to resolve
ambiguities. Sequences of SSU and ITS contain seg-
ments that are very variable. Since standard multiple
alignment programs, such as Clustal (Thompson & al.,
1994), become less reliable when sequences are highly
divergent, we used an alignment procedure employing a
linear Hidden Markov Model (HMM) as implemented in
the software SAM (Hughey & Krogh, 1996;
http://www.cse.ucsc.edu/research/compbio/sam. html).
Regions that were not aligned with statistical confidence
were excluded from the subsequent phylogenetic analy-
sis.
The data were analysed using a Bayesian approach
(Larget & Simon, 1999; Huelsenbeck & al., 2000).
Posterior probabilities were approximated by sampling
trees using a Markov Chain Monte Carlo (MCMC)
method. The posterior probabilities of each branch were
calculated by counting the frequency of trees that were
visited during the course of the MCMC analysis. The
Crespo & al. • Coscinocladium, an overlooked lichen genus 53 (2) • May 2004: 405–414
406
Fig. 2. Original description of Variolaria gaditana
(Clemente, 1807: 295).
1
Transl.: Western [European] botanists are commended to study in more detail this lichen plant.
program MRBAYES 2.01 (Huelsenbeck & Ronquist,
2001) was employed to sample the trees, and the analy-
sis was performed assuming the general time reversible
model (Rodriguez & al., 1990), including estimations of
invariant sites and assuming a discrete gamma distribu-
tion with six rate categories (GTR+I+G) for the com-
bined analyses. A run with one million generations start-
ing with a random tree and employing 12 simultaneous
chains for mitochondrial and eight for ITS sequences
was executed; every 100
th
tree was saved into a file. We
used the default settings for the priors on the rate matrix,
branch lengths, gamma shape parameter, and the propor-
tion of invariable sites. A Dirichlet distribution was
assumed for the base frequency parameters, and an unin-
formative prior was used for the topology (default set-
tings).
We plotted the log-likelihood scores of sample
points against generation time, and considered that sta-
tionarity was achieved when the log-likelihood values of
the sample points reached a stable equilibrium value
(Huelsenbeck & Ronquist, 2001). The initial 400 trees
were discarded as burn-in before stationarity was
reached. Using PAUP 4.0b10 (Swofford, 2002), majori-
ty-rule consensus trees were calculated from 9600 trees
sampled after reaching likelihood convergence to calcu-
late the posterior probabilities of the tree nodes. Unlike
nonparametric bootstrap values (Felsenstein, 1985),
these are estimated probabilities of the clades under the
assumed model (Rannala & Yang, 1996) and hence pos-
terior probabilities equal to and above 95 are considered
significant support. Phylogenetic trees were drawn using
TREEVIEW (Page, 1996).
The polarity of the characters was assessed by out-
group comparisons using Protoparmelia badia and
Rhizoplaca bullata in the mitochondrial tree. Due to its
position as a member of the well-supported Buellia
group in the mitochondrial tree, Dirinaria applanata was
chosen as outgroup for the ITS rDNA region analysis.
RESULTS
Molecular studies. — MtSSU rDNA analysis.
Insertions of varying lengths were present in Dirinaria
applanata and Heterodermia leucomela. In addition, a
small insertion of 14 bp was detected in Lecanora
muralis, Protoparmelia badia, and Rhizoplaca bullata. A
matrix of 704 aligned nucleotide position characters was
used for the analysis, after 133 positions ambiguously
aligned and all insertions were excluded.
Two major monophyletic groups were discovered
(Fig. 3). The first comprises the three Coscinocladium
gaditanum collections, and the species of Anaptychia,
Heterodermia, Phaeophyscia, Phaeorrhiza, Physcia,
Physconia, and Rinodina studied, together with Buellia
lindingeri (99 posterior probability). And the second,
groups Amandinea, Calicium, Cyphellium, Diplotomma
(incl. Diploicia), Dirinaria, Pyxine, Tholurna, Texo-
sporium, and all other Buellia species studied (100 pp).
Within the Physcia group, a well-supported mono-
phyletic group includes the two species of Anaptychia as
the sister group of a branch where Phaeorrhiza sarep-
tana is basal in relationship with the three samples of
Physconia.
Within the Buellia group, a well-supported clade
joins Diplotomma canescens as the basal sister group of
Dirinaria applanata, which is also the basal sister group
of the three species of Pyxine.
Nuclear ITS rDNA sequences analysis.
Sixteen
samples, including Mobergia calculiformis and Physcia
tenella var. maritima, the most similar sequences in the
BLAST search, were included in the analysis; also other
sequences belonging to other taxa in the Physcia group
clade were included (see Fig. 3). A matrix of 526
nucleotides was obtained, of which 72 ambiguous posi-
tions were excluded in the analysis.
Three main groups were found (Fig. 4). Although
without significant support (85 pp), Coscinocladium
gaditanum, Mobergia calculiformis and Rinodina
sophodes grouped in the same clade. C. gaditanum and
M. calculiformis samples formed two independent
monophyletic clades, in both cases with 100 pp.
Morphological studies. —
Of the 11 collections
under the name Lecanora lisbonensis in PO, the thalli of
all conformed to that of Coscinocladium gaditanum as
typified here. Three were found to have Lecanora-type
apothecia intimately mixed with the sterile lichen, push-
ing between cracks in the placodioid lobes (PO 991L,
1563L and 2317L); all three have detailed annotations
and sketches in Sampaio’s hand, with dimensions of
apothecia and ascospores conforming to those in the pub-
lished account (Sampaio, 1921). The apothecia did not
arise from a well-defined thallus and none were found to
originate in the placodioid lobes. Squash mounts were
made in water, and all three have ascospores 10.5–12 ×
4.5–5(–6) µm and have thalline and apothecial characters
which conform to L. flotowiana Spreng., as understood
by Fröberg (1997). However, we recognize that the spore
dimensions are in the upper end of the range given by
Foucard (1990; 7–14 × 3–7 µm) and Purvis & al. (1992;
(7–)8.5–14 × (3–)4–7 µm) as regards length in the L. dis-
persa group, to which L. flotowiana belongs, and that the
complex is yet to be studied critically in the
Mediterranean region.
The rare apothecia reported by Werner (1976) were
quite different from those of L. flotowiana. According to
Werner’s original notes and drawings, which are now
held by X. L., the collection he thought had apothecia
Crespo & al. • Coscinocladium, an overlooked lichen genus53 (2) • May 2004: 405–414
407
was from Rabat, and we located and examined this mate-
rial (Morocco, Rabat, en contre-bas de l’hopitâl Marie-
Feuillet sur sables agglomérées, avec Caloplaca auran-
tia, 8 April 1933, leg. R. G. Werner, BC-Werner); the
notes, measurements and drawings Werner made corre-
spond so closely to those he published that there is no
doubt this is the collection on which he based his con-
clusion that the species belonged in Buellia. The speci-
men consists of a whitish thallus partially overgrown by
C. gaditanum. We were unable to find any apothecium,
but Werner’s description of the apothecia, paraphyses
and ascospores leads us to consider this may well be the
species generally known in the Iberian peninsula as
Diplotomma ambiguum (Ach.) Flagey or a closely relat-
ed species, and not a lichenicolous fungus. Werner’s
transfer of Sampaio’s epithet was therefore evidently
based on this mixed collection. We note that the name D.
ambiguum was placed as a synonym of D. alboatrum
(Hoffm.) Th. Fr. by Nordin (2000), but we feel that his
treatment may have been too broad.
Werner (1976) also mentioned pycnidia in a different
collection of C. gaditanum (as “L. lisbonensis”) which
we also located and examined (Morocco, “Sidi Moussa,
rochers maritimes calcaires, à 500 m de l’océan”, Jan.
1932, leg. J. Gattefossé, BC-Werner). These seem to be
genuine pycnidia of the species; according to Werner’s
drawings they are immersed, globose, with pleurogenous
conidiogenous cells (Vobis, 1980: type VI) and cylindri-
Crespo & al. • Coscinocladium, an overlooked lichen genus 53 (2) • May 2004: 405–414
408
Fig. 3. Mitochondrial SSU rDNA majority-rule consensus tree based on 9600 trees from a B/MCMC tree sampling pro-
cedure. Posterior probabilities equal to or above 95 are given above the branches.
Buellia group
Physica group
cal conidia he measured as 3.8–5 × 1.3 µm. These struc-
tures are very similar to those known in Physcia and
Physconia (Vobis, 1980), and consistent with the rela-
tionships that emerged from our molecular studies.
TAXONOMY
Coscinocladium Kunze, Flora 29: 768. 1846.
Type species: C. occidentale Kunze [= C. gaditanum
(Clemente) A. Crespo, Llimona & D. Hawksw.].
Coscinocladium gaditanum (Clemente) A. Crespo,
Llimona & D. Hawksw., comb. nov.
≡ Variolaria gaditana Clemente, Ens. Veg. Andalucia:
295. 1807 [basionym]. – Neotypus (hic designatus):
Spain, Andalucía, Cádiz, Calle Honduras, on walls
of the old city, 15 Feb 2003, leg. A. Crespo, C.
Bencomo & J. F. de Bobadilla (MAF 9855).
≡ Pertusaria communis var. gaditana (Clemente)
Colm., Enum. Rev. Pl. Hisp.-lusit. 5: 833. 1889.
= Coscinocladium occidentale Kunze, Flora 29: 768.
1846. – Lectotypus (hic designatus): [Spain,
Andalucía, Cádiz, 1844/45], “pl. Willkomm 987”
(UPS L-82639).
= Ricasolia cesatii [var.] γ plumbea Bagl., Nuovo
Giorn. Bot. Ital. 11: 70. 1879. – Holotypus: Italy,
Sardinia, leg. Moris (destroyed fide Tavares, 1958).
≡ Solenopsora cesatii var. plumbea (Bagl.) Zahlbr.,
Cat. Lich. Univ. 5: 755. 1928.
= Lecanora lisbonensis G. Samp., Brotéria, sér. Bot.
19: 33. 1921. – Lectotypus (hic designatus):
Portugal, [Estremadura], S. Martinho do Porto,
rochedos marítimos, 12 Oct. 1917, leg. A. Ricardo
Jorge (PO 1567L).
Crespo & al. • Coscinocladium, an overlooked lichen genus53 (2) • May 2004: 405–414
409
Fig. 4. Nuclear ITS rDNA majority-rule consensus tree based on 9600 trees from a B/MCMC tree sampling procedure.
Posterior probabilities equal to or above 0.95 are given above the branches.
≡ Psoroma lisbonense (G. Samp.) G. Samp., Lich. Port.
Exs. No. 85. 1923.
≡ Placodium lisbonensis (G. Samp.) Klem., Nova
Hedwigia 9: 488. 1965.
≡ Buellia lisbonensis (G. Samp.) Werner, Bull. Soc.
Bot. France 123: 438. 1976.
Etymology. —
The generic name is based on the
Greek words “kóskinon” (sieve) and “kládos” (branch).
The name is spelled as “Coscinacladium” in several pub-
lications (e.g., Poelt, 1969; Nimis & Poelt, 1987) but we
see no compelling orthographic reason to replace the “o”
of the original publication. See below for the origin of
the specific epithet “gaditanum”.
Description. —
Thallus crustose-placodioid,
orbicular, forming rosettes 1–2(–3) cm diam, often con-
fluent and forming extensive patches, frequently over-
growing other lichens, usually abundantly pruinose,
whitish grey to almost white, with a bluish tinge around
and in the central part. Peripheral lobes usually well-dif-
ferentiated, 1–2(–3) × 0.5–1 mm, contiguous to laterally
subimbricate, tips only slightly broadened, covered by a
thick, white, coarsely crystalline pruina, the pruina some-
times poorly developed towards the apices and then with
a whitish grey colouration. Central parts of the thallus
irregular and often uneven, with minute fissures delimit-
ing convex areoles. Soralia arising on the lobe surfaces,
starting to burst out at the base of the lobes as protruding
groups of soredia, later becoming well-circumscribed,
discrete, not confluent even where they are denser in the
older more central parts of the thallus, 0.2–0.25(–0.32)
mm diam, sometimes empty of soredia (most probably
after periods of heavy rain) and then appearing as con-
cave circular depressions. Soredia spherical, 50–55(–90)
µm diam, lead-grey to brownish grey, surface smooth,
covered with thin pruina. Cortex covered and inter-
spersed with coarse crystals, composed of hyphae
arranged perpendicular to the surface with the upper 1–3
cells brownish and ca 3.5 µm wide, the soredia also cov-
ered with a layer of brownish angular cells individually
6–10 µm diam covered by a variably developed layer of
minute hyaline crystals. Ascomata not known.
Conidiomata (reported by Werner 1976; see above) pyc-
nidial, immersed, globose. Conidiogenous cells pleu-
rogenous (Vobis, 1980: type VI). Conidia short-cylindri-
cal, simple, hyaline, 3.8–5 × 1.3 µm
1
.
Chemistry. —
Thallus and medulla K-, C-, KC-,
PD-, and I-. Zeorin has been detected by t.l.c. (G. Paz-
Bermúdez, specimen annotations in PO). We have been
able to confirm this, we have also found an unknown
compound in material from the Empúres (L’Escala,
Catalonia). The unknown substance belongs to RF class-
es 5–6 in Solvent A, 6 in B, and 7 in C (Elix & Ernst-
Russell, 1993), has a yellow fluorescence under 365 nm
UV-light. A yellow or dull yellow-red fluorescence was
also evident under the same UV light in most intact thal-
li checked.
Illustrations. —
Tavares (1956: 134, pl. 1, Figs
1–2), Martellos & Nimis (2003, in colour), Fig. 1.
Typifications. —
The original locality for
Variolaria gaditana given by Clemente (1807, Fig. 2)
was “frequentis, in muris Gades”. “Gades” is the latin-
ized version of “Gadir”, the Phoenician name for what is
now the city of Cádiz (Andalucía), reputedly founded
around 1100 BC and the oldest city in Europe (Williams,
2000). No original material could be located amongst
Clemente’s material in MAD, but what is certainly the
same species still grows on the walls of the old city in
Cádiz and we therefore designate a collection from the
original locality as neotype here. We also note that the
original place of publication of Clemente’s name was
incorrectly indicated by Zahlbruckner (1927–1928) to
have been in Acharius (1814: 133), despite Acharius hav-
ing correctly cited Clemente’s publication.
Interestingly, Kunze (1846a) gave the original local-
ity of Coscinocladium occidentale as “In urbe Gades ad
muros et saxa arenosa marina copiose”. Despite the sim-
ilarity in the localities, habitat, and actual description,
Kunze did not mention Clemente’s name at all here nor
in the subsequently separately printed version of the
work (1846b). Kunze’s herbarium in LZ was destroyed
in World War II, but three collections of this species were
sent by Kunze to E. M. Fries and are now preserved in
UPS. R. Santesson was in no doubt that they were the
same as the species called Lecanora lisbonensis in
Portugal, and he sent them on loan to Tavares (1956) who
concurred; we also agree that the three are conspecific.
UPS (L-82445) does not have the name on the original
label but is from “ad rupes mari propinq. pr. Gades leg.
Willkomm Kze.”; UPS (L-82639) is labelled only
“Coscinocladium occidentale m. pl. Willk. 987 Hispan.
Kze”; and UPS (L-74641) has the information “987(98)
Coscinoclad. (scrips. Kunze!) Coscinocladium occiden-
tale Kze! (spec. originale!) SüdSpanien, leg. Willkomm,
comm. Auerswald” and is from the “Herb. Rel. W. v.
Zwackh”. However, in this last collection, only “987(98)
Coscinoclad.” is in Kunze’s hand, the notation “987(89)”
being the entry (and collection) numbers in Kunze
(1846a, b). All three are on the same friable rock and are
probably parts of a single original collection. We select
as lectotype that which has the binomial in Kunze’s hand
as it is well-developed (five fragments) even though
Gades is not mentioned specifically.
In the original account of L. lisbonensis, numerous
Portuguese localities were mentioned by Sampaio (1921)
Crespo & al. • Coscinocladium, an overlooked lichen genus 53 (2) • May 2004: 405–414
410
1
We did not see any pycnidia, and Werner’s observations are in need of confirmation.
with varying degrees of detail, but it is unclear to what
extent these were actual collections or field observations.
Eight collections in PO were listed as “syntypes” in the
catalogue of Sampaio’s lichen types in PO compiled by
Paz-Bermúdez & al. (2002). We do not, therefore, repeat
that information here, but note that by an oversight mate-
rial distributed in Lich. Portugal No. 85, which was not
collected until January 1922, was stated to be a syntype,
and that the two collection numbers “991bL” and
“991aL” in their paper should be corrected to PO 5608bL
and PO5608a respectively. A total of 11 specimens
remain under this name in PO, but none of the pre-1921
packets are from Lisbon or its immediate vicinity. As lec-
totype we select a collection definitely cited in the proto-
logue, which is well-developed (six rock fragments, one
of which has no lichen) and lacks intermixed Lecanora
apothecia (see above).
Ecology. —
Usually on porous, soft, calcareous
rocks, including cemented sand or sandstones, more
rarely terricolous or on more compacted calcareous or
other alkaline rocks (e.g., metabasite). Also frequent on
mortar, plasterwork, and tiles in old buildings and walls.
Mainly along the coast, halotolerant on the seashore. It
generally occurs with species of Aspicilia, Caloplaca,
Diplotomma, Lecanora and (or) Verrucaria; examples of
relevé including the species are presented in Table 3.
Distribution. —
The currently known distribution
(Fig. 5) is certainly incomplete, but follows the western
Mediterranean coast and also the Atlantic coast from
Portugal (Beira Litoral) to Morocco (near Safi). The east-
ern limit appears to be around Linosa and Lampedusa in
southern Italy (Nimis, 2003). It reaches Macaronesia,
with a single record from the Selvages Islands (Tavares,
1958).
Identification. —
Coscinocladium gaditanum
could be confused with five superficially similar crus-
tose-placodioid lichens. (1) Diplotomma canescens
(Dicks.) Flotow (syn. Diploicia canescens (Dicks.)
DeNot.; see Molina & al., 2002) which has thicker, more
robust thalli with better developed marginal lobes, a
more greenish-grey colour, gradually coalescing (not
persistently discrete) soralia, and which is K+ yellow. (2)
Caloplaca teicholyta (Ach.) J. Steiner with ash-grey thal-
li, shorter and less well-demarcated lobes, the central
parts of the thallus being occupied by minute isidia with
no soralia. (3) Aspicilia radiosa (Hoffm.) Poelt &
Leuckert which has a grey thallus, usually with brown
apothecia, no soralia, and is K+ red. (4) Solenopsora
candicans (Dicks.) J. Steiner, with a purer white thallus,
lacking soredia and pruina, PD+ red, and usually with
apothecia producing 1-septate hyaline ascospores. And
(5) Buellia zoharyi Galun which is always on soil, fre-
quently sterile, but with pure white K+ red lobes.
DISCUSSION
The mtSSU rDNA has proved to be sufficiently con-
servative to elucidate relationships of lichen taxa at the
generic level (Crespo & al., 2001; Wedin & al., 2002).
On the basis of the morphological habit of the thallus, the
kind of photobiont, and the current systematic circum-
scription, the species was compared with other taxa from
Lecanorales suborders Lecanorinae (Lecanoraceae and
Physciaceae, including Caliciaceae; Wedin & Grube,
2002) and Teloschistinae (Teloschistaceae; Eriksson &
al., 2001). The mtSSU rDNA majority rule consensus
tree (Fig. 3) shows that C. gaditanum is not a member of
Lecanoraceae, but belongs in the Physcia group (Wedin
& al., 2002), along with species of Anaptychia,
Heterodermia, Phaeophyscia, Physconia, etc.
The three samples of C. gaditanum grouped (100
pp), but no relationship between those samples and the
others included was resolved in the mitochondrial tree.
Diplotomma canescens, a morphologically similar
species, is placed in the other monophyletic group (i.e.,
the Buellia group).
A well-supported clade (100 pp) included
Anaptychia as the sister group of Physconia and
Phaeorrhiza. A relationship between Anaptychia and
Physconia had already been suggested by Poelt (1965)
on morphological grounds and by Nordin & Mattson
(2001) and Grube & Arup (2001) on molecular charac-
Crespo & al. • Coscinocladium, an overlooked lichen genus53 (2) • May 2004: 405–414
411
Fig. 5. The distribution of Coscinocladium gaditanum.
Sources: Literature: Clemente (1807), Kunze (1846),
Colmeiro (1867), Sampaio (1921), Werner (1955), Tavares
(1956), Poelt (1958), Tavares (1958), Klement (1965), Poelt
(1969), Werner (1976), Llimona (1980), Houmeau & Roux
(1984), Llimona & Egea (1984), Nimis & Poelt (1987),
Breuss (1988), Boqueras & al. (1989), V
zda (1989),
Bricaud & Roux (1990), Nimis (1993), Nimis & al. (1994),
Lumbsch & Feige (1995, 1996), Egea & Llimona (1997),
Paz-Bermúdez & al. (2002), Nimis (2003), and unpub-
lished records of A. Crespo, A. Gómez Bolea, and X.
Llimona. Herbaria: HERBESS, PO, BC-Werner, BCN.
ters. Dirinaria applanata, a widespread tropical species
with lecanorine apothecia, was included in the study as
the genus has several placodioid species that have some
superficial resemblance to C. gaditanum. However, D.
applanata was included in the Buellia group. A well-sup-
ported clade (100 pp) places Diplotomma canescens as
the basal sister group to Dirinaria applanata and the
three species of Pyxine. Surprisingly, Diplotomma
canescens did not nest with D. venustum, a result incon-
gruent with previous ITS analyses (Molina & al., 2002)
and indicating that further studies on the relationships of
those species are required, perhaps utilising additional
genes.
We conclude that Coscinocladium belongs to
Physciaceae but is distinct from the other genera with
which we have been able to compare it at the molecular
level. However, it may be most closely allied to the rela-
tively recently described Central and North American
genus Mobergia H. Mayrhofer & Sheard (Mayrhofer &
al., 1992). Unfortunately no mtSSU rDNA sequences are
available for any species of that genus in GenBank and
no fresh material was available for extraction.
Nevertheless, the information from an ITS tree analysing
also several species of the Physcia group, including
Mobergia calculiformis and Physcia tenella var. mariti-
ma (the closest sequences in the BLAST search), does
not contradict the hypothesis that Coscinocladium is a
monophyletic independent genus. Moreover, both genera
are not very similar morphologically.
ACKNOWLEDGEMENTS
We are indebted to Prof. Dr. Rolf Santesson (Uppsala) and Dr.
R. Moberg (Uppsala) for sending us the Kunze material in UPS on
loan; Prof. Dr. J. A. Elix (Camberra) for sending important sam-
ples for phylogenetic analysis; Dr. Paloma Blanco (Madrid) for
providing photocopies of some historical papers; Dr. Mercedes
Barbero (Barcelona) for chemical data; Drs. Antonio Gómez-
Bolea, Pere Navarro-Rosinés (Barcelona), Claude Roux
(Marseille), and Prof. Dr. Guido B. Feige (Essen) for distribution-
al data and suggestions; Dr. Elisa Folhadela and Dr. Graciela Paz-
Bermúdez for their kind hospitality and arranging access to
Sampaio’s collections in Porto; and Ms. Clara Bencomo and Dr.
Javier Fernández de Bobadilla for their warm hospitality in Jerez
de la Frontera and for facilitating the arduous collection of the neo-
type in the old city of Cádiz. This work has been supported by
grants REN2001-1272/GLO and BOS 2001-0869-C04-02 of the
Ministerio de Ciencia y Tecnología de España, and was undertak-
en while D.L.H. and O. B. were supported, respectively, by the
Programa Ramón y Cajal and by a grant from the same ministry.
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12 3 45
Slope (º) 5 5 10 10 3
Exposure SE S N W W
Coverage (%) 80 90 95 90 75
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