The genus Catapyrenium s. lat. (Verrucariaceae) in the Iberian Peninsula and the Balearic Islands

Abstract

A taxonomic treatment of the genera included in Catapyrenium s. lat. in the Iberian Peninsula and the Balearic Islands is provided, based on study of c . 2000 specimens from both herbaria and fresh material collected by the authors from 2005 to 2009 in numerous localities.
A total of 33 species belonging to six of the eight genera included in Catapyrenium s. lat. are present in the region, which correspond to c . 80% of species from Europe. The genera are Anthracocarpon Breuss, Catapyrenium Flot. ( Catapyrenium s. str.), Heteroplacidium Breuss, Involucropyrenium Breuss, Neocatapyrenium H. Harada and Placidium A. Massal. The genus Neocatapyrenium is reported for the first time from this region. Clavascidium liratum , the only representative of the genus Clavascidium in the region, has been reduced to synonymy with Anthracocarpon virescens .
Heteroplacidium acervatum, H. congestum and Neocatapyrenium cladonioideum are new to Europe. Involucropyrenium pusillum , only previously known from Austria, I. waltheri , an arctic-alpine species, and Neocatapyrenium latzelii , known from Croatia and Greece, are cited for the first time in the Iberian Peninsula. Placidium subrufescens and P. tenellum, two mainly Mediterranean species, are also new records from the Iberian Peninsula. Placidium boccanum is here for the first time cited in Spain, and Anthracocarpon virescens , Heteroplacidium contumescens , Placidium imbecillum and P. michelii are new to Portugal. Most of the remaining species showed a considerable expansion of their known ranges. Placidium pyrenaicum is reduced to synonymy with P. velebiticum .
Keys to genera and species occurring in the Iberian Peninsula and the Balearic Islands are presented.

Full text

The genus Catapyrenium s. lat. (Verrucariaceae)inthe
Iberian Peninsula and the Balearic Islands
María PRIETO, Gregorio ARAGÓN and Isabel MARTÍNEZ
Abstract: A taxonomic treatment of the genera included in Catapyrenium s. lat. in the Iberian
Peninsula and the Balearic Islands is provided, based on study of c. 2000 specimens from both herbaria
and fresh material collected by the authors from 2005 to 2009 in numerous localities. A total of 33
species belonging to six of the eight genera included in Catapyrenium s. lat. are present in the region,
which correspond to c. 80% of species from Europe. The genera are Anthracocarpon Breuss, Catapy-
renium Flot. (Catapyrenium s. str.), Heteroplacidium Breuss, Involucropyrenium Breuss, Neocatapyrenium
H. Harada and Placidium A. Massal. The genus Neocatapyrenium is reported for the first time from this
region. Clavascidium liratum, the only representative of the genus Clavascidium in the region, has been
reduced to synonymy with Anthracocarpon virescens.Heteroplacidium acervatum, H. congestum and
Neocatapyrenium cladonioideum are new to Europe. Involucropyrenium pusillum, only previously known
from Austria, I. waltheri, an arctic-alpine species, and Neocatapyrenium latzelii, known from Croatia
and Greece, are cited for the first time in the Iberian Peninsula. Placidium subrufescens and P. tenellum,
two mainly Mediterranean species, are also new records from the Iberian Peninsula. Placidium
boccanum is here for the first time cited in Spain, and Anthracocarpon virescens,Heteroplacidium
contumescens,Placidium imbecillum and P. michelii are new to Portugal. Most of the remaining species
showed a considerable expansion of their known ranges. Placidium pyrenaicum is reduced to synonymy
with P. velebiticum. Keys to genera and species occurring in the Iberian Peninsula and the Balearic
Islands are presented.
Key words: Anthracocarpon,Catapyrenium,Clavascidium, distribution, ecology, Heteroplacidium,
Involucropyrenium, keys, lichens, Neocatapyrenium,Placidium
Introduction
The concept of ‘Catapyrenium s. lat.’ used
here includes squamulose, squamulose-
areolate or nearly crustose species; thalli are
heteromerous with a variable degree of dif-
ferentiation, and different types of tissues,
both in the upper cortex, medulla and lower
cortex. Attachment organs are mainly rhizo-
hyphae and rhizines (rhizinomorphs sensu
Malone 1977; Wagner & Letrouit-Galinou
1988). The photobiont is a green alga. Asco-
mata are perithecia, with involucrellum
present or not; asci are cylindrical or clavate,
8-spored, with ascospores simple and colour-
less. Pycnidia can be present or not and both
Dermatocarpon and Endocarpon types have
been observed in the group. Members of
Catapyrenium s. lat. occur mostly on soil,
sometimes directly on rocks or in crevices
and on ledges, and even living as parasites;
most species occur on calcareous substrata.
They are distributed mainly in temperate,
semiarid or cold areas of both hemispheres
(Breuss 1990a; Harada 1993; Breuss
1996a).
Traditionally, many species included in
Catapyrenium s. lat. have been placed in other
genera such as Endocarpon,Dermatocarpon or
Verrucaria (e.g. Endocarpon cinereum or
Dermatocarpon boccanum) (Persoon 1794;
Acharius 1810; Servı´t 1955). Nevertheless,
Breuss (1990a) emended the genus Cata-
pyrenium Flot. to accommodate only the
squamulose species with simple ascospores
and without hymenial algae, recognizing six
groups at infrageneric level (for a historical
M. Prieto, G. Aragón and I. Martı´nez: Área de Bio-
diversidad y Conservación, Universidad Rey Juan
Carlos, C/Tulipán s/n, 28933 Móstoles, Madrid,
SPAIN. Email: maria.prieto@urjc.es.
The Lichenologist 42(6): 637–684 (2010) © British Lichen Society, 2010
doi:10.1017/S0024282910000319

review on the early systematics of Catapy-
renium s. lat. see Breuss 1990a), and thus
recognising the considerable anatomical and
morphological variability present in the
group. Some years later, a total of eight dif-
ferent genera was segregated by Harada
(1993) and Breuss (1996a) based on combi-
nations of characters such as the type of
pycnidium, ascus shape and arrangement of
the ascospores, thallus anatomy and mor-
phology (structure of the upper cortex and
type of attachment organs) and presence or
absence of an involucrellum. These authors
recognized Anthracocarpon Breuss, Catapy-
renium Flot. (Catapyrenium s. str.), Clavascid-
ium Breuss, Heteroplacidium Breuss,
Involucropyrenium Breuss, Neocatapyrenium
H. Harada, Placidium A. Massal and Sclero-
pyrenium H. Harada.
Recent molecular studies have been car-
ried out in order to understand relationships
between members of Verrucariales at different
levels (e.g. Heiðmarsson 2003; Gueidan et al.
2007; Amtoft et al. 2008; Savic´ et al. 2008;
Prieto et al. 2010). The results of these mol-
ecular analyses have shown that the main
genera included in the concept of Catapy-
renium s. lat. (except the genus Clavascidium)
are independent and unrelated entities
(Gueidan et al. 2007, 2009; Savic´ et al.
2008), thus confirming that the variability of
the group is related to evolutionary patterns.
Catapyrenium s. lat. has been poorly
studied in the Iberian Peninsula with only
scattered references in some floristic studies
or in publications of some newly described
species (e.g. Breuss & Etayo 1992; van den
Boom & Giralt 1996; Navarro-Rosinés et al.
1996; Aragón 2003); although Breuss
(1990a) in his monograph of the genus
included this region, the study was mainly
focused on central Europe. As the group has
remained poorly understood in the Iberian
Peninsula, with little collected material, and
because of recent taxonomic changes in the
group, an exhaustive revision of all taxa in the
group in this region was necessary. The work
formed part of the ‘Flora Liquenólogica
Ibérica’ project.
Material and Methods
This study is based mainly on material collected by the
authors in the Iberian Peninsula and the Balearic Islands
between 2005 and 2009. The collection comprises
c. 1700 specimens deposited at MA.
In addition, Iberian collections from BCC, BCN, BP,
LEB, LISU, MA, MACB, MAF, SANT, TFC, VAL
and VIT and European and North American herbaria
ABL, ARIZ, ASU, B, BM, COLO, GB, H, HAL, L, LI,
NY, PRM, S and TUR were revised. Some personal
collections (C. Keller, C. Scheidegger and G. Aragón)
have also been revised. A total of c. 2000 specimens were
studied. A list of the material studied can be seen online
(http://www.escet.urjc.es/biodiversos/espa/personal/
isabel/isabel_e.htm).
Observations and measurements were made using a
Nikon SMZ-800 dissecting microscope and an Olympus
BX 51 microscope. The sections (14–20 m) of thalli
were made using a Leica CM 1850 UV freezing micro-
tome and were observed and measured in water and
sometimes in lactophenol cotton blue.
For anatomical studies, ten samples per species were
analysed (when available), and ten measurements of
each specimen on different squamules were carried out.
The limited material of some species and the poor con-
dition of others, led to a lower number of measurements
in some cases. Measurements are expressed as the
mean ± standard deviation (SD) with the extremes in
parentheses; length/breadth ratios (l/b) were calculated
for both ascospores and conidia. Only data from the
Iberian material were included in the descriptions. Fre-
quently, measurements of asci and paraphyses were not
enough for descriptive purposes, in these cases we in-
cluded published data. Most variables were normally
distributed, in cases where they were not normal, trans-
formations were carried out.
For the statistical analysis we conducted ANOVA in
order to understand the differences between closely re-
lated species using SPSS Statistics 17.0.
Distributional maps were drawn with ArcView GIS
3.1, based on UTM coordinates (WGS84 Datum).
For each taxa, we include data of the basionym, type
specimens and their location, but not previously pub-
lished synonyms (see Breuss 1990aand Harada 1993).
Key to genera
1 Perithecia between the squamules (sometimes in the margin), not immersed in the
thallus, with involucrellum .................Involucropyrenium
[Pycnidia not very common, Endocarpon-type]
Perithecia immersed in the thallus, without involucrellum ............2
638 THE LICHENOLOGIST Vol. 42

2(1) Upper cortex cinereum-type, thin (5–40 m) and poorly delimited from the algal
layer, with small, roundish-subangular cells of 4–10 m diam. ..........
...................................Catapyrenium
[Pycnidia absent]
Upper cortex clearly delimited from the algal layer, from 20 to c. 100 m thick, with
roundish-angular cells 4–15 m diam. ......................3
3(2) Pycnidia Endocarpon-type, rhizohyphae present or not, with rhizines or stipe-like
structures ......................................4
Pycnidia Dermatocarpon-type, rhizohyphae always present, rhizines present or not
...........................................5
4(3) Exciple and rhizines brown to carbonaceous, rhizohyphae dark, lower cortex absent
........................... Anthracocarpon virescens
[including Clavascidium liratum]
Exciple not carbonaceous, rhizohyphae absent, squamules attached to the substra-
tum by the basal ends or by rhizines, lower cortex clearly delimited from the
medulla ............................Neocatapyrenium
5(3) Squamules small, 0·3–3 mm, thallus entirely paraplectenchymatous or subpara-
plectenchymatous, upper cortex 20–55 m thick, with cells of 5–13 m, laminal
pycnidia, asci clavate ..................... Heteroplacidium
Squamules larger, (1) 2–8 (15) mm, thallus more differentiated, upper cortex
20–100 m thick, with cells of 5–17 m, laminal or marginal pycnidia, asci
cylindrical at least when young, clavate in some rhizinate species . Placidium
Taxonomic treatment
Anthracocarpon Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996); type: A. virescens (Zahlbr.) Breuss
[= Clavascidium liratum (Breuss) Breuss].
The thallus is squamulose, with an
anatomy very similar to the genus Placidium
but characterized by the presence of
Endocarpon-type pycnidia, perithecia with a
black to carbonaceous exciple (at least on the
top, e.g., A. caribaeum Breuss), and rhizo-
hyphae and rhizines as attachment organs.
Of the three species reported worldwide,
only A. virescens occurs in Europe.
Anthracocarpon virescens (Zahlbr.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Dermatocarpon virescens Zahlbr.,
Österr. Botan. Z.68: 69 (1919); type: Jugoslawien, Dal-
matien, West Meleda, erdige Kalkfelsspalten über Porto
Soline, c. 200 m, 17/03/1910, J. Baumgartner (W!—
holotype).
Clavascidium liratum (Breuss) Breuss, Ann. Naturhist.
Mus. Wien, Ser. B, Bot. Zool.98 (Suppl.):41 (1996).—
Catapyrenium liratum Breuss, Linzer biol. Beitr.23(2):
533 (1991); type: Mallorca, Sierra del Norte (Serra de
Tramuntana), Col de Hono zwischen Buñolá und
Alaró, G. B. Feige nº 6856 (HAL—holotype!; LI—
isotype!).
(Figs 1A & 2A)
Thallus squamulose; squamules up to
5 mm, scattered to contiguous, sometimes
overlapping, flattened to convex, rounded
to slightly lobed; upper surface cream-
coloured, pale, medium to dark brown,
with conspicuous black perithecial ostioles
(Fig. 1A); lower surface pale, brown or black
with dark rhizohyphae and carbonaceous
rhizines, simple or branched, one to several
per squamule. Anatomy: thallus (180)
235 ± 35 (325) m thick, with or with-
out epinecral layer [up to 45 m when present
sulcate or not (Fig. 2A)]; upper cortex
paraplectenchymatous, (37) 58 ± 15
(100) m thick, with cells (6) 9·5 ± 2·4
(15) m diam. Algal layer (52) 74 ± 15
(112) m thick, with cells (4) 6·7 ± 1·5
(13) m diam. Medulla prosoplectenchy-
matous, (45) 93 ± 30 (162) m thick, with
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 639

F. 1.Habit of some species of Catapyrenium s. lat. A, Anthracocarpon virescens;B,Catapyrenium psoromoides;
C, Heteroplacidium divisum;D,Involucropyrenium llimonae;E,Neocatapyrenium cladonioideum;F,Placidium
imbecillum.
640 THE LICHENOLOGIST Vol. 42

F. 2.Anatomy of Catapyrenium s. lat. species, A–E, cross sections of thalli; F cross-section of pycnidia.
A, Anthracocarpon virescens showing the sulcate epinecral layer; B, Catapyrenium psoromoides showing the cinereum-
type upper cortex (uc); C, Heteroplacidium divisum;D,Heteroplacidium imbricatum, showing two perithecia;
E, Involucropyrenium waltheri showing two perithecia with involucrellum (inv); F, Endocarpon-type pycnidia in
Neocatapyrenium cladonioideum.
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 641

elongated cells (2) 3·8 ± 0·9 (6) m diam.;
lower cortex absent. Rhizohyphae (2·5)
3·6 ± 0·7 (5) m, dark; rhizines up to 300 m
thick, simple or branched.
Perithecia up to 500 m wide, pyriform
to subglobose, with a brown to black and
carbonaceous exciple, 20–55 m thick. Asci
clavate, 60–90 × 14–19 m; ascospores biseri-
ate, simple, (14) 17·5 ± 1·6 (21) × (6)
7·4 ± 0·8 (9) m, l/b ratio (1·7) 2·4 ± 0·4
(3·5).
Pycnidia Endocarpon-type; conidia bacilli-
form, (5) 6·7 ± 0·9 (9) × (1) 1·1 ± 0·2 (1·5)
m, l/b ratio (4) 6·2 ± 1·3 (9).
Notes.Clavascidium liratum was described
by Breuss (1991) and differentiated from A.
virescens mainly by the presence of a thicker
and sulcate epinecral layer and a brown exci-
ple in C. liratum (rather than carbonaceous as
in A. virescens). Both shared a prosoplecten-
chymatous medulla, dark rhizohyphae and
black rhizines. The type material of C. lira-
tum was morphologically very similar to other
specimens of A. virescens although with the
upper surface paler.
Endocarpon-type pycnidia are present in A.
virescens, but they were not observed in the
type specimen of Clavascidium liratum; this
last species was known only from the type
locality in the Balearic Islands. Recently, we
collected material in this locality belonging to
the typical form of A. virescens. Two more
specimens collected in Portugal showed a
pale upper surface, with a sulcate epinecral
layer, and a non carbonaceous exciple, being
similar to C. liratum. These specimens pre-
sented Endocarpon-type pycnidia, until now
never observed in C. liratum.
A molecular analysis of these Portuguese
specimens, together with the type specimen
of C. liratum and two typical samples of A.
virescens showed that all formed a mono-
phyletic clade (results not shown). This
analysis supports the decision taken here
to synonymize Clavascidium liratum with
Anthracocarpon virescens.
Due to the presence of rhizines, the car-
bonaceous exciple and the colour and shape
of the squamules, A. virescens resembles
Endocarpon pusillum Hedw.; however, this
last species is characterized by peri-
thecia with hymenial algae and muriform
ascospores.
Ecology. The species shows preferences
for soil and rock ledges on calcareous and
gypsiferous substrata in shrublands with
Buxus sempervirens L., Lavandula latifolia
Medik., Lycium sp, Rosmarinus officinalis L.
and Thymus sp. in dry and open habitats; we
have also collected it in Pinus halepensis Mill.
forests. Anthracocarpon virescens has been
frequently found together with Placidiopsis
cinerascens (Nyl.) Breuss, P. custnani (A.
Massal) Körb. or Placidium pilosellum.
In the area studied, A. virescens was found
between 170 and 1200 m altitude.
Distribution. The species is relatively fre-
quent in xeric areas of the eastern Iberian
Peninsula, being also present in the Balearic
Islands (Fig. 3A). New data extend the
known distribution of the species in the
Iberian Peninsula, that was previously known
mainly from the east coast of Spain. The
specimen collected in Portugal is the first
record of A. virescens from this country.
Anthracocarpon virescens is widely distrib-
uted in the Mediterranean region, being
known from Croatia (Breuss 1990a), Greece
(Christensen et al. 1997; Sipman & Raus
1999), Italy (Nimis & Martellos 2004),
Spain (Breuss 1990a), Tunisia (Breuss
1994a) and Turkey (Breuss & John 2004;
Tufan et al. 2005), growing from sea level to
1400 m altitude.
Catapyrenium Flot.
Bot. Zeitung 8: 361 (1850); type: C. cinereum (Pers.)
Körb.
The thallus is composed of small squam-
ules, 1–4 mm wide, forming a more or less
continuous thallus; there is a cinereum-type
upper cortex, i.e. thin (5–40 m), poorly
delimited from the algal layer, paraplecten-
chymatous, with small, roundish-subangular
cells 4–10 m diam. (Fig. 2B). The algal
layer is not clearly delimited above and
below, being more or less diffuse; the algal
642 THE LICHENOLOGIST Vol. 42

cells are small (2·5–10 m). The asci are
clavate, 8-spored, with ascospores biseriate,
simple, colourless, 12–24 × 5–9 m. Occa-
sionally, we have found some pseudoseptate
ascospores (see Prieto et al. 2010). Pycnidia
absent. There are seven species world-wide,
F. 3.Distribution of Catapyrenium s. lat. species in Iberia. A, Anthrocarpon virescens;B,Catapyrenium cinereum;
C, C. daedaleum;D,C. psoromoides;E,Heteroplacidium acervatum (C); H. compactum (■), H. congestum ()and
H. contumescens (:); F, H. divisum (:)andH. phaeocarpoides (■).
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 643

after the recent exclusion of C. dactylinum
from the genus (Prieto et al. 2010). Four species are present in Europe, three of which
are here reported from the Iberian Peninsula.
Key to Catapyrenium species
1 Squamules with dark margins; lower cortex well-developed, paraplectenchymatous,
medulla composed of spherical hyphae; ascospores 14–24 × 6–9 m ......
....................................C. cinereum
Squamules without dark margins; lower cortex absent; medulla loose, composed of
a mixture of elongated and spherical hyphae ..................2
2(1) On soil; ascospores 15–24 × 5–9 m................C. daedaleum
On bark; ascospores 12–18 × 5–7 m ..............C. psoromoides
Catapyrenium cinereum (Pers.) Körb.
Syst. Lich. Germ.: 325 (1855).—Endocarpon cinereum
Pers., Nueue Ann. Bot.1: 28 (1794); type: Germania
[‘Reperi hanc speciem prope Schanzfeld: circa die alte
kirche & prope montem Meisner’] (H-ACH nº 851 –
type?!).
Thallus squamulose; squamules tightly con-
tiguous, forming a continuous thallus, small,
(0·5) 1–3 mm wide, finely lobulate; upper
surface whitish, rarely greenish grey or
brownish, pruinose, with dark margins;
lower surface black, with dark rhizohyphae.
Anatomy: thallus (100) 198 ± 45 (312) m
thick, with or without epinecral layer, up to
30 m when present; upper cortex (5) 18 ± 7
(35) m thick, poorly delimited from the
algal layer, with small, roundish-subangular
cells (5) 6·7 ± 0·9 (8) m diam. Algal layer
75–175 m thick, distributed over almost the
entire thallus, with scattered cells or in clus-
ters, algal cells (3) 5·3 ± 1 (8) m diam.
Medulla up to 125 m thick, composed of
globular cells 3–12 m diam.; lower cortex
paraplectenchymatous, blackish, (7) 19 ± 5
(30) m thick, with roundish-angular cells in
two or three layers, (7) 8·6 ± 1·3 (12) m
diam. Rhizohyphae thin, (3) 3·8±0·6 (5) m,
brown to black.
Perithecia up to 350 m wide, subglobose;
exciple initially pale, later brown to black, up
to 30 m diam. Asci clavate, c. 65–
70 × 16–22 m (Breuss 1993), ascospores
biseriate, simple (occasionally pseudosep-
tate), (14) 19·4 ± 1·9 (24) × (6) 7·6 ± 0·9
(9) m, l/b ratio (1·7) 2·6 ± 0·4 (3·3).
Pycnidia absent.
Notes. The type was not located with
certainty, but a specimen in H-ACH (nº 851)
named as Endocarpon tephroides from
‘Germania’ and annotated as ‘Endocarpon
cinereum Pers.’ probably represents the type.
Catapyrenium cinereum,C. daedaleum and
C. psoromoides, are the only members of
Catapyrenium s. str. present in the Iberian
Peninsula. These three species have very
similar anatomical characters; however, C.
cinereum is easily recognized by the usually
whitish, dark rimmed squamules, and is the
only one with a lower cortex clearly delimited
and a medulla composed of globular cells.
Catapyrenium daedaleum also differs in the
longer and wider ascospores, and in thallus
thickness (thinner in C. cinereum); see
Table 1 for further differences. Catapyrenium
psoromoides is easily differentiated by its
corticolous habit.
Ecology.Catapyrenium cinereum grows pre-
dominantly in alpine grasslands, over calcif-
ereous soils or in fissures, frequently mixed
with mosses, but is also found in Juniperus
thurifera L. forests. The species occurs at
high altitudes, from 1270 to 2350 m,
together with Catapyrenium daedaleum and
Involucropyrenium waltheri.
Distribution.Catapyrenium cinereum is
found in montane areas of the Iberian
Peninsula, mainly in northern Spain, but
with occurrences in the central and southern
mountains (Fig. 3B).
Records from Andorra, Granada, La Rioja
(Casares & Llimona 1984; Azuaga &
644 THE LICHENOLOGIST Vol. 42

T 1.Results from ANOVA comparing similar species. Measurements are expressed as mean ± standard deviation in µm and ranges in parentheses
Catapyrenium cinereum C. daedaleum P Heteroplacidium acervatum H. contumescens P
Thallus thickness (100) 198±45 (312) (162) 242±48 (375) *** (260) 337±44 (410) (200) 297±64 (400) ns
Epinecral layer (0) 11±6·6 (30) (2) 9·4±4·9 (25) * up to 15 up to 10 -
Upper cortex (5) 18±7 (35) (5) 19±8 (40) ns (25) 35±6 (45) (25) 40±8 (55) ns
Upper cortex cells (5) 6·7±0·9 (8) (5) 6·3±0·9 (8) ns (6) 8±1·1 (10) (5) 7·3±1·2 (11) *
Algal layer (75) 117±33 (175) (62) 95±23 (150) ns (70) 123±28 (175) (50) 104±31 (175) ns
Algal layer cells (3) 5·3±1 (8) (2·5) 5±1·1 (7) * (5) 8±2·2 (15) (6) 8·4±1·5 (12) ns
Medulla (30) 71±33 (125) (37) 77±29 (140) ns (112) 174±65 (300) (80) 132±44 (237) ns
Medulla cells (3) 7·9±1·8 (12) (3) 5·3±1·3 (8) *** (5) 7·6±1·4 (10) (5) 7·9±1·3 (10) ns
Lower cortex (7) 19±5 (30) – – – – –
Rhizohyphae (3) 3·8±0·6 (5) (2·5) 3·5±0·8 (6) ** (3) 3·9±0·4 (5) (4) 4·5±0·7 (6) **
Ascospore length (14) 19·4±1·9 (24) (15) 18·5±1·7 (24) ** (11) 12·3±0·9 (13) (11) 12·6±1 (15) ns
Ascospore width (6) 7·6±0·9 (9) (5) 6·2±0·9 (9) *** (5·5) 5·8±0·2 (6) (5·5) 6·6±0·8 (8) **
Ascospore length/breadth
ratio
(1·7) 2·6±0·4 (3·3) (2) 3±0·5 (4·2) *** (1·8) 2·1±0·2 (2·4) (1·4) 1·9±0·3 (2·7) *
Conidium length – – – (4) 5·5±0·6 (7) (3) 3·7±0·5 (4·5) ***
Conidium width – – – (1) 1·2±0·2 (1·5) (1) 1·3±0·2 (1·5) *
Conidium length/breadth
ratio
– – – (3·3) 4·9±0·9 (6) (2) 3±0·6 (4) ***
Heteroplacidium divisum H. imbricatum P Placidium fingens P. squamulosum P
Thallus thickness (100) 149±26 (200) (150) 203±38 (300) *** (240) 284±31 (330) (240) 343±45 (450) ns
Epinecral layer up to 10 up to 10 – (5) 13·7±7·5 (25) (0) 20·6±11·8 (50) ns
Upper cortex (20) 28±6 (45) (22) 30±5 (40) ns (35) 52±13 (77) (37) 70±16 (100) *
Upper cortex cells (5) 8·2±1·8 (12) (4) 7·9±2·4 (13) ns (8) 11±2 (15) (6) 10·9±2·5 (17) ns
Algal layer (30) 60±17 (100) (57) 90±21 (137) *** (75) 107±25 (150) (62) 113±24 (162) ns
Algal layer cells (6) 8·7±1·6 (12·5) (5) 8·7±2 (12) ns (6) 8·9±1·9 (13) (6) 9±1·6 (14) ns
Medulla (25) 60±19 (87) (50) 83±26 (132) ** (37) 72±29 (137) (62) 119±30 (175) *
Lower cortex – – – (17) 41±16 (62) (25) 43±13 (80) ns
Lower cortex cells – – – (9) 11·1±1·9 (15) (9) 11·9±1·7 (15) ns
Rhizohyphae (3) 3·9±0·7 (5) (3) 4·5±0·7 (5) * (5) 5·2±0·4 (6) (5) 5·8±0·5 (7) ns
Ascospore length (11) 12·5±1 (14) (11) 14±1·7 (17) * (12) 15·4±2·2 (18) (12) 14·4±1·3 (17) ns
Ascospore width (6) 6±0·6 (7) (5) 6±0·6 (7) ns (5) 6±0·5 (7) (6) 6·7±0·5 (8) ***
Ascospore length/breadth
ratio
(1·8) 2±0·2 (2·3) (1·6) 2·4±0·4 (3·2) * (1·9) 2·6±0·5 (3·3) (1·6) 2·2±0·3 (2·8) ***
Conidium length (3·5) 4·1±0·4 (5) – – (4) 4·5±0·6 (6) (2·5) 3·4±0·5 (5) ***
Conidium width (1) 1·2±0·2 (1·5) – – (1) 1·2±0·2 (1·5) (1·3) 1·8±0·2 (2) ***
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 645

T 1.Continued
Heteroplacidium divisum H. imbricatum P Placidium fingens P. squamulosum P
Conidium length/breadth
ratio
(2·3) 3·6±0·8 (4·5) – – (2·7) 3·9±0·6 (5) (1·25) 2±0·4 (2·9) ***
Placidium imbecillum P. rufescens P Placidium imbecillum P. subrufescens P
Thallus thickness (190) 279±44 (380) (300) 410±64 (600) *** (190) 279±44 (380) (300) 377±41 (450) ***
Epinecral layer (0) 14·6±7·8 (40) (0) 12·4±6·5 (40) ns (0) 14·6±7·8 (40) (2·5) 13·4±5·8 (25) ns
Upper cortex (28) 54±14 (88) (33) 61±12 (88) ns (28) 54±14 (88) (37) 62±17 (95) ns
Upper cortex cells (5) 10·4±2·6 (16) (5) 10±2·7 (18) ns (5) 10·4±2·6 (16) (5) 9·7±2·8 (17) ns
Algal layer (42) 100±25 (175) (63) 110±27 (200) ns (42) 100±25 (175) (62) 103±21 (150) ns
Algal layer cells (4) 8·8±2·1 (15) (7) 10±1·4 (14) * (4) 8·8±2·1 (15) (6) 8·7±1·9 (13) ns
Medulla (30) 70±19 (107) (75) 154±44 (275) *** (30) 70±19 (107) (63) 134±26 (200) ***
Lower cortex (20) 41±14 (87) (38) 81±19 (138) *** (20) 41±14 (87) (25) 47±16 (87) ns
Lower cortex cells (6) 11·4±2·6 (16) (8) 13·8±2·7 (23) ** (6) 11·4±2·6 (16) (5) 11·7±3·4 (18) ns
Rhizohyphae (4) 5·1±0·6 (6) (5) 6·6±0·8 (8) *** (4) 5·1±0·6 (6) (4) 4·6±0·8 (6) ns
Ascospore length (9) 13·1±1·7 (18) (14) 17·8±1·8 (22) *** (9) 13·1±1·7 (18) (9) 12·3±1·9 (17) *
Ascospore width (5) 6·1±0·7 (8) (6) 7·9±1 (11) *** (5) 6·1±0·7 (8) (5) 5·7±0·5 (7) *
Ascospore length/breadth
ratio
(1·5) 2·2±0·3 (3) (1·5) 2·3±0·4 (3·5) ns (1·5) 2·2±0·3 (3) (1·6) 2·2±0·4 (3·2) ns
Conidium length (3) 4·2±0·9 (6) (3) 3·7±0·5 (5) ** (3) 4·2±0·9 (6) (3) 5±0·9 (7) ***
Conidium width (0·5) 1·1±0·2 (1·5) (1) 1·8±0·3 (2·5) *** (0·5) 1·1±0·2 (1·5) (1) 1·1±0·1 (1·5) ns
Conidium length/breadth
ratio
(2) 4±1 (6) (2) 2·1±0·5 (3) *** (2) 4±1 (6) (3) 4·6±1 (7) *
Placidium lacinulatum P. squamulosum P Placidium michelii P. squamulosum P
Thallus thickness (220) 321±48 (440) (240) 343±45 (450) ns (160) 200±24 (240) (240) 343±45 (450) ***
Epinecral layer (0) 8·6±4·8 (25) (0) 20·6±11·8 (50) *** (0) 8·4±4 (20) (0) 20·6±11·8 (50) ***
Upper cortex (37) 67±14 (92) (37) 70±16 (100) ns (25) 45±11 (67) (37) 70±16 (100) ***
Upper cortex cells (6) 8·6±1·5 (13) (6) 10·9±2·5 (17) *** (8) 11·5±2·1 (17) (6) 10·9±2·5 (17) ns
Algal layer (55) 95±21 (155) (62) 113±24 (162) ** (37) 69±16 (105) (62) 113±24 (162) ***
Algal layer cells (5) 9·2±1·8 (15) (6) 9±1·6 (14) ns (6) 9·5±1·5 (12) (6) 9±1·6 (14) ns
Medulla (62) 146±36 (227) (62) 119±30 (175) * (37) 76±19 (125) (62) 119±30 (175) ***
Lower cortex – (25) 43±13 (80) – (25) 37±16 (63) (25) 43±13 (80) ns
Lower cortex cells (5) 8·3±1·4 (11) (9) 11·9±1·7 (15) *** (10) 12·4±1·8 (16) (9) 11·9±1·7 (15) ns
Rhizohyphae (3) 4·3±0·6 (5) (5) 5·8±0·5 (7) *** (4) 4·7±0·7 (6) (5) 5·8±0·5 (7) ***
646 THE LICHENOLOGIST Vol. 42

T 1.Continued
Placidium lacinulatum P. squamulosum P Placidium michelii P. squamulosum P
Ascospore length (11) 13·4±1·2 (16) (12) 14·4±1·3 (17) ** (10) 12·6±1·3 (15 (12) 14·4±1·3 (17) ***
Ascospore width (6) 7·4±0·6 (9) (6) 6·7±0·5 (8) *** (4) 5·5±0·5 (6) (6) 6·7±0·5 (8) ***
Ascospore length/breadth
ratio
(1·4) 1·8±0·2 (2·7) (1·6) 2·2±0·3 (2·8) *** (1·7) 2·3±0·4 (3·1) (1·6) 2·2±0·3 (2·8) ns
Conidium length (3) 3·8±0·5 (5) (2·5) 3·4±0·5 (5) * (3) 3·3±0·3 (4) (2·5) 3·4±0·5 (5) ns
Conidium width (1·2) 1·7±0·3 (2) (1·3) 1·8±0·2 (2) ns (1·2) 1·4±0·1 (1·5) (1·3) 1·8±0·2 (2) **
Conidium length/breadth
ratio
(1·5) 2·2±0·4 (3·3) (1·25) 2±0·4 (2·9) ns (2) 2·3±0·3 (2·9) (1·25) 2±0·4 (2·9) ns
Placidium pilosellum P. rufescens P Placidium pilosellum P. squamulosum P
Thallus thickness (210) 270±30 (330) (300) 410±63 (600) *** (210) 270±30 (330) (240) 343±45 (450) ***
Epinecral layer (7·5) 20·7±8·9 (62·5) (0) 12·4±6·5 (40) *** (7·5) 20·7±8·9 (62·5) (0) 20·6±11·8 (50) ns
Upper cortex (22) 54±14 (95) (33) 61±12 (88) ns (22) 54±14 (95) (37) 70±16 (100) ***
Upper cortex cells (6) 10±1·9 (14) (5) 10±2·7 (18) ns (6) 10±1·9 (14) (6) 10·9±2·5 (17) ns
Algal layer (62) 90±14 (125) (63) 110±27 (200) ** (62) 90±14 (125) (62) 113±24 (162) ***
Algal layer cells (5) 8·6±1·7 (14) (7) 10±1·4 (14) ** (5) 8·6±1·7 (14) (6) 9±1·6 (14) ns
Medulla (62) 105±23 (170) (75) 154±44 (275) *** (62) 105±23 (170) (62) 119±30 (175) ns
Lower cortex (18) 37±18 (56) (38) 81±19 (138) *** (18) 37±18 (56) (25) 43±13 (80) ns
Lower cortex cells (7) 10·3±1·9 (14) (8) 13·8±2·7 (23) ** (7) 10·3±1·9 (14) (9) 11·9±1·7 (15) ns
Rhizohyphae (4) 4·9±0·5 (6) (5) 6·6±0·8 (8) *** (4) 4·9±0·5 (6) (5) 5·8±0·5 (7) ***
Ascospore length (12) 14·2±1·4 (17) (14) 17·8±1·8 (22) *** (12) 14·2±1·4 (17) (12) 14·4±1·3 (17) ns
Ascospore width (5) 6·5±0·5 (7·5) (6) 7·9±1 (11) *** (5) 6·5±0·5 (7·5) (6) 6·7±0·5 (8) ns
Ascospore length/breadth
ratio
(1·7) 2·2±0·3 (3·2) (1·5) 2·3±0·4 (3·5) * (1·7) 2·2±0·3 (3·2) (1·6) 2·2±0·3 (2·8) ns
Conidium length (2) 3·2±0·5 (4) (3) 3·7±0·5 (5) ** (2) 3·2±0·5 (4) (2·5) 3·4±0·5 (5) ns
Conidium width (1·2) 1·7±0·2 (2) (1) 1·8±0·3 (2·5) * (1·2) 1·7±0·2 (2) (1·3) 1·8±0·2 (2) ns
Conidium length/breadth
ratio
(1·3) 2±0·3 (2·7) (2) 2·1±0·5 (3) ns (1·3) 2±0·3 (2·7) (1·25) 2±0·4 (2·9) ns
Placidium rufescens P. squamulosum P Placidium rufescens P. subrufescens P
Thallus thickness (300) 410±64 (600) (240) 343±45 (450) *** (300) 410±64 (600) (300) 377±41 (450) ns
Epinecral layer (0) 12·4±6·5 (40) (0) 20·6±11·8 (50) *** (0) 12·4±6·5 (40) (2·5) 13·4±5·8 (25) ns
Upper cortex (33) 61±12 (88) (37) 70±16 (100) ns (33) 61±12 (88) (37) 62±17 (95) ns
Upper cortex cells (5) 10±2·7 (18) (6) 10·9±2·5 (17) ns (5) 10±2·7 (18) (5) 9·7±2·8 (17) ns
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 647

T 1.Continued
Placidium rufescens P. squamulosum P Placidium rufescens P. subrufescens P
Algal layer (63) 110±27 (200) (62) 113±24 (162) ns (63) 110±27 (200) (62) 103±21 (150) ns
Algal layer cells (7) 10±1·4 (14) (6) 9±1·6 (14) ns (7) 10±1·4 (14) (6) 8·7±1·9 (13) *
Medulla (75) 154±44 (275) (62) 119±30 (175) *** (75) 154±44 (275) (63) 134±26 (200) ns
Lower cortex (38) 81±19 (138) (25) 43±13 (80) *** (38) 81±19 (138) (25) 47±16 (87) ***
Lower cortex cells (8) 13·8±2·7 (23) (9) 11·9±1·7 (15) ** (8) 13·8±2·7 (23) (5) 11·7±3·4 (18) **
Rhizohyphae (5) 6·6±0·8 (8) (5) 5·8±0·5 (7) *** (5) 6·6±0·8 (8) (4) 4·6±0·8 (6) ***
Ascospore length (14) 17·8±1·8 (22) (12) 14·4±1·3 (17) *** (14) 17·8±1·8 (22) (9) 12·3±1·9 (17) ***
Ascospore width (6) 7·9±1 (11) (6) 6·7±0·5 (8) *** (6) 7·9±1 (11) (5) 5·7±0·5 (7) ***
Ascospore length/breadth
ratio
(1·5) 2·3±0·4 (3·5) (1·6) 2·2±0·3 (2·8) ns (1·5) 2·3±0·4 (3·5) (1·6) 2·2±0·4 (3·2) ns
Conidium length (3) 3·7±0·5 (5) (2·5) 3·4±0·5 (5) ns (3) 3·7±0·5 (5) (3) 5±0·9 (7) ***
Conidium width (1) 1·8±0·3 (2·5) (1·3) 1·8±0·2 (2) ns (1) 1·8±0·3 (2·5) (1) 1·1±0·1 (1·5) ***
Conidium length/breadth
ratio
(2) 2·1±0·5 (3) (1·25) 2±0·4 (2·9) ns (2) 2·1±0·5 (3) (3) 4·6±1 (7) ***
Placidium rufescens P. velebiticum P Placidium squamulosum P. tenellum P
Thallus thickness (300) 410±64 (600) (240) 312±47 (430) *** (240) 343±45 (450) (200) 283±50 (400) ***
Epinecral layer (0) 12·4±6·5 (40) (0) 5·2±2·3 (10) *** (0) 20·6±11·8 (50) (2·5) 21·8±11 (42·5) ns
Upper cortex (33) 61±12 (88) (25) 49±9 (67) *** (37) 70±16 (100) (45) 66±14 (100) ns
Upper cortex cells (5) 10±2·7 (18) (7) 10·6±2·5 (16) ns (6) 10·9±2·5 (17) (8) 11·5±2·2 (15) ns
Algal layer (63) 110±27 (200) (55) 85±18 (125) *** (62) 113±24 (162) (40) 93±24 (155) *
Algal layer cells (7) 10±1·4 (14) (4) 9±2·4 (13) ** (6) 9±1·6 (14) (6) 9·1±1·5 (12) ns
Medulla (75) 154±44 (275) (70) 137±43 (250) ns (62) 119±30 (175) (47) 91±24 (150) *
Lower cortex (38) 81±19 (138) (27) 49±13 (77) *** (25) 43±13 (80) (25) 35±17 (62) ns
Lower cortex cells (8) 13·8±2·7 (23) (6) 13±3 (20) ns (9) 11·9±1·7 (15) (7) 9·9±2 (12) *
Rhizohyphae (5) 6·6±0·8 (8) (4) 5·4±0·6 (6) *** (5) 5·8±0·5 (7) (3) 4±0·6 (5) ***
Ascospore length (14) 17·8±1·8 (22) (11) 14±1·3 (17) *** (12) 14·4±1·3 (17) (11) 13·2±1·1 (15) **
Ascospore width (6) 7·9±1 (11) (5) 6·3±0·5 (7) *** (6) 6·7±0·5 (8) (5) 5·9±0·7 (7·5) ***
Ascospore length/breadth
ratio
(1·5) 2·3±0·4 (3·5) (1·7) 2·2±0·2 (2·8) ns (1·6) 2·2±0·3 (2·8) (1·6) 2·3±0·3 (3) ns
Conidium length (3) 3·7±0·5 (5) (2·5) 3·9±0·7 (5) ns (2·5) 3·4±0·5 (5) (2·5) 3·2±0·5 (4) ns
Conidium width (1) 1·8±0·3 (2·5) (1) 1·7±0·4 (2) * (1·3) 1·8±0·2 (2) (1·5) 1·8±0·2 (2) ns
Conidium length/breadth
ratio
(2) 2·1±0·5 (3) (1·2) 2·5±0·9 (4·2) ** (1·25) 2±0·4 (2·9) (1·3) 1·8±0·4 (2·7) ns
*** = P<0·001; ** = P<0·01; * = P<0·05; ns = no significant differences.
648 THE LICHENOLOGIST Vol. 42

Gómez-Bolea 1998; Burgaz et al. 2007) and
Portugal (Sampaio 1924; Jansen 1993) could
not be revised and are not included in the
map (Fig. 4B).
Catapyrenium cinereum is an arctic-alpine
taxon, found in Africa, Asia, Australia,
Europe, New Zealand and North and South
America (Breuss 1993, 2001a, 2002a;
Harada 1993; Egea 1996).
Catapyrenium daedaleum (Kremp.)
Stein
In Cohn, Krypt.-Fl. Schlesien (Breslau)2(2):312
(1879).—Endocarpon daedaleum Kremp., Flora 38: 66
(1855); type: BRD, Bayern, Berchtesgadener Alpen,
Steinthal und Vorderberg zwischen Hochkalter und
Kammerlinghorn, c. 5800–6000', 1854, Rauchenberger
(M—lectotype); BRD, Karwendel (bei Mittenwald),
5009', 1849, Krempelhuber (WU—syntype).
Thallus squamulose; squamules tightly con-
tiguous, adnate or with slightly raised mar-
gins, forming a continuous thallus, small,
1–4 mm wide, finely crenulate; upper surface
faintly pruinose, greenish grey to brownish,
never darker at margins; lower surface dark,
with dark rhizohyphae. Anatomy: thallus
(162) 242 ± 48 (375) m thick; upper cortex
(5) 19 ± 8 (40) m thick, poorly delimited
from the algal layer, with small, roundish-
subangular cells of (5) 6·3 ± 0·9 (8) m
diam.; epinecral layer up to 25 m. Algal
layer not clearly delimited, diffuse, c. 62–
150 m thick, with algal cells of (2·5) 5 ±
1·1 (7) m diam. Medulla up to c. 140 m
thick, composed mainly of elongated cells
of (3) 5·3 ± 1·3 (8) m diam., loose, becom-
ing darker below; lower cortex lack-
ing. Rhizohyphae dark, (2·5) 3·5 ± 0·8
(6) m.
Perithecia up to 300 m wide, subglobose;
exciple up to 30 m, hyaline to brown, darker
at the ostiole. Asci clavate, c. 75–85 × 17–20
m (Breuss 1993); ascospores biseriate, sim-
ple (occasionally pseudoseptate), (15)
18·5 ± 1·7 (24) × (5) 6·2 ± 0·9 (9) m, l/b
ratio (2) 3 ± 0·5 (4·2).
Pycnidia absent.
Notes. Many herbarium specimens ident-
ified as C. daedaleum were found to be C.
cinereum. Both species can sometimes be
confused because of their similar thallus
morphology, but the lack of a lower cortex in
C. daedaeleum is a good diagnostic character
(see above for further details).
Ecology.Catapyrenium daedaleum lives pre-
dominantly in alpine grasslands, on calcare-
ous soils and in rock fissures, mixed with
bryophytes; it has also been collected in
Fagus sylvatica L. and Juniperus thurifera for-
ests. The species was collected from 1000 to
2300 m altitude, occasionally with Cata-
pyrenium cinereum and Involucropyrenium
waltheri.
Distribution.Catapyrenium daedaleum
presents now a wider distribution than pre-
viously known in the Iberian Peninsula
(Aragón 2003). The species has a similar
distribution to C. cinereum, as both species
occur in mountainous areas; nevertheless,
C. daedaleum is more common in the
Mediterranean region than C. cinereum
(Fig. 3C). It has been recently recorded in
Portugal for the first time by Paz-Bermúdez
et al. (2009).
According to Breuss (1993) and Nimis &
Martellos (2004) the species has a more pro-
nounced arctic-alpine distribution than C.
cinereum, although in the Iberian Peninsula
we have observed the opposite pattern,
C. daedaleum being more common in the
southern mountains than C. cinereum.
C. daedaleum has an arctic-alpine distri-
bution, known from Asia (Aptroot et al.
2003), Europe (Breuss 1990a), North
America (Goward et al. 1994; revised in this
study, hb. COLO) and South America
(Breuss 1993).
Catapyrenium psoromoides (Borrer)
R. Sant.
In Hawksworth, James & Coppins Lichenologist 12: 106
(1980).—Verrucaria psoromoides Borrer, in Hooker &
Sowerby, Engl. Bot. [Suppl. 1, tab. 2612, Fig. 1] (1831);
type: England, on elm at Hurstpierpoint and on ash at
Beeding, Sussex, Borrer (K—holotype).
(Figs 1B & 2B)
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 649

F. 4.Distribution of Catapyrenium s. lat. species in Iberia. A, Heteroplacidium imbricatum;B,Involucropyrenium
llimonae (:), I. nuriense (C), I. pusillum (■), I. tremniacense(), I. waltheri (), Neocatapyrenium cladonioideum (O)
and N. latzelii (+); C, Placidium adami-borosi (■), P. boccanum (:)andP. fingens (C); D, P. imbecillum;
E, P. lachneum;F,P. lacinulatum.
650 THE LICHENOLOGIST Vol. 42

Thallus squamulose; composed of tightly
contiguous to slightly overlapping squamules
forming a continuous thallus; squamules
adnate and lobate, up to 4 mm wide; upper
surface greyish brown to brown (Fig. 1B),
pruinose (mostly on external parts of the
lobes); lower surface with blackish-
brown rhizohyphae. Anatomy: thallus (140)
211 ± 58 (330) m thick; upper cortex (5)
16·8 ± 8·2 (32·5) m thick, paraplectenchy-
matous, with roundish-subangular cells (4)
6·9 ± 1·5 (10) m diam.; epinecral layer thin,
up to 15 m. Algal layer not clearly delim-
ited, diffuse, c. 45–112 m thick, with cells
(4) 6·4 ± 1·5 (10) m diam. Medulla loose,
(50) 72 ± 18 (175) m thick, composed
mainly of elongated hyphae of (3) 5·2 ± 1·4
(8) m diam., with some spherical cells and
interhyphal spaces; lower cortex lacking.
Rhizohyphae brown to black, (3) 3·4 ± 0·5
(4) m.
Perithecia pyriform, up to 200 m wide;
exciple hyaline to pale brown, darker on the
ostiole, up to c.25m thick; asci clavate,
55–65 × 13-15 m (Breuss 1990a), asco-
spores biseriate, hyaline, simple (occasionally
pseudoseptate), (12) 15 ± 1·6 (18) × (5)
5·6 ± 0·6 (7) m, l/b ratio (2·1) 2·7 ± 0·4
(3·6).
Pycnidia absent.
Notes.Catapyrenium psoromoides is the only
corticolous species of Catapyrenium s. str. It
is very similar to C. daedaleum from which it
differs in its significantly smaller ascospores
(15 ± 1·6 × 5·6 ± 0·6 minC. psoromoides
versus 18·5 ± 1·7 × 6·2 ± 0·9 minC. dae-
daleum) and in its ecology and distribution
(C. daedaleum is terricolous and arctic-alpine
whereas C. psoromoides is corticolous and
from temperate climates).
Ecology.Catapyrenium psoromoides is a ni-
trophilous species growing on ancient trees
in open forests (Aragón & Sarrión 2003); we
have found it on Fraxinus angustifolia Vahl,
Juniperus thurifera,Olea europaea L. and on
different species of Quercus (Q. faginea Lam.,
Q. faginea subsp. broteroi (Cout.) A. Camus,
Q. ilex subsp. ballota (Desf.) Samp., Q.
pyrenaica Willd. or Q. robur L.). The species
has been collected from 500 to 1400 m alti-
tude.
Although C. psoromoides has been found
growing only on trees in the Iberian
Peninsula, in other parts of Europe the
species also occurs on mosses (Breuss
1990a).
Distribution. We recorded few specimens
only of C. psoromoides in the Iberian
Peninsula, maybe because the species can be
overlooked due to its small size. Based on its
habitat preferences, we suspect the species
probably has a wider distribution. Despite
this, new data extend the distributional area
of the species in the Iberian Peninsula
(Fig. 3D). Records from Navarra (Etayo &
Breuss 1996), Palencia (López de Silanes
et al. 1998) and Zaragoza (Etayo et al. 1993)
could not be localized and are not included
on the map.
Catapyrenium psoromoides is known from
temperate Asia, eastern Africa, Europe, New
Zealand and SW North America (Breuss
2002a).
Heteroplacidium Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996); type: H. imbricatum (Nyl.) Breuss.
The thallus is crustose-areolate to
squamulose, composed of small squamules
or areoles (1–3 mm), with a thallus anatomy
entirely paraplectenchymatous or subpara-
plectenchymatous (Fig. 2C). Perithecia are
conspicuous and generally have the exciple
turning brown when mature. Asci clavate
with biseriate ascospores, pycnidia are lami-
nal of Dermatocarpon-type and with oblong-
ellipsoidal to cylindrical conidia. Differ-
ences from Placidium are mainly based on
squamule size, thallus anatomy and asci. Dif-
ferences between species are few because of
the weak differentiation of tissues and
consequently lack of diagnostic characters.
The species grow on soil and rock
(although some parasitic species have been
recently included in the genus), living in
warm-temperate regions.
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 651

Until now, ten species have been reported
worldwide, six of which were present in
Europe. As a result of this study the number
of species rises to eight in Europe, seven of
which are present in the Iberian Peninsula.
Key to Heteroplacidium species
1 Thallus areolate to subsquamulose, epilithic, parasitic or not . . H. compactum
[Conidia bacilliform (5–7 m long), ascospores 11–18 × 8–10 m, medulla (sub)paraplectenchymatous]
Thallus squamulose, epilithic or not, never parasitic ................2
2(1) Squamules densely imbricate ............................3
Squamules not densely imbricate ..........................4
3(2) Thallus 100–200 m thick, ascospores 11–14 × 6–7 m .......H. divisum
Thallus 150–300 m thick, ascospores 11–17 × 5–7 m .... H. imbricatum
4(2) Squamules very small, 0·5–1 mm ............... H. phaeocarpoides
[ascospores 11–13 × 8–10 m, conidia oblong-ellipsoidal]
Squamules up to 3 mm ...............................5
5(4) Conidia bacilliform, 4–7 × 1–1·5 m, ascospores 11–13 × 5–6 m, rhizohyphae 3–5
m wide .............................. H. acervatum
Conidia oblong-ellipsoid to subcylindrical, 3–4·5 × 1–1·8 m, ascospores 11–
15 × 5·5–8 m long, rhizohyphae up to 6·5 m wide ..............6
6(5) Ascospores 12–15 × 8–10 m, rhizohyphae 5–6·5 m wide ....H. congestum
Ascospores 11–15 × 5–5–8 m, rhizohyphae 4–6 m wide . . H. contumescens
Heteroplacidium acervatum (Breuss)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Catapyrenium acervatum Breuss,
Linzer biol. Beitr.28: 525 (1996); type: España, Islas
Baleares, Mallorca, Serra de Llevant, Puig de sa Font bei
Son Cervera, 200–270 m, 15/04/1987, O. Breuss (W—
holotype; LI—isotype!).
Thallus epilithic, composed of tightly
contiguous to sometimes overlapping
squamules, forming compact cushion-like
aggregations; squamules 0·3–3 mm wide,
irregularly rounded to slightly lobate, con-
vex; upper surface pale brown to brown,
matt, greyish towards the margins; lower sur-
face brown, with bundles of rhizohyphae
forming a stipe-like organ. Anatomy:thallus
(260) 337 ± 44 (410) m thick; upper cortex
(25) 35 ± 6 (45) m thick, paraplectenchy-
matous, with small roundish-angular cells
(6) 8 ± 1·1 (10) m diam.; epinecral layer thin
or lacking, up to 15 m when present. Algal
layer 70–175 m, with cells of (5) 8 ±
2·2 (15) m diam. Medulla (112) 174 ± 65
(300) m thick, composed of spherical hy-
phae of (5) 7·6 ± 1·4 (10) m diam., the
lower ones turning brownish; lower cortex
not clearly delimited from the medulla.
Rhizohyphae hyaline, (3) 3·9± 0·4 (5) m,
aggregated in central parts of the squamules
giving the aspect of a stipe.
Perithecia (sub)globose, up to 350 m
wide, exciple up to 30 m, pale except at the
ostiole which is brown. Asci clavate, 50–
60 × 11–15 m; with biseriate ascospores,
simple, (11) 12·3 ± 0·9 (13) × (5·5) 5·8 ±
0·2 (6) m, l/b ratio (1·8) 2·1 ± 0·2 (2·4).
Pycnidia laminal; conidia bacilliform, (4)
5·5 ± 0·6 (7) × (1) 1 ± 0·2 (1·5) m, l/b
ratio (3·3) 4·9 ± 0·9 (6).
Notes. This species is characterized by its
growth form of compact cushions of aggre-
gated squamules; although we collected
three specimens which did not show this
typical form. Because their anatomical char-
acters are similar to H. acervatum as well as
molecular data (M. Prieto et al. unpub-
lished), we classify these three specimens as
H. acervatum. A few specimens from Murcia
652 THE LICHENOLOGIST Vol. 42

(MUB) with the typical growth form of com-
pact cushions of aggregated squamules and
previously classified as H. contumescens be-
long to H. acervatum.
Heteroplacidium acervatum and H. contu-
mescens, can be differentited mainly by the
shape of conidia (cylindrical, 5·5 ± 0·6 m
long in H. acervatum and shortly cylindrical
to oblong, 3·7 ± 0·5 minH. contumescens),
and width of the ascospores (5·8 ± 0·2 min
H. acervatum versus 6·6 ± 0·8 minH. con-
tumescens); moreover, the rhizohyphae are
thinner in H. acervatum (3·9 ± 0·4 m versus
4·5 ± 0·7 minH. contumescens) (but see
Table 1 for further differences). The
cushion-like growth of H. acervatum is very
characteristic, but not always present, as
mentioned above.
Ecology.Heteroplacidium acervatum grows
directly on calcareous rocks or over a slight
accumulation of soil; it has been collected in
xeric areas dominated by Chamaerops humilis
L., from 200 to 400 m altitude.
Distribution. Until now the species has
been found in very few localities in the Bal-
earic Islands (Breuss 1996b); we have found
it sparsely distributed in Murcia province,
located in south-eastern Spain (Fig. 3E), and
is recorded here for the first time in the
Iberian Peninsula and also continental
Europe.
Heteroplacidium compactum (A.
Massal.) Gueidan & Cl. Roux
Bull. Ass. Fr. Lichénologie 33(1):25 (2008).—Placidium
compactum A. Massal., Misc. Lichenol.: 62 (1856); type:
Germany, Arnold.
Thallus epilithic, areolate to subsquamu-
lose, areoles up to 2 mm diam., angular to
rounded, flat to slightly convex; upper sur-
face medium to dark brown, dull to some-
what shiny. Anatomy: thallus up to 600 m
thick; upper cortex 10–25 m thick, with
cells 5–8 m diam., the uppermost layer with
a brown tinge; epinecral layer up to 20 m
thick. Algal layer 100–200 m thick, with
cells 8–15 m diam. Medulla of globular cells
of 7–10 m thick; lower cortex not clearly
delimited. Rhizohyphae 4–6 m.
Perithecia nearly spherical, 250–400 m
wide, with pale exciple, darkening with age.
Asci clavate 55–70 × 13–20 m; ascospores
11–18 × 8–10 m.
Pycnidia laminal; conidia bacilliform,
5–7 × c. 1·5 (2) m.
Notes. Recently, Verrucaria compacta and
V. fuscula have been combined into the genus
Heteroplacidum (Gueidan et al. 2007; Roux
2008). The two species have been considered
to be the same taxon by some authors
(Fröberg 1989; Santesson 1993), with H.
compactum the autotrophic form of H.
fusculum; or they have been retained as differ-
ent species by other authors (Clauzade &
Roux 1985; Breuss 1994b). Breuss (1994b)
conducted a detailed investigation and con-
cluded that the main differences between the
two taxa were as follows. Heteroplacidium
compactum has bacilliform conidia (5–7 m
long), longer ascospores (11–18 m), a
cellular texture in the medulla, and is either
autotrophic or is parasitic on various crus-
tose lichens. In contrast, H. fusculum has
shortly cylindrical conidia (3–5 m long), a
medulla with some cylindrical hyphae,
shorter ascospores (9–14 m) with a thick
wall (1 m), and is parasitic on Aspicilia
calcarea (L.) Körb., or later becoming
autotrophic.
Only three specimens we revised corre-
sponded clearly to H. compactum with bacil-
liform conidia and long ascospores, and
were either parasitic or non-parasitic. The
remainder showed characters intermediate
between H. compactum and H. fusculum; for
example they were parasitic on Aspicilia cal-
carea with long ascospores but with either
shortly cylindrical or longer, bacilliform
conidia. Nevertheless, all the specimens liv-
ing on A. calcarea had a thick ascospore
wall. These observations lead us to doubt
the distinction between these two species,
but more material is necessary in order to
clarify this question.
Few measurements could be made, due to
the small number of specimens available;
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 653

consequently, the description above is based
on published data (Breuss 1994b).
Ecology. The species has been collected
from acid rocks and lava; between 30 and
1000 m altitude.
Distribution.Heteroplacidium compactum is
very sparsely distributed in the Iberian
Peninsula, occurring only in eastern Spain
(Fig. 3E). The species has been previously
reported from Asia, Europe and northern
Africa (Breuss 1994b).
Heteroplacidium congestum (Breuss &
McCune) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Catapyrenium congestum Breuss &
McCune, Bryologist 97: 365 (1994); type: USA, Idaho,
Owihee Co, Atriplex confertifolia-Artemisia spinescens
flats, Castle Creek Road, 4·5 km NW of junction with
Mudflat Road, 1095 m, vi 1991, McCune (OSC—
holotype; LI —isotype!).
Thallus densely squamulose or subver-
ruculose; squamules small, 0·5–2 mm wide,
tightly contiguous and scarcely overlapping,
rounded to slightly lobate, flat to slightly
convex; upper surface medium to blackish
brown, matt; lower surface pale. Anatomy:
thallus 250–400 m thick, upper cortex
20–40 m, of roundish angular cells 5–9 m
diam., epinecral layer of 10–20 m. Algal layer
70–100 m, with cells of 6–13 m diam.
Medulla composed of globular cells of
6–10 m diam.; lower cortex hardly discern-
ible, of more closely packed cells. Rhizo-
hyphae colourless, thick-walled, 5–6·5 m
diam.
Perithecia subglobose, up to 400 (500) m
wide; exciple pale when young, dark brown at
maturity. Asci clavate, 65–85 × 20–25 m;
ascospores biseriate, broadly ellipsoid to sub-
globose, 12–15 × 8–10 m.
Pycnidia laminal,conidia oblong ellipsoid
to subcylindrical, 3–4 × 1·5–1·8 m.
Notes. The combination of densely aggre-
gated (but not overlapping) small squamules,
thick walled rhizohyphae and broadly ellip-
soidal ascospores distinguish this species
from H. acervatum and H. contumescens,
which have narrower ascospores and thinner
rhizohyphae.
The material collected in the Iberian
Peninsula was too sparse for detailed exami-
nation, so details of morphology and
anatomy are based on Breuss & McCune
(1994).
Ecology.Heteroplacidium congestum has
been collected on calcareous soils in dry open
habitats together with Placidium squamulo-
sum. The species was found at 380 m alti-
tude.
Distribution. The species is recorded here
for the first time in the Iberian Peninsula
(Fig. 3E), constituting the first record from
Europe; it was previously known only from
North America, where it was considered to
be endemic.
Heteroplacidium contumescens (Nyl.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Endocarpon contumescens Nyl.,
Flora 61: 341 (1878); type: Algeria, Biskra, 1878, Norrlin
(H-NYL!—lectotype; H-NYL!—isotypes).
(Fig. 3E)
Thallus squamulose; squamules 1–3 mm
wide, scattered or contiguous to hardly over-
lapping, rounded to slightly lobate, flat or
with deflexed to revolute margins; upper sur-
face pale to medium brown, matt; lower sur-
face pale to brown. Anatomy: thallus (200)
297 ± 64 (400) m thick, upper cortex (25)
40 ± 8 (55) m thick, paraplectenchyma-
tous, with small roundish-angular cells (5)
7·3 ± 1·2 (11) m diam.; epinecral layer thin
or lacking, up to 10 m when present. Algal
layer (50) 104 ± 31 (175) m, with cells (6)
8·4 ± 1·5 (12) m diam. Medulla (80)
132 ± 44 (237) m, composed of spheri-
cal hyphae of (5) 7·9 ± 1·3 (10) m diam.,
the lower ones turning brownish; lower
cortex not clearly delimited from the
medulla. Rhizohyphae colourless, (4) 4·5 ±
0·7 (6) m.
654 THE LICHENOLOGIST Vol. 42

Perithecia globose to subglobose, up to
400 m wide; exciple up to 30 m, pale or
darkening at maturity. Asci clavate, 60–
70 × 15–18 m; ascospores biseriate, simple,
(11) 12·6 ± 1 (15) × (5·5) 6·6 ± 0·8 (8) m,
l/b ratio (1·4) 1·9 ± 0·3 (2·7).
Pycnidia laminal; conidia oblong-ellipsoid
to subcylindrical, (3) 3·7 ± 0·5 (4·5) × (1)
1·3 ± 0·2 (1·5) m, l/b ratio (2) 3 ± 0·6 (4).
Notes. Differences between H. contumes-
cens and H. acervatum have been previously
discussed under that species and are shown
in Table 1. Some herbarium specimens
revised in this study were previously ident-
ified as H. imbricatum or H. divisum, but both
these species are easily distinguishable from
H. contumescens by their imbricate squam-
ules. The species could also be confused with
H. compactum or H. fusculum, but these are
areolate-subsquamulose and with wider
ascospores than H. contumescens, and are
parasitic or free-living.
Ecology.Heteroplacidium contumescens has
been collected on sandy soils or directly on
rock with an accumulation of soil; the species
can live on both calcareous or acid substrata
and even on volcanic rocks; it was collected
predominantly in dry habitats from 200 to
700 m altitude.
Distribution. The species occurs mainly in
coastal areas of south-eastern Spain, with
some occurrences in central regions of the
Iberian Peninsula. We have collected it for
the first time in Portugal (Fig. 3E).
Most of the samples studied here are from
herbaria and were collected from 1975 to
1990; during our field trips very few speci-
mens of H. contumescens were collected pos-
sibly because of habitat destruction and
modification that has affected the south-east
Mediterranean coast. For this reason, we be-
lieve that H. contumescens should be treated
as an endangered species in the Iberian
Peninsula, or at least a detailed population
study of this species should be carried out.
Heteroplacidium contumescens is a rare
species with scattered occurrences in
Australia, central Asia and the Mediterranean
region (Breuss 2001b).
Heteroplacidium divisum (Zahlbr.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Dermatocarpon divisum Zahlbr.,
Österr. Bot. Z.59: 349 (1909); type: Jugoslawien,
Dalmatien, Ragusa (Dubrovnik), Mokosica, c. 200 m,
21 ix 1908, A. Latzel (W!—lectotype; PRM—isotype).
(Figs 1C & 2C)
Thallus composed of densely imbricate
squamules; squamules small and thin,
1–3 mm wide and c. 0·2 mm thick, flat to
undulate, entire to crenulate (Fig. 1C), with
margins free and without rhizohyphae; upper
surface greyish or greenish to pale or medium
brown, somewhat pruinose; lower surface
whitish, usually blackening, attached by
rhizohyphae accumulated at the centre of the
squamules. Anatomy: thallus (100) 149 ± 26
(200) m wide, upper cortex (20) 28 ± 6
(45) m thick, with roundish angular cells
(5) 8·2 ± 1·8 (12) m diam.; epinecral layer
very thin or lacking. Algal layer (30) 60 ± 17
(100) m thick, with cells (6) 8·7 ± 1·6
(12·5) m diam. Medulla (25) 60 ± 19 (87)
m thick, with many globular cells (6)
7·9 ± 2·0 (15) m diam.; lower cortex not
clearly delimited from the medulla. Rhizo-
hyphae colourless to slightly brownish,
(3) 3·9 ± 0·7 (5) m, usually concentrated
on the middle of the squamule, giving the
appearance of a stipe.
Perithecia (sub)globose, up to 500 m
wide; exciple pale to brown, darkened
towards the ostiole, up to 25 m thick. Asci
clavate, with biseriate ascospores, simple,
(11) 12·5 ± 1 (14) × (6) 6 ± 0·6 (7) m, l/b
ratio (1·8) 2 ± 0·2 (2·3).
Pycnidia laminal; conidia subcylindrical,
(3·5) 4·1 ± 0·4 (5) × (1) 1·2 ± 0·2 (1·5) m,
l/b ratio (2·3) 3·6 ± 0·8 (5).
Notes.Heteroplacidium divisum and H.
imbricatum are the only species in the genus
with imbricate squamules. Of the two,
H. divisum has significantly thinner squa-
mules (149 ± 26 m versus 203 ± 38 min
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 655

H. imbricatum) and consequently the algal
layer and medulla are also thinner; in
addition, ascospores are shorter in H.
divisum (12·5 ± 1 minH. divisum versus 14
±1·7 minH. imbricatum) (see Table 1 for
further differences). Some herbarium speci-
mens belonging to H. imbricatum have pre-
viously been erroneously identified as H.
divisum.
Ecology.Heteroplacidium divisum has been
collected on calcareous rocks, in fissures and
cavities. Most specimens were collected in
semiarid or dry regions and only the speci-
men from the Balearic Islands was found in a
very humid place on damp rocks; from 100 to
600 m altitude.
Distribution. The species is very scarce in
the Iberian Peninsula, with scattered occur-
rences in the south-east coast of Spain and
the Balearic Islands (Fig. 3F). Heteroplacid-
ium divisum is very rare worldwide, distrib-
uted mainly in Mediterranean regions,
occurring in Croatia, France and Turkey
(Breuss 1990a,b, 1998). A specimen revised
by us from LI represents a new record from
Italy.
Heteroplacidium imbricatum (Nyl.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Endocarpon imbricatum Nyl., Bot.
Not.1853: 161; type: Frankreich, Dept. Hérault, ad
Mireval prope Monspelium (H-NYL!—lectotype,
H-NYL!—isotype).
Thallus squamulose, composed of densely
imbricate squamules; squamules 1–3 mm
wide, rounded to lobate or crenulated,
slightly curved to concave or convex, some-
times curved up with free margins and with-
out rhizohyphae; upper surface greyish or
greenish brown to brown, somewhat pru-
inose at margins; lower surface whitish to
pale brown, attached by rhizohyphae accu-
mulated in the centre of the squamule.
Anatomy: thallus (150) 203 ± 38 (300) m
thick; upper cortex (22) 30 ± 5 (40) m
thick, with roundish angular cells of (4)
7·9 ± 2·4 (13) m diam.; epinecral layer very
thin or lacking, up to 10 m when present.
Algal layer (57) 90 ± 21 (137) m thick, with
cells (5) 8·7 ± 2 (12) m diam. Medulla (50)
83 ± 26 (132) m thick, composed of globu-
lar cells (4) 7·6 ± 1·7 (11) m diam.;
lower cortex not clearly delimited from
the medulla. Rhizohyphae colourless, (3)
4·5 ± 0·7 (5) m, accumulated on the mid-
dle of the squamule, giving the appearance of
a stipe.
Perithecia (sub)globose, up to 350 m
wide, exciple pale to brown, darkening
towards the ostiole, up to 25 m thick
(Fig. 2D). Asci clavate; ascospores biseriate,
simple, (11) 14 ± 1·7 (17) × (5) 6 ± 0·6
(7) m, l/b ratio (1·6) 2·4 ± 0·4 (3·2).
Pycnidia laminal; conidia subcylindrical,
4–5 × c.1m.
Notes. This species has often been con-
fused with members of the genus Placidium,
from which it differs mainly in the habitus,
with squamules distinctly imbricate, and
smaller than in Placidium; moreover, Hetero-
placidium has an entirely (sub)paraplecten-
chymatous thallus (Fig. 2C).
Specimens reported from the provinces
Zamora (Terrón et al. 2000) and Asturias
(Aragón et al. 2007) belong to Placidium ten-
ellum and Placidium sp., respectively. One
record from Almerı´a (Egea & Llimona 1981)
proved to be H. contumescens.
Ecology.Heteroplacidium imbricatum occurs
mainly on calcareous rocks with some soil
accumulation, and in fissures or cavities, fre-
quently sharing a habitat with Placidium
rufescens; from 100 to 850 m altitude.
Distribution.Heteroplacidium imbricatum is
a common species in coastal areas of
Mediterranean Spain, with some records
from the more interior provinces and the
Balearic Islands (Fig. 4A). New data re-
ported here extends the distributional area of
the species in both Spain and Portugal. On
the other hand, records from Alicante and
Madrid (Barreno & Merino 1981; Alonso
et al. 1989) could not be verified and the
species has not been collected by us in these
provinces.
656 THE LICHENOLOGIST Vol. 42

The species is not very common world-
wide, known only from southern Europe,
Macaronesia and northern Africa (Breuss
1988).
Heteroplacidium phaeocarpoides (Nyl.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):40 (1996).—Endocarpon phaeocarpoides Nyl.,
Flora 62: 358 (1879); type: Frankreich, Dépt. Bouches-
du-Rhone, in regione Massiliensi, Le Ciotat, 06/1879,
A. Taxis (H-NYL!—holotype).
Thallus squamulose; squamules very small
(0·5–1 mm), scattered to contiguous, flat;
upper surface greyish brown, matt, pruinose.
Anatomy: thallus 150–250 m thick, upper
cortex 20–30 m thick, paraplectenchyma-
tous, with cells 7–11 m diam.; epinecral layer
thin, c.10m. Algal layer occupying almost
the entire thallus, algal cells 8–19 m diam.
Medulla undifferentiated; lower cortex of
roundish cells. Rhizohyphae colourless,
4–5 m.
Perithecia up to 450 m wide, globose;
exciple pale. Asci oblong-clavate, 60–
70 × 16–20 m; ascospores biseriate, simple,
11–13 × 8–10 m.
Pycnidia laminal; conidia oblong-ellipsoid,
2·5–3·5 × 1–1·5 m.
Notes. The species has been reported once
from Spain (Breuss 1990b); however, this
specimen was not found, and no further
specimens have been found during field
work. The measurements are based on the
reference cited.
Ecology.Heteroplacidium phaeocarpoides
was reported as growing on sandy soils at
350 m altitude in a semi-arid region (Breuss
1990b).
Distribution. Only one record has been
reported from Almerı´a (Breuss 1990b)
(Fig. 3F). Although this specimen has not
been revised by us we have included it in the
map as it constitutes the only one from this
region.
The species is known from France, Spain,
Tunisia (Breuss 1994a), Macaronesia
(Boom & Etayo 2006), North Africa (Breuss
1994a) and Ukraine (M. Prieto et al., unpub-
lished, revised by us from herbarium LI).
Involucropyrenium Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):38 (1996); type: I. waltheri (Kremp.) Breuss.
The thallus is composed of small squam-
ules (0·1–3 mm), thin (50–300 m), and
with a thallus anatomy similar to the genus
Catapyrenium (i.e. upper cortex of cinereum-
type, medulla not clearly defined, algal layer
occupying almost the entire thallus, clavate
asci, biseriate ascospores, etc). The main
diagnostic characters for this genus are the
position of the perithecia, situated between
the squamules (see Fig. 1D) and the pres-
ence of an involucrellum (complete, dimidi-
ate or apical) (Fig. 2E); pycnidia have not
been observed except in I. squamulosum
(Brand & Van den Boom) Breuss, and were
described as unilocular (Endocarpon-type)
(Roux 2005).
The species occur on calcareous and
gypsiferous soils, in rock fissures or directly
on limestones, also on artificial substrata in
ruderal conditions, such as old bricks or
mortar. They occupy semi-arid to alpine and
temperate environments.
The genus is distributed mainly in Europe,
with a single species present in North
America or Asia (I. waltheri). It comprises
eight species, five of which are reported here
from the Iberian Peninsula. Two species (i.e.
I. llimonae and I. nuriense) are known only
from this region.
The genus must be regarded as poorly
known, since several species are known only
from their type localities, or in no more than
a handful of sites, and are probably often
confused with species of Verrucaria.
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 657

Key to Involucropyrenium species
1 Involucrellum partial, not enclosing the entire perithecium ............2
Involucrellum entire, enclosing the perithecium ..................3
2(1) Thallus epilithic, of densely imbricate squamules; upper cortex not paraplectenchy-
matous, with cells of irregular shape; involucrellum dimidiate, only reaching
half-way down the perithecia; exciple colourless, ascospores 11–14 × 7–9·5 m.
.....................................I. nuriense
Thallus consisting of more or less dispersed squamules; upper cortex paraplecten-
chymatous, with roundish-angular cells; involucrellum apical; exciple brown to
black; ascospores 13–18 × 6–8 m...............I. tremniacense
3(1) Rhizohyphae dark, hypothallus well developed ............ I. waltheri
[Ascospores 15–23 × 7.5–11 m. Artic-alpine distribution]
Rhizohyphae colourless, without hypothallus ...................4
4(3) Thallus consisting of dispersed squamules with a very characteristic shape, more or
less irregular to digitate-coralloid; involucrellum dark brown, 30-60 m thick;
ascospores 14–22 × 7–11 m.....................I. llimonae
Thallus of more crowded squamules, sometimes overlapping; involucrellum black,
25–35 m, thicker in the upper part; ascospores 19–24 × 9–14 m .......
.....................................I. pusillum
Involucropyrenium llimonae (Etayo,
Nav.-Ros. & Breuss) Breuss
Beitr. Naturk. Oberösterreichs 13: 213 (2004).—
Catapyrenium llimonae Etayo, Nav.-Ros. & Breuss, Can-
dollea 51: 140 (1996); type: España, Navarra, Rada, la
Bárdena Blanca, sobre suelos yesı´fero-arcillosos, 300–
400 m, 28 v 1990 J. Etayo (LI!—holotype).
Thallus squamulose; squamules small, (0·1)
0·3–0·5 (0·7) mm, dispersed, irregularly
shaped to digitate-coralloid; upper surface
brown. Anatomy: thallus (50) 70–150 m
thick; upper cortex 10-20 m thick,
paraplectenchymatous, with roundish-
subangular cells (3) 4–7 m; epinecral layer
thin 10–15 m. Algal layer occupying almost
the entire thallus, with cells of 6–10 m diam.
Medulla and lower cortex absent. Rhizo-
hyphae colourless, 3–5 m.
Perithecia dispersed to aggregated, devel-
oped between the squamules (Fig. 1D), glo-
bose to slightly conical, 0·2–0·4 mm. Exciple
colourless to brown, 15–20 m with a com-
plete dark brown involucrellum, 30–60 m
thick. Asci clavate, 55–80 × 15–21 m;
ascospores biseriate, broadly ellipsoid, (14)
16–18·2–21 (22) × (7) 9–9·7–11 m
(Navarro-Rosinés et al. 1996).
Pycnidia not seen.
Notes. This species is easily recognized by
the irregular to digitate-coralloid squam-
ules. Involucropyrenium pusillum is a similar
species but with a thicker involucrellum and
larger ascospores (19–24 × 9–14 m). Two
further species, I. terrigenum (Zschacke)
Breuss and I. sbarbaronis (Servı´t) Breuss
(neither of which has been found in the
Iberian Peninsula) have smaller asco-
spores than I. llimonae (15–20 × 6–8 m
in I. sbarbaronis; 10–12 × 7–8·5 minI.
terrigenum) and a nearly crustose thallus
rather than clearly dispersed squamules as
in I. llimonae. Only one specimen has been
found up to now in the Iberian Peninsula;
for this reason we have included the
measurements based on Navarro-Rosinés
et al. (1996).
Ecology.Involucropyrenium llimonae has
been collected on very eroded and weathered
gypsiferous and clayey soils, at the base of
shrubs.
Distribution. This species has been found
only in the type locality, situated in the north-
ern part of Spain, in a semi-arid environment
at 300–400 m altitude (Fig. 4B).
658 THE LICHENOLOGIST Vol. 42

Involucropyrenium nuriense
(Nav.-Ros. & Breuss) Breuss
Beitr. Naturk. Oberösterreichs 13: 214 (2004).—
Catapyrenium nuriense Nav.-Ros. & Breuss, Candollea
51: 144 (1996); type: España, Cataluña, Prov. Girona,
Ripollés, Queralbs, Vall de Núria, Forat de l’Embut,
cerca del Pla dels Eugassers, 2300–2400 m, paredes
verticales de roca calcárea orientados al norte, 1 viii
1984, P. Navarro-Rosinés (BCC!—holotype).
Thallus epilithic, almost continuous, 0·5–2
cm diam.; squamules densely imbricate, (0·5)
1–2 (3) mm wide, irregularly shaped to sub-
lobulate. Anatomy: thallus 150–200 m
thick; upper cortex 30–40 m thick, not
clearly paraplectenchymatous, with cells ir-
regularly arranged and elongated, (3) 5–7·5
(8·5) × (1·5) 2–4 m; epinecral layer absent
or very thin (6–12 m). Algal layer 100–150
m, with algal cells 7–10 m diam. Medulla
not differentiated; lower cortex 15 m thick,
dark brown, cells with thick walls. Rhizo-
hyphae colourless, 4 m thick.
Perithecia globose, 0·2–0·5 (0·7) mm, dis-
persed to aggregated, developed in the mar-
ginal region of the squamules, rarely in the
centre; exciple colourless, 20–30 m thick;
involucrellum black, dimidiate (only reach-
ing to the middle of the perithecia); periphyses
30–40 × 1·5 m. Asci clavate 55–65 × 15–20
m; ascospores biseriate, ovoid, broadly ellip-
soid or subglobose, 11–12·5–14 × (7) 8·2–9
(9·5) m (Navarro-Rosinés et al. 1996).
Pycnidia not seen.
Notes.Involucropyrenium nuriense is the
only species in the genus without a clearly
paraplectenchymatous upper cortex. More-
over, the involucrellum is dimidiate, not
entire, a character that approaches I. tremnia-
cense, although this species has an apical
involucrellum. The two species also differ
in ascospore size (11–14 × 7–9·5 minI.
nuriense versus 13–18 × 6–8 minI. tremnia-
cense) and in their ecology, I. tremniacense
being a terricolous species growing exclu-
sively in the Mediterranean region (see
below).
Only one specimen has been found in the
Iberian Peninsula and for this reason we have
included the measurements based on
Navarro-Rosinés et al. (1996) in the descrip-
tion.
Ecology.Involucropyrenium nuriense is a
saxicolous species and has been found at very
high altitudes (2300–2400 m), on vertical
limestone facing N-NE.
Distribution. The species is known only
from the type locality, collected in the eastern
Pyrenees, north east of Spain (Fig. 4B).
Involucropyrenium pusillum Breuss &
Türk
Beitr. Naturk. Oberösterreichs 13: 214 (2004); type:
Österreich, Oberösterreich, Puglalm SE vom Hengst-
pass, 920 m, auf Moosen über Kalkblöcken, 5 x 1987,
S. Wagner (LI!—holotype).
Thallus squamulose; squamules small, 0·2–
0·5 mm wide, greenish brown, matt, con-
tiguous to aggregated and often overlapping,
slightly lobulate, closely adnate to the sub-
stratum or with slightly raised margins.
Anatomy: thallus 100–150 m thick, almost
entirely (sub)paraplectenchymatous; upper
cortex not always clearly delimited, up to
20 m thick, with cells 3–8 m diam.; epi-
necral layer lacking or very thin, up to
10 m when present. Algal layer occupy-
ing almost the entire thallus, with algal cells
5–8 m diam. Medulla and lower cortex not
differentiated. Rhizohyphae hyaline, thin,
2·5–3 m diam.
Perithecia developed between the squam-
ules, sometimes in groups, nearly globose,
0·25–0·40 mm diam. Exciple hyaline to
brown, c.25m thick, ostiole paler. Involu-
crellum entire, black, 25–35 m thick,
thicker in the upper part. Periphyses 20–
30 × 1–1·5 m. Asci clavate, 75–85 × 20–25
m; ascospores (19) 21·2 ± 1·7 (24) × (9)
12·2 ± 1·3 (14) m.
Pycnidia absent.
Notes. The ascospores of the Iberian
material are slightly wider than in the de-
scription of I. pusillum in Breuss & Türk
(2004), although the other characteristics
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 659

agree with this species. Involucropyrenium
pusillum apparently looks like I. squamulosum,
but the involucrellum in I. pusillum com-
pletely encloses the exciple, whereas in I.
squamulosum it only reaches the base of the
perithecium. Moreover, the ascospores are
larger in I. squamulosum (24–27 minI.
squamulosum versus 19–24 minI. pusillum).
Other species in the genus with an entire
involucrellum are I. waltheri, clearly distin-
guishable by its hypothallus, and I. llimonae
with smaller ascospores and a thicker involu-
crellum. Involucropyrenium terrigenum and I.
sbarbaronis also develop an entire involucrel-
lum, but in both cases the ascospores are
shorter and narrower (10–12 × 7–8·5 in I.
terrigenum and 15–20 × 6–8 in I. sbarbaronis);
however, these two species have not been
found in the Iberian Peninsula.
Ecology. This species has been collected on
wall mortar, at 850 m altitude, growing with
Placidium boccanum. Breuss & Türk (2004)
reported it only from the type locality, grow-
ing on mosses above limestone at 920 m
altitude. Although the habitat in which we
collected the species agrees with that of I.
squamulosum, a saxicolous species common
on artificial substrata, the measurements
of the ascospores and the anatomy of the
involucrellum correspond with I. pusillum.
Distribution. We have collected I. pusillum
in Portugal (Fig. 4B), widening considerably
the distribution of the species previously
known only from the type locality in Austria.
Involucropyrenium tremniacense (A.
Massal.) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):38 (1996).—Catapyrenium tremniacense A.
Massal., Lotos 6: 79 (1856); type: Italien, Veneto,
Tregnano, 1855, M. Brojo (M—lectotype; BM, O, S,
W!—isotypes).
Thallus squamulose to subcrustose; squam-
ules 0·3–1·5 (2) mm, roundish to slightly
lobulate, flattened to slightly convex, epru-
inose, closely adpressed to the substratum;
upper surface beige to pale brown. Anatomy:
thallus 100–150 m thick; upper cortex para-
plectenchymatous, 10–25 m thick, with
roundish-angular cells 6–9 m diam.; epi-
necral layer 20–50 m. Algal layer 60–120 m
thick, with cells 6–10 m diam. Medulla and
lower cortex not differentiated. Rhizohyphae
colourless, c.4m thick.
Perithecia frequent, developed between the
squamules, globose, up to 350 m diam.;
exciple brown to black, with an apical involu-
crellum. Asci clavate; ascospores biseriate,
ellipsoid, 13–17 (18) × 6–7 (8) m.
Notes. This species is characterized by the
combination of incomplete involucrellum
and thallus of dispersed squamules (or sub-
crustose). Additional differences from other
species in the genus are discussed above.
Ecology.Involucropyrenium tremniacense
was found in open grassland on calcareous
substrata in semi-arid regions, from 90 –
c. 400 m altitude.
Distribution. The species has been found in
two distant localities in Spain, one located
in the north and the other in the south-east
(Fig. 4B).
The worldwide range of the species
reaches south-central Europe and the
Mediterranean region (Breuss 1990a; Nimis
& Martellos 2004).
Involucropyrenium waltheri (Kremp.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):38 (1996).—Verrucaria waltheri Kremp., Flora
38: 69 (1855); type: BRD, Bayern, Karwendel, 26 viii
1850, Krempelhuber (M—lectotype; WU—isotype).
Thallus squamulose, with a nearly crustose
appearance; squamules less than 1·5 mm
diam., brown to grey, matt, flattened to
slightly convex, contiguous, forming a crust;
margins entire to crenate-lobulate, sur-
rounded by a dark hypothallus; lower surface
dark, with 4–5 m thick, dark rhizohyphae.
Anatomy: thallus (70) 100–200 (300) m
thick; with a paraplectenchymatous upper
cortex, 10–25 m thick; epinecral layer thin or
absent, up to 25 m when present. Algal layer
distributed over nearly the entire thallus,
660 THE LICHENOLOGIST Vol. 42

with algal cells 6–11 m. Medulla not clearly
differentiated; lower cortex composed of
polygonal cells, dark; rhizohyphae dark
brown, 4–5 m thick.
Perithecia globose, often aggregated; exciple
brown to black with an entire carbonaceous
involucrellum, developed in the hypothallus
between the squamules. Asci clavate, with
biseriate ascospores (15) 17–21 (23) × (7·5)
8–10 (11) m.
Notes. This species is easily recognized by
its nearly crustose thallus, with brown to grey
squamules and perithecia developed from a
black hypothallus. It is the only species in the
genus with dark rhizohyphae. Sometimes it
can resemble Catapyrenium cinereum, from
which it differs in the perithecia that are not
immersed in the squamules and by the
presence of the involucrellum.
Ecology.Involucropyrenium waltheri was
collected on calcareous soils, together with
C. cinereum, between bryophytes, in alpine
grasslands at 2300 m.
Distribution. This species was erroneously
cited in Spain by Aragón et al. (2006), but the
revised samples (hb. Aragón 3663/97 and
2299/97) corresponded to young specimens
of Endocarpon sp. Consequently, the only
record in the Iberian Peninsula was found
during this study in the Pyrenees on alpine
grassland (Fig. 5B).
The species has an arctic-alpine distri-
bution, occurring mainly in north and central
Europe (Breuss 1990a) with some occur-
rences in Asia (Sun et al. 2008) and North
America (Breuss & McCune 1994).
Neocatapyrenium H. Harada
Nat. Hist. Res.2: 129 (1993); type: N. cladonioideum
(Vain.) Harada.
The thallus has a cushion-like growth,
composed of imbricate squamules; thallus
anatomy is similar to Placidium, but the
genus is characterized by Endocarpon-type
pycnidia with cylindrical conidia (Fig. 2F),
the lack or reduction of rhizohyphae and the
attachment to the substratum by rhizines or
by the basal end of the squamule. Peri-
thecia have colourless exciples and asci are
clavate with biseriate ascospores. There are
five species reported world-wide, three of
which are present in Europe. In the Iberian
Peninsula, we have found two species.
All species are rare and have limited
distributions (Breuss 2005).
Key to Neocatapyrenium species
1 Squamules fixed to the substratum by elongated basal ends .............
.................................N. cladonioideum
Squamules fixed to the substratum by pale rhizines .......... N. latzelii
Neocatapyrenium cladonioideum
(Vain.) H. Harada
Nat. Hist. Res.2(2):129 (1993).—Siphula cladonioides
Vain., Bot. Mag. (Tokyo)35: 47 (1921); type: Japan,
Honshu, Gunma-ken (as Prov. Kozuke), Mt. Myôgi, on
rock, 19 iv 1916, A. Yasuda 142 (TUR—holotype).
Thallus squamulose, with a cushion-like
growth; squamules lobate, 2–5 mm wide, con-
vex, imbricate, attached to the substratum by
basal ends only; upper surface flesh coloured
to beige, matt or a little glossy, epruinose;
lower surface pale brown (Fig. 1E); rhizo-
hyphae and rhizines lacking. Anatomy: thallus
(360) 407 ± 53 (480) m thick, upper cortex
(35) 43 ± 5 (50) m, paraplectenchymatous,
with cells (7) 7·6 ± 0·9 (9) m diam.; epi-
necral layer very thin or lacking, up to c.12
m thick when present. Algal layer (50)
67 ± 15 (87) m, with algal cells (7) 10 ± 1·9
(12) m diam. Medulla prosoplectenchyma-
tous, (175) 206 ± 44 (287) m thick, with
elongated hyphae of 3–5 m diam., with
some spherical cells in the uppermost and
lowermost parts of the medulla; lower cortex
clearly delimited from the medulla, (42)
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 661

55 ± 8 (62) m thick, with rounded to
angular cells of (5) 7·4 ± 1·1 (9) m, the
lowermost brownish.
Perithecia pyriform to globose, up to 400
m wide; exciple 30–40 m pale, sometimes
brownish in the upper part. Asci clavate;
ascospores biseriate, simple, (15) 15·9 ± 1·2
(18) × (6) 6·9 ± 0·4 (7·5) m, l/b ratio (2)
2·3 ± 0·3 (2·6).
Pycnidia laminal, immersed, inconspicu-
ous, Endocarpon-type; conidia bacilliform,
5–7 × c.1 m.
F. 5.Anatomy of Placidium species. A, cross-section of a marginal Dermatocarpon-type pycnidium of Placidium
semaforonense; B–D, cross-sections of the thalli; B, Placidium squamulosum showing the medulla (m) composed of
globular cells (r = rhizohyphae); C, Placidium lachneum showing the prosoplectenchymatous medulla (m) and the
lower cortex of anticlinally arranged cells (lc); D, Placidium velebiticum, showing the prosoplectenchymatous medulla
composed of elongated cells and the lower cortex clearly delimited.
662 THE LICHENOLOGIST Vol. 42

Notes. Anatomically, the specimen col-
lected in Spain is very similar to the speci-
mens described by Harada (1993), although
the squamules are thicker and wider, and
more lobate rather than sublinear as Harada
described; in addition our specimen has a
thin epinecral layer whereas the species was
described as lacking this layer. We did not
feel these differences were sufficient to justify
describing a new species, so we include this
specimen as N. cladonioideum. The species is
easily recognizable by its cushion-like growth
with imbricate squamules.
Ecology.Neocatapyrenium cladonioideum
has been collected on calcareous rocks and
soils (specimens from Asia are reported from
lava or granitic rocks). It has been found
together with Heteroplacidium imbricatum and
Placidium pilosellum, at 850 m altitude.
Distribution. The record reported here
from the Segura mountains, south-east Spain
(Fig. 4B), together with N. latzelii (see
below), constitute the first records of this
genus for the Iberian Peninsula.
Neocatapyrenium cladonioideum was pre-
viously known only from Asia, being here for
the first time reported from Europe, and
having a different ecology to specimens from
Asia.
Neocatapyrenium latzelii (Zahlbr.)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):41 (1996).—Dermatocarpon latzelii Zahlbr.,
Österr. Botan. Z.68: 70 (1919); type: Jugoslawien, Dal-
matien, Slano, Feldmauern, 3m, 4 i 1909, A. Latzel nº
799 (W!—holotype).
Thallus squamulose, with a cushion-like
growth; squamules lobate, 3–7 mm wide, im-
bricate; attached to the substratum only by
pale rhizines, rhizohyphae lacking; upper
surface pale to dark brown, matt; lower sur-
face pale brown. Anatomy: thallus 300–500
m thick, upper cortex 50–70 m thick, para-
plectenchymatous, with cells 5–8 m diam.;
epinecral layer thin, up 10–15 m thick. Algal
layer up to 100 m thick, with cells 7–12 m
diam. Medulla prosoplectenchymatous, with
elongated hyphae of 3–5 m diam., with
some spherical cells in the uppermost and
lowermost parts of the medulla; lower cortex
clearly delimited from the medulla, with
rounded to angular cells 7–10 m.
Perithecia pyriform to globose, up to 550
m wide, with a pale exciple. Asci clavate;
ascospores biseriate, simple, 13–17 × 6·5–
8·5 m.
Pycnidia laminal, immersed, inconspicu-
ous, Endocarpon-type; conidia bacilliform,
4·5–8·5 × 1–1·5 m.
Notes. The species is easily identified by its
cushion-like growth form, imbricate squam-
ules, the lack of rhizohyphae and the pres-
ence of pale rhizines. Only one specimen has
been found in the Iberian Peninsula, so we
have included measurements based on the
literature (Breuss 1990a).
Ecology. In the Iberian Peninsula the
species was collected in cavities and fissures
of calcareous rocks, in dry habitats, at
c. 450 m altitude.
Distribution.Neocatapyrenium latzelii has
been previously recorded in Croatia and
Greece (Breuss 1990a; Sipman & Raus
2002). This record constitutes the first from
Spain, and, together with N. cladonioideum,
the only representatives of the genus in the
Iberian Peninsula (Fig. 4B).
Placidium A. Massal.
Symmicta 75 (1855); type: P. michelii A. Massal.
Thallus squamulose, with squamules (1)
2–8 (15) mm wide, upper cortex 20–100 m
thick, with cells 5–17 m diam.; medulla
and lower cortex usually well developed
(Figs. 3B, 3C, 3D). Asci are usually cylindri-
cal at least when young, but clavate in some
species with rhizines. Pycnidia laminal or
marginal, Dermatocarpon-type, with oblong-
ellipsoidal to cylindrical or bacilliform co-
nidia (Fig 3A). Rhizines present or not.
Placidium species inhabit soil, rocks and
fissures, ledges and cavities, rarely trees [e.g.
P. arboreum (Schweinitz ex Michener)
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 663

Lendemer and P. corticola (Räsäsen) Breuss];
usually over calcareous substrata, although
there are some species occurring on gypsifer-
ous or acid substrata or even on artificial
substrata.
The genus comprises 32 species world-
wide, 16 of which have been reported from
Europe. In this study, we report 14 species
occurring in the Iberian Peninsula and the
Balearic Islands.
Key to Placidium species
1 Rhizines present (rhizohyphae also present), few to many per squamule .....2
Rhizines absent ...................................3
2(1) Pycnidia laminal; medulla composed of globular cells, ascospores 11–16 × 6–9 m
.................................. P. lacinulatum
Pycnidia marginal; medulla prosoplectenchymatous, ascospores 13–22 × 7–12 m
.................................P. semaforonense
3(1) Medulla composed of globular cells ........................4
Medulla prosoplectenchymatous ......................... 11
4(3) Pycnidia laminal ...................................5
Pycnidia marginal ................................. 10
5(4) Thallus thin, < 250 m; perithecial wall black, thin, <20 m thick . . P. michelii
[Lower side black; conidia oblong-ellipsoidal, 3–4 × 1·2–1·5 m;
ascospores 10–15 × 4–6 m, rhizohyphae 4–6 m]
Thallus thicker, >250 m; perithecial wall pale, >20 m thick ..........6
6(5) Conidia oblong-ellipsoid, 2·5–5 × 1·2–2 m, l/b ratio 1·25–2·9 ..........7
Conidia cylindrical to bacilliform, more than 4 m long × 1–2 m, l/b ratio 2·5– 5
...........................................9
7(6) Mainly epilithic, on mortar or calcareous rock; squamules with black margins, lower
side black throughout ....................... P. boccanum
[rhizohyphae 4–6 m; ascospores 11–15 × 5–8 m]
Mainly terricolous, on soils, rock ledges and in fissures; squamules without black
margins, lower side pale to brown, rarely blackish ................8
8(7) Squamules up to 6 mm; lower cortex cells 11–15 m, rhizohyphae 5–7 m; asco-
spores 12–17 × 6–8 m.................... P. squamulosum
Squamules up to 3·5 mm; lower cortex cells 7–12 m, rhizohyphae 3–5 m;
ascospores 11–15 × 5–7·5 m ................... P. tenellum
9(6) Thallus 240–330 m, rhizohyphae 5–6 m; conidia 4–6 × 1–1·5 m; ascospores 12–
18 × 5–7 m ............................. P. fingens
Thallus 350–550 m, rhizohyphae 6–8 m; conidia 5–8 × 1·5–2 m; ascospores
15–24 × 7·5–11·5 m...................... P. norvegicum
10(4) Thallus 300–600 m thick, rhizohyphae thick, 5–8 m; ascospores 14–22 × 6–11
m..................................P. rufescens
Thallus 210–330 m thick, rhizohyphae thin, 4–6 m; ascospores 12–17 × 5–7·5 m
................................... P. pilosellum
11(3) Pycnidia laminal, or laminal and marginal in the same squamule . P. velebiticum
[including P. pyrenaicum]
Pycnidia marginal only .............................. 12
664 THE LICHENOLOGIST Vol. 42

12(11)Lower cortex of anticlinally arranged angular cells ................ 13
Lower cortex of irregularly arranged angular cells ................15
13(12)Lower cortex without black pigment; rhizohyphae thin 5–6 m; conidia oblong-
ellipsoidal .............................P. velebiticum
Lower cortex with black pigment; rhizohyphae thick, of 6–8 m thick; conidia
bacilliform ....................................14
14(13)Squamules > 6 mm, with a thin margin; ascospores 14–17 × 6–7·5 m; conidia
4·5–6·5 × 1–1·5 m...................... P. adami-borosi
Squamules < 6 mm, with a thick margin; ascospores 12–18 × 6–8 m; conidia
5–7 × 1–2 m............................P. lachneum
15(12)Squamules large, up to 12 mm wide, almost foliose; conidia oblong-ellipsoid, 2·5–
5 × 1–2·5 m, l/b ratio 1·2–4 ..........................16
Squamules up to 6 mm wide; conidia cylindrical to bacilliform, 3–7 × 0·5–1·5 m,
l/b ratio 2–7 ...................................17
16(15)Thallus 300–600 m thick; rhizohyphae thick, 5–8 m; ascospores 14–22 × 6–11
m..................................P. rufescens
Thallus 270–360 m thick, rhizohyphae thin, 5–6 m; ascospores 13–17 × 6–7 m
...................................P. velebiticum
17(15)Thallus thin, 190–380 m thick, medulla 30–107·5 m; conidia 3–6 × 0·5–1·5 m
...................................P. imbecillum
Thallus 300–450 m thick, medulla 63–200 m; conidia 3–7 × 1–1·5 m.....
..................................P. subrufescens
Placidium adami-borosi Szatala
Ann. Mus. Nat. Hungar.7: 271 (1956); type: Hungaria,
comit. Heves, in rupibus andesit. Orient. supra vallem
Csatorna-völgy montis Remetebérc prope Mátraufüred,
c. 750 m, 5 vi 1951, Á. Boros (BP—holotype!; hb
Veˇzda— topotype).
Thallus squamulose; squamules 3–10 mm
wide, scattered to contiguous, mainly over-
lapping, with upturned margins, flat to con-
cave, roundish to slightly lobate, the margins
sometimes irregularly lobate and thin; upper
surface pale to medium brown, sometimes
with a reddish tinge, matt or a little glossy;
lower surface pale to brown. Anatomy:
thallus 400–700 m thick; upper cortex
50–80 m, paraplectenchymatous, with
cells of 8–13 m; epinecral layer up to 15 m
thick. Algal layer 100–130 m, with cells of
8–14 m diam. Medulla 150–250 m thick,
prosoplectenchymatous, with filamentous
hyphae of 3–5 m diam.; lower cortex clearly
delimited from the medulla, 50–60 m thick,
composed of angular cells, of 12–17 (20) m
wide, conglutinated in vertical rows, the low-
ermost cells with black pigment (Fig. 3C).
Rhizohyphae thick, of 6–8 m diam., colour-
less.
Perithecia broadly pyriform, up to 550 m
wide; exciple pale, 30–35 m. Asci cylindrical,
80–90 × 14–18 m, ascospores uniseriate,
simple, 14–17 × 6–7·5 m.
Pycnidia marginal, inconspicuous; conidia
bacilliform, (4·5) 5·8 ± 0·8 (7) × (1)
1·3 ± 0·2 (1·5) m) (conidia measurements
are based on the Iberian specimens).
Notes. Only two specimens have been
found in the Iberian Peninsula; thus, descrip-
tion of the species is based on previous
studies (Breuss 1990a).
This species is almost identical to P.
lachneum from which it differs mainly in the
diameter of the squamules [3–10 mm in
P. adami-borosi and 2–7 (10) mm in P.
lachneum] and thickness of the margin (thin
in P. adami-borosi and thick in P. lachneum);
the lower cortex is not so evident in the
margins in P. adami-borosi. The most import-
ant difference reported is the ecology and
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 665

distribution. Placidium adami-borosi grows at
lower altitudes in southern Europe and west-
central Asia, whilst P. lachneum is reported as
an arctic-alpine lichen distributed in Asia,
northern Europe, North and South America,
at higher altitudes (Breuss 2002b). The
minor morphological differences could be
explained by the plasticity of the species in
response to their different habitats. More-
over, data from a preliminary molecular
analysis (M. Prieto et al. unpublished) do not
support P. adami-borosi and P. lachneum as
distinct species. However, at present we have
decided to maintain P. adami-borosi as an
independent species pending a forthcoming
wider study.
Ecology.Placidium adami-borosi has been
collected from 1300 to 1650 m altitude in
cavities among mosses, or in crevices, on
calcareous substrata. In Europe the species
has been reported growing on calcareous and
siliceous substrata, from 620 to 1800 m.
Distribution.Placidium adami-borosi has
been collected in widely separated localities
in northern and southern Spain (Fig. 4C).
The species is distributed in Europe,
mainly in the Mediterranean region, and
Asia. It has been reported from Bulgaria
(Breuss 1990a), Cyprus (Breuss 1990a;
Litterski & Mayrhofer 1998), France (Breuss
1990a), Greece (Spribille et al. 2006),
Hungary (Breuss 1990a), Italy (Breuss
1990a,b; Nimis & Martellos 2004), Spain
(Breuss 1990a) and Tajikistan (Kudratov &
Mayrhofer 2002).
Placidium boccanum (Servít) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):38 (1996).—Dermatocarpon boccanum Servı´t,
Rozpr. C
ˇeskoslov. Akad. Veˇd.65 (3):23 (1955); type:
Jugoslawien, Hrvatska, Dalmatia, Hercegnovi, inter
Trebesin et Kameno, 250 m, calc., 1929, M. Servít
(PRM!—holotype).
Thallus squamulose, mainly epilithic; squa-
mules up to 4 mm wide, scattered to contigu-
ous, rarely somewhat overlapping, adpressed
to the substratum or with slightly upturned
margins, roundish to slightly lobate, flat to
fairly convex; upper surface pale to dark
brown, matt or a little glossy, with a black
margin; lower surface black throughout.
Anatomy: thallus (210) 313 ± 66 (400) m
thick, upper cortex (50) 75 ± 12 (87) m
thick, paraplectenchymatous, with cells (6)
9 ± 2·1 (13) m diam.; epinecral layer up to
35 m thick. Algal layer (75) 95 ± 14 (112)
m thick, with cells (10) 12·5 ± 1·6 (15) m
diam. Medulla (37) 109 ± 52 (162) m thick,
composed of globose cells of 9–13 m diam.;
lower cortex not clearly delimited from the
medulla, (25) 29 ± 4 (33) m thick, com-
posed of more densely aggregated rounded to
angular cells of 10-15 m, the lowermost
with dark pigment. Rhizohyphae (4)
4·9 ± 0·6 (6) m thick, colourless.
Perithecia pyriform, up to 500 m wide;
exciple pale, 30–35 m thick. Asci cylindri-
cal, (60) 66 ± 5 (73) × (10) 11·2 ± 1·2 (13)
m; ascospores uniseriate, simple, (11)
13·2±1·1 (15) × (5) 6·4 ± 0·7 (8) m, l/b
ratio (1·7) 2·1 ± 0·3 (2·8); periphyses 18–
40 × 2–3 m.
Pycnidia laminal, immersed; conidia
oblong-ellipsoid, 3–4 × c. 1·5 m.
Notes.Placidiun boccanum is characterized
by its epilithic growth, being, together with P.
rufescens, the only species of the genus occur-
ring directly on rocks. The species is ana-
tomically very similar to P. squamulosum,
which also has a medulla composed of glo-
bose cells, laminal pycnidia and oblong-
ellipsoidal conidia. However, in addition to
its epilithic habit, P. boccanum differs from
P. squamulosum in the black margins of the
squamules, and the lower surface that is con-
tinuously black, while the lower surface in P.
squamulosum is usually pale to dark brown,
rarely black. They also differ in the size of
rhizohyphae (4·9 ± 0·6 minP. boccanum
versus 5·8 ± 0·5 minP. squamulosum) and
ascospores (13·2 ± 1·1 m long in P. bocca-
num versus 14·4 ± 1·3 minP. squamulo-
sum).
The differentiation from P. tenellum is
more difficult, because both species have
relatively thin and small squamules, but they
differ mainly in the thinner rhizohyphae of P.
tenellum (4 ± 0·6 m versus 4·9 ± 0·6 m
666 THE LICHENOLOGIST Vol. 42

thick in P. boccanum). The presence of the
black margin in P. boccanum and the eco-
logy can also be useful as a distinguishing
feature.
Another similar species, with laminal
pycnidia, oblong-ellipsoidal conidia and a
medulla composed of globular cells is P.
michelii, which is clearly distinguished from
P. boccanum by its dark and thin exciple,
thinner thallus and upper cortex (313 ±
66 m thick and 75 ± 12 m thick, respect-
ively, in P. boccanum versus 200 ± 24 and
45±11m thick in P. michelii); moreover,
the ascospores are wider in P. boccanum
(6·4 ± 0·7 m versus 5·5 ± 0·5 minP.
michelii).
Placidium fingens and P. norvegicum have
larger ascospores and bacilliform conidia
(see below).
Ecology.Placidium boccanum lives on
mortar, rocks and consolidated soils, on
both acid and basic substrata. Only three
specimens have been found in the Iberian
Peninsula; one of them living on mortar of a
castle wall, the second was collected on a
rock in oak forest of Quercus ilex subsp. bal-
lota, and the third was found on acid soil
which was very consolidated due to a high
proportion of small grains of mica and
quartz.
Although in Europe P. boccanum has been
collected at low altitudes (70–250 m), in the
Iberian Peninsula the altitudinal range in-
creases from 500–1450 m.
Distribution.Placidium boccanum is a very
rare species in the Iberian Peninsula. The
record from Portugal was previously re-
ported by Carvalho (1997), whilst in Spain it
has been found for the first time in two scat-
tered localities (Fig. 4C), situated in central
and south-western Spain.
In Europe, P. boccanum has been little col-
lected in the west and south (Breuss 1990a;
Purvis et al. 1992), thus showing a
mediterranean distribution and reaching
the Macaronesian Region (Breuss 1996c). It
has been recently recorded in Turkey (Halici
& Aksoy 2009).
Placidium fingens (Breuss) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):38 (1996).—Catapyrenium fingens Breuss,
Stapfia 23: 79 (1990); type: Islas Canarias, Tenerife, Las
Cañadas, La Fortaleza, 2050–2150 m, auf bemooster
Erde, 21vii1987, O. Breuss (LI!—holotype).
Thallus squamulose; squamules up to 6 mm
wide, thin, c. 0·25–0·35 mm thick, scattered
to contiguous, fully adpressed to the substra-
tum with slightly raised margins, lobate;
upper surface pale to medium brown, matt;
lower surface pale to brownish, sometimes
with hyphal outgrows in the margins of the
squamules. Anatomy: thallus (240) 284 ± 31
(330) m thick, upper cortex (35) 52 ± 13
(77) m thick, paraplectenchymatous, with
cells (8) 11 ± 2 (15) m diam.; epinecral layer
thin, up to 25 m. Algal layer (75) 107 ± 25
(150) m thick, with cells (6) 8·9 ± 1·9
(13) m diam. Medulla (37) 72 ± 29
(137) m thick, composed of globular cells
7–13 m diam. Lower cortex not clearly de-
limited from the medulla, (17·5) 41·4 ± 16·1
(62·5) m thick, with rounded to angular
cells of (9) 11·1 ± 1·9 (15) m, the lower-
most with dark pigment. Rhizohyphae (5)
5·2 ± 0·4 (6) m thick, colourless.
Perithecia broadly pyriform, up to 500 m
wide, with a pale exciple. Asci cylindrical;
ascospores uniseriate, simple, (12) 15·4 ± 2·2
(18) × (5) 6 ± 0·5 (7) m, l/b ratio (1·9)
2·6 ± 0·5 (3·3).
Pycnidia laminal, immersed; conidia cylin-
drical to bacilliform, (4) 4·5 ± 0·6 (6) × (1)
1·2 ± 0·2 (1·5) m, l/b ratio (2·7) 3·9 ± 0·6
(5).
Notes. Anatomically, North American
populations of P. fingens are distinct from the
Iberian ones; American specimens have
smaller ascospores (11–15 × 5–6·5 m) and
longer conidia (5–7 m). In addition, the
ecology is rather different, as some of the
specimens from North America are from
bark, whereas European specimens do not
occur on trees. It is not clear whether
these two populations represent different
species as only a small amount of material
was available; a more detailed analysis is
needed.
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 667

In the Iberian Peninsula, Placidium fingens
is characterized by its bacilliform conidia
4–6 × 1–1·5 m, which are thus significantly
different in length, breadth and length/
breadth ratio from the remaining species of
Placidium with laminal pycnidia. Although
this difference is significant it is not always
easy to observe. Consequently, Iberian
populations are difficult to distinguish from
P. squamulosum and, because conidium
shape is one of the main diagnostic charac-
ters, it is necessary to have specimens with
fully developed pycnidia for correct identifi-
cation. However, the upper cortex and
medulla are significantly thicker in P. squa-
mulosum. Ascospores are narrower in P.
fingens and, consequently, the length/breadth
ratio is also different (see Table 1).
Other characters distinguish P. fingens
from the remaining species with laminal
pycnidia. Placidium boccanum can be easily
distinguished by its epilithic habit (see pre-
vious description for further details), P.
michelii has a dark exciple and thinner squa-
mules, the ascospores are significantly
shorter (12·6 ± 1·3 minP. michelii versus
15·4 ± 2·2 minP. fingens) and the length/
breadth ratio is also significantly smaller. Pla-
cidium tenellum has smaller squamules (up to
6mminP. fingens versus up to 3·5 mm in P.
tenellum), significantly thinner rhizohyphae
(5·2 ± 0·4 minP. fingens versus 4 ± 0·6 m
in P. tenellum), and significantly shorter
ascospores than P. fingens (15·4 ± 2·2 m
in P. fingens versus 13·2 ± 1·1 minP.
tenellum).
The only other species with laminal pyc-
nidia and bacilliform conidia is P. norvegi-
cum, which can be distinguished by thallus
thickness (0·35–0·55 mm in P. norvegicum
versus 0·24–0·33 mm in P. fingens), conidia
length (5–8 minP. norvegicum versus 4–6 in
P. fingens) and ascospore size (15–24 × 7·5–
11·5 minP. norvegicum versus 12–18 × 5–7
minP. fingens). The growth form of P.
norvegicum and the ecology are also import-
ant distinguishing characters (see below).
Placidium fingens has an external morpho-
logy similar to P. pilosellum, as some speci-
mens of both taxa have rhizohyphae on the
margins of the squamules, but distinguishing
between these two species is very easy when
pycnidia are present, as P. pilosellum has mar-
ginal pycnidia.
Ecology. In the Iberian Peninsula P. fingens
was found mainly in mountainous areas with
occurrences in more xeric and lower lands;
the species has been found on calcareous
soils, shelves and fissures, from 976 to
2000 m altitude. However, in North
America the species has been reported grow-
ing on soil and bark in warm, dry places.
Distribution. The species was described
from the Macaronesian region and cited by
Seaward & Arvidsson (1997) in southern
Spain, the latter constituting the first record
from Spain (and from Europe). Unfortu-
nately, a voucher for this record could not be
examined.
In the Iberian Peninsula, it is sparsely
distributed and we have not found it in
Portugal. Placidium fingens extends along
the northern part of Spain, being rarer to the
south (Fig. 4C). These records represent the
northern limit of this species in the European
continent.
Placidium fingens is also distributed in SW
North America (Breuss 2002b), occurring in
dry and warm places. One specimen from
Argentina has been found (LI 297524) dem-
onstrating its presence in South America; this
specimen is closer to the Iberian than North
American specimens.
Placidium imbecillum (Breuss) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.): 38 (1996).—Catapyrenium imbecillum Breuss,
Stapfia 23: 80 (1990); type: Österreich, Steiermark,
Totes Gebirge, am Vorderen Lahngangsee, 1500 m, 11
vii 1982, O.Breuss (hb. Breuss nº 2644—holotype).
(Fig. 1F)
Thallus squamulose; squamules up to 5 mm
wide, contiguous to densely aggregated and
sometimes slightly overlapping, thin (c. 0·2–
0·4 mm); roundish to lobate, with wavy and
raised margins. Upper surface light to me-
dium brown, matt or a little glossy; lower
surface pale, upturned margins without
668 THE LICHENOLOGIST Vol. 42

rhizohyphae (Fig. 1F). Anatomy: thallus
(190) 279 ± 44 (380) m thick, upper cortex
(28) 54 ± 14 (88) m thick, paraplectenchy-
matous, with cells of (5) 10·4 ± 2·6 (16) m
diam.; epinecral layer sometimes lacking, up
to 40 m when present. Algal layer (42)
100 ± 25 (175) m thick, with cells (4)
8·8 ± 2·1 (15) m diam. Medulla (30)
70 ± 19 (107) m thick, prosoplectenchy-
matous, with filamentous hyphae of 3–5 m
diam.; lower cortex clearly delimited from
the medulla, (20) 41·4 ± 13·8 (87·5) m
thick, with roundish to angular cells of (6)
11·4 ± 2·6 (16) m. Rhizohyphae (4)
5·1 ± 0·6 (6) m thick, colourless.
Perithecia pyriform, up to 400 m wide,
with a pale exciple, up to 30 m thick. Asci
cylindrical, 70–80 × 12–15 m (Breuss
1990a); ascospores uniseriate, simple, (9)
13·1 ± 1·7 (18) × (5) 6·1 ± 0·7 (8) m, l/b
ratio (1·5) 2·2 ± 0·3 (3).
Pycnidia marginal, relatively inconspicu-
ous, Dermatocarpon-type; conidia cylindrical
to bacilliform, (3) 4·2 ± 0·9 (6) × (0·5)
1·1 ± 0·2 (1·5) m, l/b ratio (2) 4 ± 1 (6).
Notes. The species is characterized by
its relatively thin thallus, with a medulla
that is significantly thinner (up to 100 m)
than in the other species with marginal
pycnidia, and by its bacilliform conidia.
Some specimens have hyphal outgrowths
on the margins as in P. pilosellum.Placidium
imbecillum could be confused with other
species with marginal pycnidia and a proso-
plectenchymatous medulla (e.g., P. lach-
neum,P. rufescens and P. subrufescens).
Placidium lachneum and P. rufescens differ in
their thicker thalli, medulla and lower cortex,
and in the longer and broader ascospores (see
Table 1 for differences with P. rufescens).
Placidium lachneum presents a very typi-
cal lower cortex composed of anticlinally
arranged cells. The differentiation from P.
subrufescens is more difficult, since both
species are very similar and share the same
conidial shape. Our analysis shows that
they differ mainly in thickness of thallus and
medulla; also the ascospores are longer and
wider in P. imbecillum and the conidia are
shorter (see Table 1).
The medulla of P. imbecillum is very thin,
and sometimes it is difficult to identify the
medulla type. Furthermore, in samples with-
out pycnida it could be confused with species
with medulla composed of globular cells, and
these must be differentiated on features of
thallus, rhizohyphae and ascospores.
Ecology.Placidium imbecillum is very com-
mon in the Iberian Peninsula, living on cal-
careous soils, rock fissures or ledges in
Juniperus thurifera and Quercus ilex forests
and in shrublands dominated by Buxus sem-
pervirens,Lavandula latifolia,Rosmarinus of-
ficinalis and Thymus sp., or with Herniaria
fruticosa L. and Quercus coccifera L. on gypsif-
erous soils. One specimen has been collected
in subalpine grasslands. Often it grows
together with Anthracocarpon virescens and
Placidiopsis cinerascens.
Distribution.Placidium imbecillum has pre-
viously been reported only once in the
Iberian Peninsula (Breuss 1996c). Data from
our study demonstrates that P. imbecillum is
relatively abundant, with a wide distribution,
living predominantly in central and north-
eastern parts of the Iberian Peninsula. We
have found some specimens in the south and
in the Balearic Islands and also in Portugal
(Fig. 4D), the latter constituting the first
record from Portugal.
The species is distributed in Central and
Mediterranean Europe (Breuss 1990a,b,
1994a) being recorded from Austria,
Croatia, Greece, Italy, Morocco, Slovenia,
Spain and Switzerland (Breuss 1996c; Egea
1996; Mrak et al. 2004; Spribille et al. 2006).
It is also found in North America (McCune
& Rosentreter 2007).
Placidium lachneum (Ach.) B. de Lesd.
Ann. Cryptog. Exot.5: 100 (1932).—Lichen lachneus
Ach., Lich. Svec. Prodr.: 140 (1798); type: Suecia (H-
ACH! —lectotype).
(Fig. 5C)
Thallus squamulose; squamules up to 7 mm
wide, scattered to contiguous or slightly over-
lapping, thick (c. 0·25–0·6 mm), roundish to
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 669

lobate or crenate, adpressed to the substra-
tum or with raised margins free from the
substratum. Upper surface dark brown to
red-brown, matt; lower surface black
throughout, upturned margins without
rhizohyphae. Anatomy: thallus (240)
374 ± 77 (600) m thick, upper cortex (25)
59 ± 18 (95) m thick, paraplectenchyma-
tous, with cells (6) 11·2 ± 2·7 (16) m diam.;
epinecral layer very thin or lacking, up to
c.15m when present. Algal layer (80)
113 ± 23 (162) m thick, with cells (5)
9·6 ± 2·7 (18) m diam. Medulla (50)
147 ± 49 (325) m, prosoplectenchyma-
tous, with elongated hyphae of 3–5 m
diam.; lower cortex clearly delimited from
the medulla, (30) 63 ± 25 (125) m thick,
composed of angular cells, (8) 15 ± 4·1 (28)
m × (7) 12·7 ± 4·5 (30) m, conglutinated
in vertical rows, the lowermost cells with
black pigment (Fig. 3C). Rhizohyphae thick,
(5) 6·4 ± 0·7 (8) m diam., colourless or
dark in the proximal zone.
Perithecia broadly pyriform, up to 550 m
wide, with a pale exciple c.30m. Asci cylin-
drical, 70–90 × 12–18 m (Breuss 1990a);
ascospores uniseriate, simple, (12) 15·3 ± 1·4
(18) × (6) 7·2 ± 0·7 (8) m, l/b ratio (1·5)
2·1 ± 0·3 (2·7).
Pycnidia marginal, relatively prominent;
conidia bacilliform, (4·5) 5·7 ± 0·6 (7) × (1)
1·6 ± 0·4 (2) m, l/b ratio (3) 3·8 ± 1·1 (6).
Notes. Two further varieties apart from the
type were described by Breuss (1990a). One
of them, P. lachneum var. globiferum (Breuss)
Breuss, is characterized by the presence of
pycnidia substantially larger than in the type
variety, and by auriculiform squamules. This
variety is reported from Russia (Breuss
1990a). The other variety, P. lachneum var.
oleosum (Breuss) Breuss, is characterized by
the presence of oil drops in the lower cortex
and even in the medulla, and pycnidia being
rare or sometimes completely lacking. The
distribution is similar to the type variety
(arctic-alpine), although it is less common
(Breuss 1990a). However, because the re-
ported differences are small, we have consid-
ered all specimens to belong to the same
variety. Most Iberian samples possess oil
drops in the lower cortex, and we have ob-
served this feature in other species of the
genus, including P. squamulosum and P.
pilosellum.
The species is easily distinguished by
the lower cortex anatomy, only shared by P.
velebiticum. Although P. velebiticum shares
with P. lachneum a prosoplectenchymatous
medulla with marginal pycnidia, the lower
cortex is not black pigmented and its cells are
not always in vertical rows. The main differ-
ence between them is the shape of conidia,
which are bacilliform in P. lachneum and
oblong-ellipsoid in P. velebiticum. Rhizo-
hyphae and ascospores are significantly thin-
ner in P. velebiticum (rhizohyphae 6·4 ±
0·7 m and ascospore width 7·2 ± 0·7 in P.
lachneum versus 5·3 ± 0·5 m and 6·4 ± 0·5,
respectively in P. velebiticum). Placidium lach-
neum is, together with P. rufescens,P. sema-
foronense and P. subrufescens, one of the
species with the thickest thallus of the group;
moreover, all of them have marginal pycnidia
and a prosoplectenchymatous medulla. Nev-
ertheless, P. lachneum is the only species that
has long bacilliform conidia. Differences
with P. adami-borosi have been discussed
above.
Ecology. The species has been reported
from both basic or acid substrata, although in
the Iberian Peninsula it has been found
mainly on calcareous substrata. It grows on
soil, between mosses or in fissures, in alpine
grasslands; we have also collected it in
Juniperus thurifera forests. In most cases it is
accompanied by Catapyrenium cinereum,
sharing the same ecological preferences. It
has been collected predominantly from high
altitudes, 1200 m to 2350 m.
Distribution.Placidium lachneum is distrib-
uted predominantly in northern Spain,
mainly in the Pyrenees and Picos de Europa
mountains; it is less common, in the
Mediterranean region (Fig. 4E). Records
from Granada (Casares & Llimona 1984;
Breuss 1990a) could not be confirmed.
In the Iberian Peninsula the species
has often been wrongly identified; many
670 THE LICHENOLOGIST Vol. 42

specimens have proved to be Placidium
squamulosum and P. rufescens.
Placidium lachneum has an arctic-alpine
distribution, in northern and montane re-
gions of Europe, Asia, North and South
America and in the north of Africa (Breuss
2002b).
Placidium lacinulatum (Ach.) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Endocarpon hepaticum var. lacinu-
latum Ach., Lich. Univ.: 299 (1810); type: Helvetia
(H-ACH sub Endocarpon hepaticum var. “laciniatum”—
lectotype).
Thallus squamulose; squamules up to 8 mm
wide, scattered to contiguous, rarely overlap-
ping, closely adpressed to the substratum or
with slightly raised margins, roundish to
deeply lobate, flat to concave or convex.
Upper surface pale to dark brown, matt,
epruinose or slightly pruinose; lower surface
pale to dark, rhizinate, rhizines few to many
per squamule, pale or brown, simple or
branched. Anatomy: thallus (220) 321 ± 48
(440) m thick, upper cortex (37) 67 ± 14
(92) m thick, paraplectenchymatous, cells
(6) 8·6 ± 1·5 (13) m diam.; epinecral layer
thin or lacking, up to 25 m when present.
Algal layer (55) 95 ± 21 (155) m thick, cells
(5) 9·2 ± 1·8 (15) m. Medulla (62)
146 ± 36 (227) m, composed of globular
cells (6) 9·5 ± 1·9 (13) m diam., lower cor-
tex not clearly delimited from the medulla, of
more densely aggregated cells of
(5) 8·3 ± 1·4 (11) m. Rhizohyphae (3)
4·3 ± 0·6 (5) m, colourless. Rhizines pale,
few to many per squamule, simple to
branched; up to 0·3 mm thick, and more
than 10 mm long (although it is very difficult
to remove the entire rhizine intact), but easily
broken in dry conditions and sometimes
inconspicuous.
Perithecia broadly pyriform to subglobose,
up to 600 m wide, with a pale to brown
exciple. Asci cylindrical to clavate, (55)
73·23 ± 10·9 (90) × (13) 15 ± 1·7 (18) m;
ascospores uniseriate to biseriate, simple, (11)
13·4 ± 1·2 (16) × (6) 7·4 ± 0·6 (9) m, l/b
ratio (1·4) 1. ± 0·2 (2·7).
Pycnidia laminal, immersed; conidia
oblong-ellipsoid, (3) 3·8 ± 0·5 (5) × (1·2)
1·7 ± 0·3 (2) m, l/b ratio (1·5) 2·2 ± 0·4
(3·3).
Notes. Both the algal layer and especially
the medulla are much thicker than
described by Breuss (2002b). However, the
lower cortex is often difficult to distinguish,
so that our medulla measurements appar-
ently refer to both medulla and lower cortex
together.
Breuss (1990a) distinguished two var-
ieties, var. lacinulatum and var. latisporum,
mainly on the basis of ascospore size. Ten
years later a new variety was described by
Breuss (2000), var. erythrostratum, which
has a reddish colouring of the lower part of
the medulla and wider spores than in the
type variety. Breuss (2002b) included var.
atrans (later published in Lendemer 2004),
characterized by dark brown to blackish
exciple and rhizines and subcylindrical
conidia.
The infraspecific classification of P. lacinu-
latum is still not clear. Breuss (2002b)
reported that intermediates are found and
from our point of view a complete wordwide
phylogenetic and taxonomic study based on
all species of Placidium with rhizines (e.g.
P. umbrinum, P. pseudorufescens, etc) as well
as the varieties of P. lacinulatum. (i.e. P.
lacinulatum var. atrans, var. erythrostratum,
var. lacinulatum and var. latisporum) would be
necessary in order to solve the relationships
at the species level.
In the Iberian Peninsula most specimens
corresponded to var. lacinulatum, although
ascospores were wider than in the original
description. This could be explained by the
recombination of different varieties which
are not reproductively isolated. For this
reason we consider all the specimens as the
same variety for the time being.
The species can be easily distinguished by
its rhizines, the deeply lobed squamules and
the presence of laminal pycnidia. In cases
when rhizines are inconspicuous or individ-
uals are young, the species could be confused
with P. squamulosum. However, this has
thicker epinecral and algal layers, longer and
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 671

narrower ascospores and thicker rhizohyphae
(see Table 1).
Placidium semaforonense, another rhizinate
Placidium species in the Iberian Peninsula,
has a prosoplectenchymatous medulla and
marginal pycnidia. Moreover, the ascospores
are longer and wider than in P. lacinulatum
(see below).
Ecology.Placidium lacinulatum has been
found in a wide range of habitats, mainly in
mediterranean calcareous or gypsiferous
shrublands (e.g. associated with Helianthe-
mum squamatum (L.) Dum. Cours., Reseda
suffruticosa Loefl. ex Koelp. and Bupleurum
rigidum L.); however, the species also occurs
rarely in alpine-subalpine grasslands, reach-
ing 2350 m altitude, or in the thermo-
mediterranean belt, with some records at
sea level. Placidium lacinulatum lives on
soils, fissures and ledges, showing a
preference for fine grained and bare soils.
Sometimes it was mixed with mosses and
associated with Anthracocarpon virescens,
Endocarpon loscosii Müll. Arg. and Placidio-
psis cinerascens.
Distribution. The species is very common
in Spain; it has not been found in mainland
Portugal, but we have confirmed a specimen
(in ABL) from Porto Santo in the Azores.
Until now, P.lacinulatum has been
recorded mainly from central and southern
Spain (Breuss 1990a; Llimona & Hladun
2001). Nevertheless, new data show that
the species lives predominantly in central
and north-eastern Spain, in continental
regions, with occurrences in mountainous
areas such as the Pyrenees, Picos de
Europa, Sierra de Segura, Sierra de Alcaraz
and Sierra de las Nieves (Fig. 4F).
Although we have found some specimens in
coastal areas, it is not very common and
there are reported records that we could not
revise from Cádiz, Granada (this belonging
to var. latisporum, Breuss 1990a), Murcia
and Valencia provinces.
Placidium lacinulatum is widespread, being
distributed in America, Africa, Asia and
Europe, showing clear preferences for
Mediterranean to mild temperate regions
(Breuss 2002b).
Placidium michelii A. Massal.
Sched. Crit. Lich.5: 100 (1856); type: Massalongo, Lich.
exs. Ital. 161 (W!—lectotype).
Thallus squamulose; squamules up to
5 mm, rather thin, up to c. 0·25 mm, scat-
tered to contiguous, adpressed to the sub-
stratum, roundish to slightly lobed. Upper
surface pale to dark chestnut brown, matt;
lower surface black throughout. Anatomy:
thallus thin (160) 200 ± 24 (240) m thick,
upper cortex (25) 45 ± 11 (67) m thick,
paraplectenchymatous, cells (8) 11·5 ± 2·1
(17) m diam.; epinecral layer thin or lacking,
up to 20 m when present. Algal layer (37)
69 ± 16 (105) m thick, cells (6) 9·5 ± 1·5
(12) m diam. Medulla (37) 76 ± 19 (125)
m thick composed of globular cells 6–13 m
diam.; lower cortex not clearly delimited
from the medulla, (25) 37 ± 16 (63) m, but
with more densely aggregated, rounded to
angular cells, (10) 12·4 ± 1·8 (16) m diam.,
the lowermost with dark brown pigment.
Rhizohyphae (4) 4·7 ± 0·7 (6) m thick, col-
ourless.
Perithecia pyriform, up to 350 m wide,
with a thin, 15–25 m, dark brown exciple.
Asci cylindrical, c. 60–70 × 10–13 m
(Breuss 1990a); ascospores uniseriate, simple,
(10) 12·6 ± 1·3 (15) × (4) 5·5 ± 0·5 (6) m;
l/b ratio (1·7) 2·3 ± 0·4 (3·1).
Pycnidia laminal, immersed; conidia
oblong-ellipsoid, (3) 3·3 ± 0·3 (4) × (1·2)
1·4 ± 0·1 (1·5) m, l/b ratio (2) 2·3 ± 0·3
(2·9).
Notes. The thin thallus (160–240 m)
clearly separates P. michelii from the remain-
ing species of the genus; the dark and thin
exciple is also a good diagnostic character
when perithecia are present. Some similar
species (e.g., P. fingens, P. squamulosum and
P. tenellum), living in dry habitats, could be
mistaken for P. michelii as they may have
some overmature perithecia with a darker
colour than usual.
Placidium squamulosum differs in its thicker
thallus and thallus layers (see Table 1),
672 THE LICHENOLOGIST Vol. 42

thicker rhizohyphae and conidia, and longer
and wider ascospores. Placidium tenellum also
has a relatively thin thallus, but thicker than
in P. michelii, and also differs in the thinner
rhizohyphae and broader conidia.
Differences from P. boccanum and P.
fingens are discussed above.
Ecology. Predominantly in dry, open habi-
tats in forests (Juniperus thurifera and Quercus
ilex) or shrublands (Buxus sempervirens,
Lavandula latifolia,Rosmarinus officinalis), on
calcareous soils, in rock crevices and on rock
ledges and showing a preference for sandy
soils. It has been found associated with
Anthracocarpon virescens,Placidiopsis cineras-
cens,Placidium pilosellum and P. squamulosum;
from c. 450–1100 m altitude.
Distribution.Placidium michelii was re-
corded in the Iberian Peninsula by Colmeiro
(1889) and in the early 20th century by
Sampaio (1921). Unfortunately, these speci-
mens have not been revised by us. Two fur-
ther records (LEB 4395 and 4984) from
Zamora province proved to be Placidium
squamulosum; in these specimens, the
perithecia are dark possibly because they are
overmature. However, another specimen
from the same locality is P. michelii (MACB
69463). The species was also cited in
La Rioja (Burgaz et al. 2007), but these
specimens have not been revised.
The species is sparsely distributed and
rather rare in the Iberian Peninsula. The
records are too scattered to suggest any par-
ticular distribution pattern (Fig. 6A). Never-
theless, our records constitute an increase in
the known distribution of this rare species in
the area studied and include the first record
from Portugal.
Placidium michelii is widely distributed but
scattered in Asia, Europe and North
America, and unknown in the Southern
Hemisphere (Breuss 2002b).
Placidium norvegicum (Breuss) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Catapyrenium norvegicum Breuss,
Plant Syst. Evol.159: 100 (1988); type: Norvegia,
Troms, in jugo inter Kilpisjärvi-Lyngen, 1967 J. P.
Norrlin (H!—holotype).
Thallus squamulose; squamules 3–6 mm,
upper surface medium to dark brown, matt,
rarely somewhat glossy towards the margins.
Squamules adpressed to the substratum;
contiguous but not overlapping; lower sur-
face black. Anatomy: thallus rather thick,
350–550 m, upper cortex 40–60 m, para-
plectenchymatous; epinecral layer thin. Algal
layer 100–130 m, with cells 8–14 m diam.
Medulla thick, composed of globular cells
up to 15 m. Lower cortex clearly delimited,
with roundish to polygonal cells 11–15 m,
with brown pigment between the cells.
Rhizohyphae hyaline but brownish in the
proximal zone, 6–8 m thick.
Perithecia broadly pyriform, up to 600 m
wide, with colourless to pale brown exciple,
only darkening around the ostiole. Asci cylin-
drical, 90–100 × 15–23 m, ascospores uni-
seriate, simple, (15) 17–22 (24) × (7·5)
8–10·5 (11·5) m. Periphyses 40–50
(60) × 2–3 m.
Pycnidia laminal, immersed; conidia bacil-
liform, (5) 6·1 ± 1·0 (8) × (1·5) 1·8 ± 0·2
(2) m, l/w ratio (2·5) 3·6 ± 0·7 (5·3) (co-
nidia measurements are based on the Iberian
specimen).
Notes. Only one sterile specimen was
found in the Iberian Peninsula; thus the
description of the species is based on Breuss
(1990a).
The species is characterized by its thallus
shape (forming rosettes rather than with
isolated squamules), the large and wide
ascospores and the bacilliform conidia. It
could be confused with P. rufescens, because
both species have a thick thallus; however,
P. rufescens has a prosoplectenchymatous
medulla and marginal pycnidia.
Ecology. On calcareous soil in alpine
grasslands, at 2180 m, together with Cata-
pyrenium cinereum and C. daedaleum.
Distribution.Placidium norvegicum is re-
corded from a single locality in northern
Spain (Fig. 6A).
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 673

F. 6.Distribution of Placidium species in Iberia. A, P. michelii (:) and P. norvegicum (); B, P. pilosellum;
C, P. rufescens;D,P. semaforonense;E,P. squamulosum;F,P. subrufescens.
674 THE LICHENOLOGIST Vol. 42

The species shows an arctic-alpine distri-
bution in the boreal zone of Europe and
North America, also in the Alps over 2000 m
(Breuss 1990a). This record represents the
southern limit of the species.
Placidium pilosellum (Breuss) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996). —Catapyrenium pilosellum Breuss,
Stapfia 23: 98 (1990); type: Ireland, Clare, limestone
pavement near Crusheen, crevice of limestone, grid ref.
R/3788, 17 ix 1977, A. M. Burnet (hb. Seaward nº
102259—holotype).
(Fig. 6B)
Thallus squamulose; squamules up to 6 mm
wide, scattered to contiguous and partly
overlapping, comparatively thin (0·2–
c. 0·35 mm); roundish to lobate, flat and
fully adpressed to the substratum or ± wavy
and with margins free from the substratum,
with thin hairs on the margin. Upper surface
yellowish to medium or dark brown, usually
with an orange tinge, matt; lower surface
usually pale throughout, rarely dark.
Anatomy: thallus (210) 270 ± 30 (330) m
thick, upper cortex (22) 54 ± 14 (95) m
thick, paraplectenchymatous, cells (6)
10 ± 1·9 (14) m diam.; epinecral layer thick,
up to c.65m. Algal layer (62) 90 ± 14 (125)
m thick, cells (5) 8·6 ± 1·7 (14) m diam.
Medulla (62) 105 ± 23 (170) m thick, com-
posed of globular cells (7) 10.3 ± 1.9 (16)
m; lower cortex hardly discernible from the
medulla, (18) 37 ± 18 (56) m, with more
densely aggregated cells of (7) 10·3 ± 1·9
(14) m diam., the lowermost sometimes
with a brown tinge. Rhizohyphae (4)
4·9 ± 0·5 (6) m, colourless.
Perithecia pyriform, up to 500 m wide,
with a pale exciple, up to 35 m thick. Asci
cylindrical, 70–90 × 10–15 m (Breuss
2002b); ascospores uniseriate, simple, (12)
14·2 ± 1·4 (17) × (5) 6·5 ± 0·5 (7·5) m, l/b
ratio (1·7) 2·2 ± 0·3 (3·2).
Pycnidia marginal; conidia oblong-
ellipsoid, (2) 3·2 ± 0·5 (4) × (1·2) 1·7 ± 0·2
(2) m, l/b ratio (1·3) 2 ± 0·3 (2·7).
Notes. The species is characterized by a
medulla made up of globular cells, marginal
pycnidia, oblong-ellipsoid conidia and a thin
thallus and rhizohyphae. Hyphal outgrowths
are mostly but not always present on the lobe
margin; they are not unique to this species
having also been observed in P. fingens (with
laminal pycnidia and bacilliform conidia)
and P. imbecillum (with a thinner thallus,
prosoplectenchymatous medulla, differen-
tiated lower cortex and bacilliform conidia).
When the medulla is hardly discernible
Placidium pilosellum could also be confused
with P. rufescens (with much thicker thallus
and rhizohyphae, and larger ascospores) (see
Table 1).
Placidium pilosellum and P. squamulosum
are the two most abundant species in the
Iberian Peninsula. Placidium squamulosum
has laminal pycnidia; when pycnidia are
absent the species can be distinguished on
thickness of thallus, upper cortex, algal layer
and rhizohyphae thickness (see Table 1).
We found one specimen having both mar-
ginal and laminal pycnidia in the same squa-
mule (a characteristic never reported until
now); its anatomical features and phylo-
genetic analyses (M. Prieto et al. unpub-
lished) led us to conclude that the specimen
corresponded to P. pilosellum.
Ecology.Placidium pilosellum lives in a wide
range of ecological conditions, predomi-
nantly at low altitudes, in dry open habitats
dominated by Mediterranean shrubs (e.g.,
Buxus sempervirens, Lavandula latifolia,
Rosmarinus officinalis or Thymus sp). Occa-
sionally, the species was found in forested
areas, such as Juniperus thurifera, Pinus
halepensis and Quercus ilex forests. We have
also collected it in alpine grasslands between
rock outcrops. The species lives on calcare-
ous and gypsiferous soils, in fissures and on
ledges and also on sandy soils from sea level
to 2300 m.
Distribution.Placidium pilosellum is the
most frequent species of Placidium in the
Iberian Peninsula being distributed mainly
along the eastern half of Spain, where calcar-
eous substrata dominate, although we have
also collected it in the west on calcareous
outcrops. The species is also very common in
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 675

the Balearic Islands. The new records, most
of them new at the provincial level, greatly
extend the distribution of the species in
Spain.
Until now the species was known from
Portugal in one scattered locality only in
Tras-os-Montes e Alto Douro province
(Paz-Bermúdez et al. 2009), but new data
expand its distribution to Algarve, Beira
Litoral and Estremadura provinces
(Fig. 6B).
Placidium pilosellum occurs in Australia,
Europe, the north of Africa, North America
and SW Asia (Breuss 2002b), and has been
recently recorded in South America
(Prieto et al. 2008), thus demonstrating the
cosmopolitan behaviour of the species.
Placidium rufescens (Ach.) A. Massal.
Sched. Crit. Lich.6: 114 (1956).—Endocarpon rufescens
Ach., Lich. Univ.: 304 (1810); type: Helvetia (H-
ACH—lectotype).
Thallus squamulose to almost foliose; squa-
mules large, up to 15 mm wide and up to
0·6 mm thick, densely aggregated and over-
lapping; roundish to lobate, with broad lobes
and undulate margins free from the substra-
tum, without rhizohyphae in marginal areas;
upper surface medium to dark brown, red-
brown or chestnut-brown, matt or glossy;
lower surface pale brown at the margins,
turning black toward the centre. Anatomy:
thallus (300) 410 ± 64 (600) m thick, upper
cortex (33) 61 ± 12 (88) m, paraplecten-
chymatous, with cells (5) 10 ± 2·7 (18) m
diam.; epinecral layer sometimes lacking,
up to 40 m when present. Algal layer
(63) 110 ± 27 (200) m, cells (7) 10 ± 1·4
(14) m diam. Medulla predominantly
prosoplectenchymatous, (75) 154 ± 44
(275) m thick, with elongated cells 3–5 m
diam., spherical cells usually few, but occa-
sionally numerous; lower cortex clearly de-
limited from the medulla, (38) 81 ± 19 (138)
m thick, with rounded to angular cells (8)
13·8 ± 2·7 (23) m diam. Rhizohyphae thick,
(5) 6·6 ± 0·8 (8) m, colourless.
Perithecia pyriform, up to 550 m wide,
with a pale exciple up to 30 m thick. Asci
cylindrical, 80–100 × 12–21 m (Breuss
1990a); ascospores uniseriate ascospores, sim-
ple, (14) 17·8 ± 1·8 (22) × (6) 7·9 ± 1 (11)
m, l/b ratio (1·5) 2·3 ± 0·4 (3·5).
Pycnidia marginal, relatively prominent
and abundant; conidia oblong-ellipsoid, (3)
3·7 ± 0·5 (5) × (1) 1·8 ± 0·3 (2·5) m, l/b
ratio (2) 2·1 ± 0·5 (3).
Notes.Placidium rufescens is easily recog-
nized by its large and thick squamules, large
ascospores and thick rhizohyphae; the mar-
ginal pycnidia are quite evident, with oblong-
ellipsoid conidia. The medulla is typically
prosoplectenchymatous, but in some speci-
mens the prosoplectenchymatous nature is
obscured by numerous globular cells. It is the
only species of Placidium growing directly on
rock, apart from P. boccanum.
Placidium squamulosum differs in the
dimensions of ascospores, rhizohyphae,
lower cortex cells and thallus layers
(Table 1), laminal pycnidia and medulla of
globular cells. Placidium subrufescens shares a
prosoplectenchymatous medulla and mar-
ginal pycnidia, but differs in the smaller
ascospores and bacilliform conidia (Table 1).
Placidium pilosellum and P. imbecillum also
have marginal pycnidia; for differences, see
above and Table 1. Placidium velebiticum
differs by the thinner thallus and thallus
layers, and by the thinner rhizohyphae (see
Table 1).
Ecology.Placidium rufescens occupies a wide
range of habitats, predominantly in canyons,
on rocky ground and on vertical rocks, in
Mediterranean scrublands (e.g., Buxus sem-
pervirens,Lavandula latifolia,Rosmarinus
officinalis or Thymus sp.) and forests of Abies
pinsapo Boiss., Juniperus thurifera,Pinus
halepensis,Quercus ilex or Q. pyrenaica. Also
in alpine grasslands, commonly living
directly on rocks but also on mortar, in fis-
sures and cavities and on soil. Occasionally,
we have collected it on gypsiferous soils, from
sea level to 2100 m.
Distribution.Placidium rufescens is very
common in the Iberian Peninsula, being
widely distributed in the eastern half of Spain
on calcareous soils with some occurrences on
676 THE LICHENOLOGIST Vol. 42

calcareous outcrops in the west, and in the
Balearic Islands. In Portugal it was pre-
viously cited by Paz-Bermúdez et al. (2009).
The new records extend the distributional
area of this species, being cited for the first
time in many provinces (Fig. 6C).
It is a widespread species cited in Asia,
Europe, north of Africa and North America
(Breuss 2002b).
Placidium semaforonense (Breuss)
Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Catapyrenium semaforonense
Breuss, Stapfia 23: 112 (1990); type: Kanarische Inseln,
Tenerife, Macizo de Anaga, SW-Hang des Semáforo bei
Igueste de San Andrés, 80–150 m, 10 vii 1986 (hb.
Breuss nº 4290—holotype).
(Fig. 5A)
Thallus squamulose; squamules up to 6 mm
wide, scattered to contiguous, not overlap-
ping, with the margins free from the substra-
tum and raised, sometimes up-turned and
undulate; roundish to lobate, flat to convex
or concave. Upper surface pale to dark
brown, matt, often pruinose at margins;
lower surface pale or darkening, rhizinate,
with few to many rhizines per squamule,
pale, simple or branched, sometimes anasto-
mosing. Anatomy: thallus (300) 378 ± 58
(550) m thick, upper cortex (50) 79 ± 17
(112) m thick, paraplectenchymatous, cells
(5) 8·9 ± 1·7 (12) m diam.; epinecral layer
thin or lacking, up to 20 m when present.
Algal layer (75) 124 ± 29 (200) m thick,
cells (7) 9·1 ± 1·6 (12) m diam. Medulla
(87) 175 ± 50 (375) m thick, prosoplecten-
chymatous in the upper part and with more
spherical cells towards the base, the globose
cells 7–13 m diam. and the elongated ones
3–5 m diam. Lower cortex not clearly de-
limited from the medulla, (25) 42 ± 16 (70)
m thick, of more densely aggregated cells
(6) 8·8 ± 1·4 (11) m diam. Rhizohyphae
(3) 4·2 ± 0·6 (5) m thick, colourless;
rhizines pale, few to many per squamule,
simple to branched; up to 0·35 mm thick,
and more than 10 mm long (although is very
difficult to dig up the entire rhizine), easily
breaking in dry conditions and sometimes
inconspicuous.
Perithecia broadly pyriform to globose, up
to 500 m wide, with a pale exciple, up to 40
m thick. Asci cylindrical to clavate; asco-
spores uniseriate to biseriate, simple, (13)
16·1 ± 1·8 (22) × (7) 9 ± 1 (12)m, l/b ratio
(1·4) 1·8 ± 0·2 (2·6).
Pycnidia marginal; conidia oblong-
ellipsoid, (3) 3·8 ± 0·5 (5) × (1·2) 1·8 ± 0·3
(2) m, l/b ratio (1·5) 2·1 ± 0·4 (3·3).
Notes. The species is easily recognized by
the rhizines and the lobulate squamules with
marginal pycnidia. It could be confused with
P. rufescens with a prosoplectenchymatous
medulla and marginal pycnidia, but P.
rufescens does not have rhizines (for more
differences see notes under P. rufescens).
The differences with P. lacinulatum are
discussed above.
Ecology.Placidium semaforonense grows
predominantly in xeric areas, such as dry
mediterranean scrublands with Chamaerops
humilis, Lavandula latifolia, Stipa tenacissima
L. and Tetraclinis articulata (Vahl) Mast. at
low altitudes from sea level to 1300 m alti-
tude. The species was collected mainly from
calcareous soils and shelves, with some
occurrences on gypsiferous substrata. It pre-
fers sandy soils although in the type locality it
was collected on volcanic rocks.
Distribution. Until now P. semaforonense
has been little collected in the Iberian
Peninsula, always on the east coast, and with
few records in Portugal. New data extend its
distributional area to more continental habi-
tats and to the centre of Spain, although it is
more abundant in coastal areas (Fig. 6D). It
is not very frequent in Spain, being mainly
distributed in Asia, the Macaronesian region,
Mediterranean Europe, and with some
records from Africa (Breuss 2004).
Placidium squamulosum (Ach.) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Endocarpon squamulosum Ach.,
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 677

Method. Lich.: 126 (1803); type: Germania, Sprengel
(H-ACH—lectotype).
(Fig. 5B)
Thallus squamulose; squamules up to
6 mm, scattered or contiguous to densely
aggregated or slightly overlapping, nearly
completely adnate to the substratum, some-
times with slightly raised margins; roundish
to lobate. Upper surface pale to dark brown,
matt; lower surface pale to brown, rarely
blackish. Anatomy: thallus (240) 343 ± 45
(450) m thick, upper cortex (37) 70 ± 16
(100) m thick, paraplectenchymatous, cells
(6) 10·9 ± 2·5 (17) m diam.; epinecral layer
sometimes lacking, up to 50 m when
present. Algal layer (62) 113 ± 24 (162) m
thick, cells (6) 9 ± 1·6 (14) diam. Medulla
(62) 119 ± 30 (175) m thick, composed of
globular cells of 7–15 m diam. (Fig. 3B);
lower cortex not clearly delimited from the
medulla, (25) 43 ± 12 (80) m, made up of
more densely aggregated rounded to angular
cells, (9) 11·9 ± 1·7 (15) m wide. Rhizo-
hyphae (5) 5·8 ± 0·5 (7) m thick, colourless.
Perithecia broadly pyriform, up to 650 m
wide, with a pale exciple. Asci cylindrical,
70–90 × 10–15 m (Breuss 2002b); asco-
spores uniseriate, simple, (12) 14·4 ± 1·3
(17) × (6) 6·7 ± 0·5 (8) m; l/b ratio (1·6)
2·2 ± 0·3 (2·8). Periphyses 30–40 × 3–4 m
(Breuss 2002b).
Pycnidia laminal, immersed; conidia
oblong-ellipsoidal, (2·5) 3·4 ± 0·5 (5) ×
(1·3) 1·8 ± 0·2 (2) m, l/b ratio (1·25)
2 ± 0·4 (2·9).
Notes. The species could be confused with
P. tenellum, which differs in the small squam-
ules, the thinner thallus, algal layer and me-
dulla thickness, smaller lower cortical cells,
thinner rhizohyphae and smaller ascospores
(see Table 1).
The differences between P. semaforense
and other species with laminal pycnidia are
discussed above.
Ecology.Placidium squamulosum, together
with P. pilosellum, have the widest ecological
amplitude in the Iberian Peninsula, growing
mainly in open habitats, in calcareous or
gypsiferous shrublands (with e.g., Buxus sem-
pervirens,Chamaerops humilis, Lavandula lati-
folia,Quercus coccifera,Stipa tenacissima and
Tetraclinis articulata,) and forests (with e.g.,
Juniperus thurifera,Pinus halepensis,Quercus
ilex and Q. pyrenaica). We also have collected
specimens in the coastal regions and alpine
grasslands, and occasionally in more humid
places such as Fagus sylvatica forests, grow-
ing in rock fissures. It was collected from sea
level to 2350 m altitude from calcareous and
gypsiferous soils, in rock cavities and fissures
and on ledges.
Distribution.Placidium squamulosum is
found throughout the Iberian Peninsula but
especially in the Mediterranean region and
the eastern half of the Peninsula, extending
to parts of central and northern Spain and the
Balearic Islands (Fig. 6E).
The species is thought to be the most com-
mon member of the genus worldwide, and the
most cosmopolitan species (Breuss 1993).
Placidium subrufescens (Breuss)
Breuss
Catapyrenium subrufescens Breuss, Linzer biol. Beitr.20
(2):832 (1988); type: Kanarische Inseln, Tenerife,
Montaña de la Crucita, c. 1450 m, Breuss, 12 vii 1983
(LI!—holotype).
Thallus squamulose; squamules up to
6 mm wide, contiguous to slightly overlap-
ping; roundish to lobate, with undulate mar-
gins raised from the substratum. Upper
surface light to medium brown, matt or a
little glossy; lower surface pale, margins with-
out rhizohyphae. Anatomy: thallus (300)
377 ± 41 (450) m thick, upper cortex (37)
62 ± 17 (95) m thick, paraplectenchyma-
tous, cells (5) 9·7 ± 2·8 (17) m; epinecral
layer thin, up to 25 m. Algal layer (62)
103 ± 21 (150) m thick, cells (6) 8·7 ±
1·9 (13) m diam. Medulla (63) 134 ± 26
(200) m thick, prosoplectenchymatous,
with elongated cells, 3–5 m long; lower cor-
tex clearly delimited from the medulla, (25)
47 ± 16 (87) m thick, with rounded to
angular cells of (5) 11·7 ± 3·4 (18) m.
Rhizohyphae (4) 4·6 ± 0·8 (6) m thick,
colourless.
678 THE LICHENOLOGIST Vol. 42

Perithecia pyriform, up to 500 m wide,
with a pale exciple. Asci cylindrical, 65–
80 × 10–12 m (Breuss 1990a); ascospores
uniseriate, simple, (9) 12·3 ± 1·9 (17) × (5)
5·7 ± 0·5 (7) m, l/b ratio (1·6) 2·2 ± 0·4
(3·2).
Pycnidia marginal, scattered, compara-
tively small and inconspicuous; conidia bacil-
liform, (3) 5 ± 0·9 (7) × (1) 1·1 ± 0·1 (1·5)
m, l/b ratio (3) 4·6 ± 1 (7).
Notes. The species could be confused with
P. imbecillum and P. rufescens, which share the
prosoplectenchymatous medulla and mar-
ginal pycnidia; the differences from both
species have been previously discussed (see
notes on P. imbecillum and P. rufescens for
details).
Ecology. In the Iberian Peninsula the
species lives predominantly in coastal areas,
in shrublands dominated by Chamaerops
humilis, or more interior regions in Juniperus
thurifera and Quercus ilex forests; it has
been collected on calcareous soils, and in
rock fissures and on ledges, from c. 150 m to
1300 m.
Distribution. Scattered in the Iberian
Peninsula, on the east coast , the south-east
and in the north of Spain (Fig. 6F). Recorded
here for the first time from the Iberian
Peninsula, but previously known from the
Canary Islands and Greece (Breuss 1990a;
Sipman & Raus 2002).
Placidium tenellum (Breuss) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Catapyrenium tenellum Breuss,
Stapfia 23: 126 (1990); type: Cyprus, 4 km. N Gecitköy,
on calcareous soil, 200 m, vii 1987, P. Jacobsen (LI!—
holotype).
Thallus squamulose; squamules up to
3·5 mm, scattered to contiguous, roundish
or shallowly lobate; upper surface medium to
dark brown, matt; lower surface pale to
black. Anatomy: thallus (200) 283 ± 50
(400) m thick, upper cortex (45) 66 ± 14
(100) m, paraplectenchymatous, with cells
of (8) 11·5 ± 2·2 (15) m diam.; epinecral
layer up to c.40m. Algal layer (40) 93 ± 24
(155) m thick, cells (6) 9·1 ± 1·5 (12) diam.
Medulla (47) 91 ± 24 (150) m, composed
of globular cells, 7–12 m diam. Lower cor-
tex not clearly delimited from the medulla,
(25) 35 ± 17 (62) m, made up of more
densely aggregated cells, roundish to angu-
lar, (7) 9·9 ± 2 (12) m diam. Rhizohyphae
thin, (3) 4 ± 0·6 (5) m, colourless.
Perithecia pyriform, up to 500 m wide,
with a pale exciple. Asci cylindrical, 65–
75 × 10–13 (Breuss 1990a)m; ascospores
uniseriate, simple, (11) 13·2 ± 1·1 (15) ×
(5) 5·9 ± 0·7 (7·5) m, l/b ratio (1·6)
2·3 ± 0·3 (3). Periphyses 30–35 × 3·5 m
(Breuss 1990a).
Pycnidia laminal, immersed; conidia
oblong-ellipsoid, (2·5) 3·2 ± 0·5 (4) × (1·5)
1·8 ± 0·2 (2) m, l/b ratio (1·3) 1·8 ± 0·4
(2·7).
Notes. Characterized by the small squam-
ules and thin rhizophyphae. Another diag-
nostic character is the size of the cells in the
lower cortex. These characters distinguish P.
tenellum from P. squamulosum (Table 1). The
differences between P. tenellum and other
species with laminal pycnidia have been dis-
cussed above.
Ecology.Placidium tenellum is relatively
common in coastal areas in shrublands domi-
nated by Chamaerops humilis and Tetraclinis
articulata, but rarer towards the interior; it
was collected mainly on calcareous and
sandy soils, with some occurrences in gypsif-
erous soils, usually on bare substrata. One
herbarium specimen was labelled as growing
on acid soil, but we have not found the
species on this type of substratum. It often
occurs with Endocarpon pusillum,Placidium
lacinulatum and P. semaforonense and from
sea level to 1000 m altitude, although more
frequently at low altitude.
Distribution. Relatively frequent in the
Iberian Peninsula, occurring mainly on the
Mediterranean coast, with some records
from central and north Spain (Fig. 7A). Pla-
cidium tenellum was previously cited in Spain
in an unpublished work (Navarro-Rosinés
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 679

1992) but this specimen could not be traced.
The records shown here are the first pub-
lished records from the Iberian Peninsula
and for both Portugal and Spain.
Placidium tenellum has a mainly
Mediterranean distribution with occurrences
in the Irano-Turanian Region and Southern
Africa (Breuss 1995).
Placidium velebiticum (Zahlbr. ex
Zschacke) Breuss
Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool.98
(Suppl.):39 (1996).—Dermatocarpon velebiticum
Zahlbr. ex Zschacke, In Rabenh., Kriptog.-Flora IX,
Abt. I/1: 629 (1934); type: Croatia, in monte Velebit, in
ditione “Bunjevac´ka” supra Raduc´ ad saxa umbrosa, c.
1150 m, J. Baumgartner (W!— lectotype).
Placidium pyrenaicum (Breuss & Etayo) Breuss, Ann.
Naturhist. Mus. Wien, Ser. B, Bot. Zool.98 (Suppl.):39
(1996).—Catapyrenium pyrenaicum Breuss & Etayo, Pl.
Syst. Evol.181: 257 (1992); type: España, Navarra,
Baraibar, S. Miguel de Aralar, cerca del monasterio,
Carici sylvaticae-Fagetum, 1300 m, J. Etayo, 19 iv 1991
(LI! —holotype and isotypes).
(Fig. 5D)
Thallus squamulose to almost foliose; squa-
mules large, up to 15 mm wide, relatively
thin, densely aggregated to overlapping;
more or less deeply lobate, loosely adnate,
attached to the substratum by the central
parts of the squamules with the margins free,
flat or up-turned, without rhizohyphae in the
margins. Upper surface pale brown, rarely
dark, matt; lower surface pale brown at the
margins, turning black to the centre.
Anatomy: thallus (240) 312 ± 47 (430) m
thick, upper cortex (25) 49 ± 9 (67) m
thick, paraplectenchymatous, cells (7)
10·6 ± 2·5 (16) m diam.; epinecral layer very
thin or lacking, up to 10 m when present.
Algal layer (55) 85 ± 18 (125) m thick, cells
(4) 9 ± 2·4 (13) m diam. Medulla (70)
137 ± 43 (250) m thick, prosoplectenchy-
matous, with elongated cells of 3–5 m;
lower cortex clearly delimited from the
medulla, (27) 49 ± 13 (77) m thick, with
rounded to angular cells of (6) 13 ± 3 (20)
m, irregularly arranged or in distinct vertical
rows (Fig. 3D). Rhizohyphae (4) 5·4 ± 0·6
(6) m, colourless to brownish.
Perithecia pyriform, up to 500 m wide,
with a pale exciple. Asci cylindrical, 75–
90 × 12–17 m; ascospores uniseriate, simple,
(11) 14 ± 1.3 (17) × (5) 6·3 ± 0·5 (7) m,
l/b ratio (1·7) 2·2 ± 0·2 (2·8).
Pycnidia exclusively marginal or marginal
and laminal at the same time; conidia oblong-
ellipsoid, (2·5) 3·9 ± 0·7 (5) × (1) 1·7 ±
0·4 (2) m, l/b ratio (1·2) 2·5 ± 0·9 (4·2).
Notes.Placidium pyrenaicum was reported
to differ from P. velebiticum mainly by the
occurrence of laminal pycnidia (Breuss
& Etayo 1992), but we have observed
both laminal and marginal pycnidia in the
same squamule in most specimens of
F. 7.Distribution of Placidium species in Iberia. A, P. tenellum;B,P. velebiticum.
680 THE LICHENOLOGIST Vol. 42

P. pyrenaicum. The size of conidia is not
significantly different between pycnidia in
the two different positions. Only one other
specimen has been previously reported as
having laminal and marginal pycnidia at the
same time (see P. pilosellum). This is a strik-
ing characteristic not previously reported in
the group.
Placidium pyrenaicum and P. velebiticum do
not show any clear anatomical differences, so
that the position of pycnidia is the only dis-
tinguishing character; moreover, samples
with pycnidia in both positions have been
found in the type specimen and isotypes of P.
pyrenaicum. Results from molecular analyses
showed that DNA sequences are identical in
the two species (M. Prieto et al. unpub-
lished). Based on these results it was decided
to synonymyze P. pyrenaicum with P. velebiti-
cum.
Differences between P. velebiticum and
P. rufescens have been given above (see
Table 1).
Ecology. The species has been found in
mountainous regions, growing on rocks with
soil and mixed with mosses, in fissures or
cavities and occasionally on soil. In Spain,
the species occurs mainly in shaded situa-
tions in Fagus sylvatica or Fraxinus sp.
forests, from 900 to 2100 m altitude.
Distribution. Within the Iberian Peninsula,
it is confined to northern Spain (Fig. 7B). It
has previously been cited as P. velebiticum
(Aragón et al. 2007) and P. pyrenaicum
(Breuss & Etayo 1992).
The species has a scattered European dis-
tribution, in Austria, Croatia, Spain and
Switzerland (Breuss & Etayo 1992), and has
been considered as endangered (Breuss &
Etayo 1992).
Conclusions
Catapyrenium s. lat. comprises c. 69 species in
8 genera worldwide. Forty species and 6 gen-
era are represented in Europe from which 33
species (c. 80%) in 5 genera are reported here
from the Iberian Peninsula and the Balearic
Islands.
The study of the genus Catapyrenium s. lat.
in the Iberian Peninsula demonstrates the
poor knowledge of this group in the region.
An example of this is Placidium pilosellum,a
cosmopolitan species reported only occa-
sionally from the Iberian Peninsula, but
which is actually widespread in the region,
with many new records reported here. Even
more, P. imbecillum, was previously known
from one province in the north of Spain,
but is now known to be widespread in the
eastern part of the Iberian Peninsula. In
summary, eight species are new records for
the Iberian Peninsula, three of which are
reported here for the first time for Europe.
Four species were collected for the first time
in Portugal and one in Spain.
Generally, members of the group grow on
calcareous substrata (on soils, in rock fissures
and ledges or directly on rock) predomi-
nantly in open habitats or Mediterranean
shrublands; however, some species can live
on both acid and calcareous substrata (e.g.
Heteroplacidium contumescens) or even on
gypsiferous soil (e.g. Anthracocarpon vires-
cens).
Species in this group show distinct distri-
butional patterns; from those mainly distrib-
uted in coastal areas such as Heteroplacidium
contumescens,H. imbricatum and Placidium
semaforonense to species preferentially distrib-
uted in montane regions such as Cata-
pyrenium cinereum,C. daedaleum and Placid-
ium lachneum. In the region, only one species
is corticolous (C. psoromoides). There are
species widely distributed over the eastern
part of the Iberian Peninsula, e.g. Placidium
rufescens,P. squamulosum and P. pilosellum,
the latter being one of the most abundant
species in the area. The rarer species such as
Placidium boccanum,P. michelii and P. sub-
rufescens can be sparsely distributed or known
only from a single collection, for example,
Involucropyrenium llimonae, I. nuriense, I.
pusillum, I. waltheri,Neocatapyrenium cladon-
ioideum,N. latzelii and Placidium norvegicum.
Although this study has resulted in a con-
siderable expansion of the known distri-
bution of most species, Heteroplacidium
2010 Catapyrenium s. lat. in the Iberian Peninsula—Prieto et al. 681

contumescens showed the opposite pattern,
being less collected now than previously sug-
gesting a contraction of its distribution. This
could be due to the habitat destruction suf-
fered in the main distributional area of the
species on the south-eastern coast of Spain.
For this reason this species should be
included in the Red List of endangered
species in an attempt to conserve its habitats.
We are grateful to the curators of the herbaria in ABL,
ARIZ, ASU, B, BCC, BCN, BM, BP, COLO, GB, H,
HAL, L, LEB, LI, LISU, MA, MACB, MAF, NY,
PRM, S, SANT, TFC, TUR, VAL, VIT and C. Keller
for loans of type material and other significant collec-
tions. We also thank O. Breuss for helping in several
identifications and for improving this manuscript with
his useful comments and A. Orange for the English
revision and comments on the manuscript. The first
author also thanks R. Belinchón, D. Martı´nez, I. Morell,
M.A.G. Otálora, S. Prieto and J. Torres for their help
with field work. This research was supported by a grant
from the Universidad Rey Juan Carlos to the first author,
and the research projects CGL2004-04795-C04-04 as
part of “Flora Liquenológica Ibérica” and CGL2007-
66066-C04-04 from the Spanish Ministry of Education
and Science.
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