The Lichenologist 39(1): 15–33 (2007) 2007 British Lichen Society
doi:10.1017/S0024282907005993 Printed in the United Kingdom
The lichen genus Porina in Macaronesia, with descriptions of
two new species
Emmanuël SÉRUSIAUX, Franz BERGER, Maarten BRAND and
Pieter van den BOOM
Abstract: Detailed studies on the lichen genus Porina in Macaronesia have led to a reappraisal of the
genuine identity of Porina atlantica (Erichsen) P. M. Jørg., a characteristic species that has previously
been confused with P. guaranitica, P. heterospora, P. nucula, P. mastoidea or P. rhodostoma in the
literature, and is here reported from Madeira, the Canary Islands, Ireland, France and Portugal. Two
new species are described: P. effilata Brand & Sérus. sp. nov. (known from Madeira, the Canary
Islands, Great Britain, Ireland and Portugal) and P. ocoteae Brand & Sérus. sp. nov. (restricted to La
Palma, Canary Islands, and São Jorge in the Azores). Porina leptospora Nyl. is recognized at the species
level, and P. isidiata Kalb & Hafellner is reduced into synonymy with P. atlantica. A key to all known
species of Porina in Macaronesia is provided.
Key words: Azores, Canary Islands, England, France, Gomera, Ireland, La Palma, laurisilva,
Los Tilos, Madeira, Porina effilata, Porina ocoteae, Portugal, rare species, São Jorge, Tenerife, Wales
Introduction
The checklist of lichens occurring in
Macaronesia (Hafellner 1995, 1999, 2002)
reflects the confusion of authors with regard
to the identity of several species of Porina in
the archipelagos. For example, Hafellner
(1995: 74) suspected that earlier reports of
P. mastoidea (Ach.) A. Massal. and P. nucula
Ach. referred to P. isidiata Kalb & Hafellner;
he accepted P. guaranitica Malme as a
distinct species, with P. heterospora (Fink)
R. C. Harris as a synonym. For the same
group of species, Etayo (1998: 100–101)
and McCarthy (2003: 21) accepted only one
species, viz. P. atlantica (Erichsen) P. M.
Jørg., with P. guaranitica and P. heterospora
as synonyms.
E. Sérusiaux: Plant Taxonomy & Conservation Biology
Unit, Environmental Sciences Dept., University of
Liège, Sart Tilman B22, B-4000 Liège, Belgium.
Email: e.serusiaux@ulg.ac.be
F. Berger: A-4794 Kopfing 130, Austria.
A. M. Brand: Klipperwerf 5, NL-2317 DX Leiden, The
Netherlands.
P. van den Boom: Arafura 16, NL-5691 JA Son, The
Netherlands.
As we now have large collections of Porina
from Madeira and the Canary Islands as well
as some from the Azores, we are able to
produce a more detailed taxonomic account
of the species present. The large amount
of additional relevant data now available
(Harris 1995; Lücking & Vězda 1998;
McCarthy, 1993, 2000, 2001, 2003) was
also critical for this study.
The generic concept
Almost simultaneously, Harris (1995) and
Kalb & Hafellner (1995) introduced new
generic concepts within the Porinaceae (the
valid name for the family since the typification of Porina Müll. Arg. with P. nucula
Ach.; see McCarthy 2003: 8), including
revisions of the circumscription of the wellknown genus Porina. The two taxonomic
circumscriptions are not consistent with
each other and neither has received wide
support (see McCarthy & Malcolm 1997 for
a discussion), although the genus Pseudosagedia (Müll. Arg.) M. Choisy, resurrected
by Kalb & Hafellner (1995), is now used in
several European checklists. The purpose of
16
THE LICHENOLOGIST
this paper is to examine the species present
in Macaronesia, and, to some extent, those
also present in Western Europe and is
thus species-dedicated. We have therefore
adopted a conservative generic concept
and broad delimitation of Porina, following
McCarthy (2003). We have also suggested
that the introduction of new generic concepts should take into account phylogenetic
analyses based on multiple gene DNA
sequences, when representative species of all
groups within the family have been tested
(Grube et al. 2004a; Baloch & Grube,
2006).
Methods
This study is based on collections and observations
made by the authors in Macaronesia, western Europe
and several tropical regions, as well as on the examination of herbarium material borrowed from institutions
and colleagues. The material was examined in tap
water, in lactophenol cotton-blue (LCB; FLUKA
Chemika 61335) or in Lugol’s solution (IKI; Lugol
solution SIGMA L-6146). The measurements always
refer to material mounted in water. The dimensions of
perithecia are those of the perithecia or when present
the perithecial warts (enclosing the perithecia s. str.) in
herbarium specimens.
The Species
Porina atlantica (Erichsen)
P. M. Jørg.
Graphis Scripta 12: 1 (2000).—Ocellularia atlantica
Erichsen, Hedwigia 66: 276 (1926); type: Canary
Islands, Tenerife, Llano de los Vieijos in Monte Minas
de Abajo, on Laurus canariensis, 17 iii 1919, L. Lindiger
(HBG—holotypus, not seen) [material requested but
not received].
Porina isidiata Kalb & Hafellner, Herzogia 9: 83
(1992); type: Madeira, An der Levada da Serra do Faial
entlang, etwas SW von Santo da Serra in Richtung
Camacha, in letzten Resten eines alten, jedoch stark
gestörten, NW-exponierten Eucalyptus-Waldes, an
Laurus azorica, 750 m, 13 viii 1990, K. Kalb 23484 (hb
Kalb—holotypus!, isotypus!).
Porina guaranitica auct. europ. p. p., non Malme,
Ark. Bot. 23(A): 13 (1929).
Porina heterospora auct. europ. p. p., non (Fink) R. C.
Harris, in Tucker & Harris, Bryologist 83: 12 (1980).
Porina mastoidea auct. europ. p. p., non (Ach.) Müll.
Arg., Bot. Jahrb. Syst. 6: 399 (1885).
Porina nucula auct. europ. p. p., non Ach., Syn. Meth.
Lich.: 112 (1814).
Porina rhodostoma auct. europ., non Müll. Arg., Bot.
Jahrb. Syst. 6: 398 (1885).
Vol. 39
(Figs 1A & B, 3A–D, 7A)
Thallus
epiphloeodal,
rather
thick
(c. 0·5 mm) and strongly appressed and
adherent to the bark, occasionally overgrowing liverworts and mosses, very rare on
rocks, usually not shiny, dark green to pale
greyish, in herbarium specimens usually pale
with an orange or pinkish tinge, when fresh
and alive typically orange-brown to vivid
green, rugose to verrucose, sometimes
irregularly wrinkled, typically with oxalate
crystals deposited in globose agglomerations
inside the thallus (crystallocumuli sensu
Hafellner & Kalb 1995: 163–164), which
can usually be seen as translucent dots under
high magnification, without a prothallus.
Isidia absent or locally developed on parts of
thalli, or abundant and covering large parts
of thalli, simple to coralloid, cylindrical to
most commonly globose and constricted
several times along their length and thus
appearing moniliform, usually brittle and
easily removed, concolorous with the thallus
or paler, 0·1–0·5(–1·0) mm high.
Perithecia rare to abundant, hemispherical
to subglobose, slightly but distinctly constricted at their base, rarely not, 0·5–0·9
(–1·1) mm diam., typically covered by a
thallus layer containing agglomerations of
oxalate crystals and thus appearing as if
enclosed in a large thalline wart; ostiolar
region slightly depressed, usually orange,
pinkish, reddish to brownish, rarely dark
brown, in young perithecia almost indistinct
as the covering thallus layer has not yet got
thinner so that the underlying orange involucrellum becomes visible. Involucrellum typically pale orange in section and K+ bright
orange to reddish, 30–40 m thick, readily
detached from the thallus covering layer in
sections and separated from it by a very loose
hyphal tissue around the base of mature
perithecia. Excipulum pale yellow, c. 20 m
thick. Paraphyses abundant, simple or rarely
branched near the base, not inflated at the
apices, c. 1·5 m thick. Asci cylindricalclavate, up to 20020–24 m. Ascospores
8/ascus, fusiform, never tapering towards
one end, straight or slightly curved, 7–9
(–13)-septate (one spore seen with 15
2007
Porina in Macaronesia—Sérusiaux et al.
17
F. 1. Morphology of Porina species. A & B, P. atlantica [La Palma, Barranca Gallegos, iv 2004, E. Sérusiaux s.
n. (LG)]; C & D, P. effilata, arrows point to perithecia and colours electronically exaggerated in C to make the
perithecia more conspicuous [C: Ireland, Killarney, 2000, A. M. Brand 40242 (hb Brand); D: La Palma, Cubo de
la Galga, 3 iv 1986, A. M. Brand 13302 (hb Brand)]; E & F, P. ocoteae, E, young perithecia; F, mature perithecia
[La Palma, Los Tilos, iv 2004, E. Sérusiaux s. n. (LG—holotypus)]. Scale=0·1 mm.
septa), 44–807–13 m, plus a distinct,
2–3 m thick perispore.
Pycnidia rare, usually immersed in the thallus with only the ostioles visible, 0·1–0·3 mm
diam., with a hyaline or pale orange, K+
bright orange wall. Pycnidiospores ellipsoid,
sometimes slightly tapering towards one end,
hyaline, simple, 2·5–3·01·0–1·5 m.
18
THE LICHENOLOGIST
Vol. 39
F. 2. Morphology of Porina species. A, P. guaranitica [Paraguay, 1893, G. O. A. Malme 1469 (S—holotypus)];
B, P. mastoidea [Papua New Guinea, Laing Island, 1992, P. Diederich 11644 (hb Diederich)]; C, P. nucula [Papua
New Guinea, Burbura logging site, 28 vii 1992, P. Diederich 11909b (hb Diederich)]; D, P. rhodostoma [Dominica,
Cabrit National Park, 26 iii 1998, E. Sérusiaux s. n. (LG)]. Scale=0·1 mm.
Notes. In Macaronesia, Porina atlantica is a
distinctive species, easily recognized by its
dull, verrucose thallus containing packets of
oxalate crystals, large perithecia covered by
a thallus layer with an orange to reddish
brown periostiolar region, and 7–9(–13)septate ascospores, 44–807–13 m, not
tapering towards one end and with a thick
perispore. Quite surprisingly, it has been
confused with P. effilata, newly described in
this paper, and which has a non-verrucose
thallus (lacking packets of oxalate crystals),
smaller perithecia with a smooth or slightly
tomentose surface, and ascospores typically
effilate (caudate and tapering towards the
proximal end), narrower [9–11(–13)septate, 68–837–11 m] and with a much
thinner perispore. Porina atlantica comes
close to P. nucula Ach., characterized by a
thallus that is strongly appressed and adher-
ent to the bark, verrucose, pale orange to
vivid green when fresh and without a prothallus, prominent perithecia covered by a
thallus layer with a slightly reddish ostiolar
region and broad ascospores, 7-septate and
with a thick perispore.
Porina nucula Ach. (Figs 2C, 3E & F) is
the type species of the genus, but several,
often confused taxa are clearly related. Their
taxonomy remains to be clarified, as demonstrated by the preliminary treatment of
Harris (1995: 172–175) for Florida (USA).
The lectotypification made by Harris (1995:
174) points to a species with perithecia
slightly constricted at their base and
with ascospores 40–5211–14 m, with
rounded ends. He further demonstrated a
considerable variation in ascospore size and
septation in the same group, under the
informal ‘‘sp. 674’’ [ascospores narrower,
2007
Porina in Macaronesia—Sérusiaux et al.
F. 3. Ascospores of Porina species. A–D, P. atlantica
[A, Ireland, West Cork, 20 ix 1982, P. W. James s. n.
(BM); B, La Palma, Los Tilos, iv 2004, E. Sérusiaux s.
n. (LG); C, Madeira, Chão da Ribeira, vii 2003, E.
Sérusiaux s. n. (LG); D, France, Ste Engrâce, P. van den
Boom 12821 (hb van den Boom)]; E & F, P. nucula
[E, Papua New Guinea, P. Diederich 11489 (LG, hb
Diederich); F, Papua New Guinea, P. Diederich 11909b
(LG, hb Diederich)]. Scale:=10 m.
with pointed ends, 42–53(–61)(7–)
8–11(–12) m], ‘‘sp. 6’’ [ascospores with
ends pointed, evenly tapering at both
ends, 60–7510–13(–15) m; perithecial
warts large, c. 0·6 mm in diam.], and ‘‘sp.
30308’’
(ascospores
25–335–6 m;
perithecial warts small, c. 0·3 mm in diam.).
The matter is indeed unresolved, and our
own experience with tropical material confirms that a great deal of variation is present
and that further taxa are most probably
involved. We therefore decided to adopt
the ‘‘finely cut’’ circumscription of Harris
19
(1995: 174) and Lücking & Vězda (1998:
213–214). R. Lücking kindly provided us
with data on the lectotypes designated by
R. C. Harris in H and NY. The delimitation
of the species by McCarthy (1994: 400;
2001: 140) is broader and would question
the circumscription adopted here for P.
atlantica. Until more data become available
from tropical areas, we distinguish these two
species mainly by the size of the perithecia: in P. altantica, as circumscribed here,
mature perithecia are 0·5–0·9(–1·1) mm in
diameter, while they never exceed 0·7 mm in
P. nucula.
Within the populations here assigned to P.
atlantica, septation and size of ascospores
can vary considerably: several specimens
from Macaronesia and Ireland (Fig. 3A & B)
consistently have 7-septate ascospores
measuring 44–607–11 m, while in others
the spores are more septate and longer. The
most divergent specimens are ‘Madeira,
Chão da Ribeira, vii 2003, E. Sérusiaux s. n.’
which has 9–13-septate ascospores measuring 60–7110–13 m (Fig. 3C), and
‘SW France, Ste Engrâce, P. van den
Boom 12821’ which has 9–11(–15)-septate
ascospores measuring 72–8011–13 m
(Fig. 3D). As the variation covers the whole
distributional range (as studied in this
paper) of the species and is not correlated
with any other characters, it has been
included in the general description provided
above.
Examination of the species names formerly
used for the species in the area of study. Tavares
(1952: 320) used the name ‘‘Porina mastoidea (Ach.) Mass.’’ for specimens of this
species collected in Madeira, although he
had not seen the type. However, he did
examine the type of Ocellularia atlantica
Erichsen, described from Tenerife, and he
was ‘‘inclined to think that the Tenerife
lichen is only a variant of Porina mastoidea’’.
The description provided by Erichsen
(1926: 276) reads as follows: ‘‘Thallus (. . .)
verruculoso-inaequalis, interdum aliquid
rimosus, (. . .). Apothecia solitaria in verrucis. Verrucae fertiles subglobosae, basi non
constrictae, numerosae sed non confertae,
20
THE LICHENOLOGIST
F. 4. Ascospores of Porina species. A & B, P. mastoidea [A, Papua New Guinea, P. Diederich 11491 (LG, hb
Diederich); B, Papua New Guinea, P. Diederich 11644
(LG, hb Diederich)]; C–E, P. heterospora [C, USA,
Florida, E. Sérusiaux 1765 (LG); D, USA, Louisiana,
S. C. Tucker 18481 (E); E, Guatemala, P. van den Boom
33491 (LG, hb van den Boom)]. Scale=10 m.
0·6–1 mm latae. (. . .) Sporae octonae,
decoloratae, fusiformes, 5–11 septatae,
0·062–70 mm longae et 0·018–21 mm
crassae, episporio 0·003–4 mm crasso.’’ It is
a rather accurate description of the taxon
dealt with here. Although the type collection
was not available for this study, we accept
this epithet.
Porina mastoidea (Ach.) Müll. Arg. is a
different species with a rather smooth, shiny,
thick thallus which readily peels and
becomes detached from the substratum,
and forms bullate portions, with a welldeveloped black prothallus. Its perithecia are
usually applanate and rarely constricted at
their base, typically have a conspicuous
black ostiole, and the ascospores are
7-septate, 32–666–13 m, with a 1·5–
3·5 m thick perispore (ascospore characters
following McCarthy 2001: 137; Figs 2B, 4A
Vol. 39
& B). Although the ascospores are quite
similar to those of P. nucula and P. atlantica,
P. mastoidea probably belongs to a different
taxonomic group, which corresponds to
Clathroporina Müll. Arg. sensu Harris (1995:
171–172). Porina africana Müll. Arg., P.
tetracerae (Ach.) Müll. Arg. and P. ocoteae
Brand & Sérus., newly described in this
paper, also belong to that latter group.
No material of P. atlantica was available to
Swinscow when he published his outstanding revision of the genus in the British Isles
(Swinscow 1962): indeed, both collections
he mentioned under P. nucula Ach. from
Ireland belong to P. effilata (pp. 47–49; both
collections seen in BM: Aug. 1935, D. A.
Jones s. n. and June 26, 1961, P. W. James &
T. D. V. Swinscow s. n.; see below, under P.
effilata). The drawings of ascospores (within
the ascus or standing alone) in Swinscow’s
fig. 12 are typical for P. effilata, but his
section of a perithecium is misleading as well
as part of his description of the perithecial
wall: indeed, he records that perithecia are
‘‘largely covered by a thalline layer’’ and
adds the puzzling ‘‘loculi containing plaques
of refractile material sometimes present in
thalline covering layer’’. This is never the
case with P. effilata nor in the material from
Ireland he had at hand. We suggest that
Swinscow had extracted these data on the
perithecia from other specimens he had
examined, apparently from tropical areas
and belonging to the genuine P. nucula
or other species, possibly the genuine P.
heterospora [see his sentence (p. 48): ‘‘The
description is based on the examination of
five specimens from the southern United
States of America, two from Cuba, and three
from south-west Ireland’’]. He may have
seen caudate and multi-septate ascospores
from material belonging to P. heterospora
from the US (Fig. 4C–E) and these are
indeed close to the Irish material of P. effilata
he had on his table. The drawing of the
perithecial section in Swinscow’s fig. 12 is a
further clue to the confusion of species as the
ascospores it contains are not at all caudate
and mostly 7-septate, and thus cannot
belong to either of the Irish specimens
he was studying. This mixture of data is
2007
Porina in Macaronesia—Sérusiaux et al.
suspected to be the source of the confusion
for further workers examining material of P.
atlantica and P. effilata.
Swinscow (1962: 47–49) assigned the two
collections from Ireland he was studying to
P. nucula Ach., a widespread pantropical
species (McCarthy 2003: 73–74). The differentiation of this species from P. atlantica
is quite difficult and has been discussed
above, the main conspicuous difference
being the size of mature perithecia.
The use of the epithet nucula was accepted
by Poelt & Vězda (1977: 195, 201) for
‘‘Ireland and Portugal’’ and by Champion
(1976: 30) for a collection from ‘‘Anaga’’ on
Tenerife in the Canary Islands. Both P.
atlantica and P. effilata are now recognized
in Ireland and Portugal, while only P.
atlantica is known in the Sierra de Anaga in
Tenerife.
Following the suggestion by Harris (1975:
173), the name P. heterospora (Fink) R. C.
Harris (=‘‘P. nucula auct. brit., non Ach.’’)
was adopted in the The Lichen Flora of Great
Britain and Ireland (Purvis et al. 1992: 491).
However, it is clear from the description
provided that the authors of the Flora were
working with material of both P. atlantica
and P. effilata. Indeed, meanwhile P. atlantica had been found in SW Ireland (collections made in 1982 by P. W. James and
P. M. Jørgensen) and its conspicuous and
large perithecia were noted by these
lichenologists. The description of thallus
and perithecia published in Purvis et al.
(1992) reads as follows and was clearly
inspired by Swinscow’s drawing (1962: 47):
‘‘Thallus (. . .) coarsely granular to verrucose, sometimes corrugate-wrinkled.
Perithecia 0·5–0·9 mm diam., (. . .) largely
covered by a thalline exciple containing
locules with plaque-like crystals, immersed
in thalline warts with only apex exposed’’; it
perfectly matches P. atlantica. The description of ascospores (‘‘Ascospores 40–80
8–12 m, 7–14-septate, fusiform-clavate’’)
refers to P. effilata and was also inspired by
Swinscow’s drawing.
The epithet heterospora was introduced in
the lichen flora of Macaronesia by González
et al. (1990: 106) for Tenerife and by Kalb &
21
Hafellner (1992: 82) for Madeira. The area
studied by González et al. is a part of the
Sierra de Anaga where P. atlantica occurs,
and the species is also common in all laurel
forests of Madeira.
Kalb & Hafellner (1992: 83) described P.
isidiata Kalb & Hafellner as the isidiate
counterpart of specimens they referred to P.
heterospora but are obviously identical with
P. atlantica. We suggest that P. isidiata is
merely a form of P. atlantica of no taxonomic
value. Isidiate specimens of P. atlantica have
been observed in all localities sampled
in Macaronesia, as well as in European
material. The isidia are globose to cylindrical
and sometimes form coralloid proliferations
on the thallus surface; they can be sparse
and present only on parts of the thalli and
seem to develop into coralloid outgrowths
in specimens under stress. The distinction
between abnormal outgrowths of the photobiont and genuine isidia in Porina is a difficult matter: the experience of McCarthy
(1993: 16) suggests ‘‘these structures are
manifestations of rampant growth by the
photobiont’’, an opinion shared by Harris
(1995: 171). However, the latter author
described the new Clathroporina isidiifera on
the basis for example of its ‘‘true isidia,
constricted at the base with an organization
similar to that of the thallus’’. It can indeed
be suggested that two different types of
‘‘isidia’’ can be found in the genus, one
being genuine isidia and acting as diaspores,
and the other being ‘‘adaptations to enhance
gas exchange in a permanently humid
environment’’ (Grube et al. 2004b: 1161). A
similar situation is found within populations
of another species found mainly in La Palma
(P. ocoteae; see below): many individuals do
not produce such isidioid outgrowths, but
those in apparently stressed conditions or
growing in less suitable habitats do produce
them; moreover, such thalli do not produce
perithecia. In these cases (isidioid thalli in P.
atlantica and P. ocoteae), we are convinced
that the production of ‘‘isidia’’ does not
represent a diagnostic criterion for taxonomic purposes. Otherwise, the collections
referred to P. isidiata are identical with P.
atlantica.
22
THE LICHENOLOGIST
Jørgensen (2000a) examined the type of
Ocellularia atlantica Erichsen and found it to
be identical with the populations named P.
heterospora ‘‘distributed from SW Ireland to
Macaronesia’’. The photograph provided of
the type clearly indicates that it represents
the species dealt with here as P. atlantica.
However, the only collection made by
Jørgensen in SW Ireland and with a handwritten identification label as P. atlantica
seen by us (P. M. Jørgensen no. 9121, BG!)
is typical P. effilata. He also examined
(Jørgensen 2000b) an isidiate specimen he
also had collected in SW Ireland during the
same field trip in 1982 (P. M. Jørgensen no.
9126, BG!) and accepted the epithet isidiata
for it. This specimen has a few isidia and
definitely belongs to the isidiate form of P.
atlantica.
In his revision of the saxicolous Porina
in the Southern Hemisphere, McCarthy
(1993: 49) reduced P. heterospora (Fink) R.
C. Harris into synonymy with P. guaranitica
Malme, described from rocks in Paraguay.
This opinion was adopted by van den Boom
et al. (1995: 278) for a collection from SW
France, but was questioned by Harris (1995:
173). McCarthy (2003: 21) confirmed this
view in his remarkable Catalogue of the
Porinaceae, and further accepted the older
epithet atlantica for the species. We believe
that McCarthy adopted a broad species concept for this group, which includes P. guaranitica from S America, the P. heterospora
group as studied in detail by Harris (1995:
172–175) for the south-eastern United
States, P. atlantica as circumscribed here
and the newly described P. effilata.
The type collection of P. guaranitica has
been examined (Paraguay, Paraguaría, on
basalt, 27 vi 1893, G. O. A. Malme 1469,
S—lectotype). It grows on rock, its perithecia (immersed in thalline verrucae; see
McCarthy 1993: 50, fig. 50) do not exceed
0·5 mm in diam. and have a pale orange
periostiolar region; its ascospores are
elongate-fusiform, slightly but distinctly
tapering towards one end, 10–16-septate,
74–879–10 m (plus a perispore 2 m
thick; Fig. 2A, 5A). Harris (1995: 173) also
saw the type but gave different data on spore
Vol. 39
F. 5. Ascospores of Porina species. A, P. guaranitica
[Paraguay, G. O. A. Malme 1469 (S—lectotype)]; B &
C, P. rhodostoma [B, Cuba, C. Wright, (G—holotype);
C, Dominica, 26 iii 1998, E. Sérusiaux s. n. (LG)]; D,
P. leptospora [Azores, F. Berger 15974 (hb Berger)]; E &
F, P. borreri [E, France, Ste Engrâce, 10 vii 1989, E.
Sérusiaux s. n. (LG); F, Belgium, Furfooz, P. van den
Boom 13835 (LG)]. Scale=10 m.
septation and size: .9·5 septaZ and 57–
71 m long for ten spores examined. The
specimens named P. atlantica here usually
have larger perithecia with a larger and
darker periostiolar region, and ascospores
usually with fewer septa, and never tapering
towards one end. Although we have not
studied any other specimen from South
America, we suggest that these differences
might be worth recognition at the species
rank. However, even if the populations from
South America are to be considered as conspecific with those from Macaronesia, atlantica is the older epithet and can be used for
the populations found in Macaronesia and
Western Europe.
In his study of the Trichotheliaceae from
Florida (USA), Harris (1995: 172–175)
described P. heterospora (Fink) R. C. Harris
with clavate spores having a long tapering
2007
Porina in Macaronesia—Sérusiaux et al.
tail,
(9–)11–13-septate,
85–11012–
14 m. We can confirm such a description
on the basis of specimens from SE USA and
Guatemala (see list below; Fig. 4C–E). Such
shape, septation and size are completely
different from either species dealt with here
(P. atlantica or P. effilata) and the epithet
heterospora can thus be excluded from the
flora of Europe and Macaronesia. Porina
heterospora is said (Harris 1995: 173) to be
common in the southern coastal plain of SE
USA, as well as Cuba and Venezuela.
McCarthy & Malcolm (1996: 549) introduced the epithet ‘‘aff. rubrostoma Müll.
Arg.’’ (a mispelling of rhodostoma Müll.
Arg.) for a collection from Tenerife (Las
Montañas de Anaga). The corresponding
specimen has not been examined but we
assume it belongs to P. atlantica, as two
specimens have been seen from the very
same area in Tenerife. McCarthy (2003: 84)
has also used the epithet rhodostoma for the
specimen from SW Ireland named isidiata by
Jørgensen (2000b), which we here refer to
P. atlantica (see above). The epithet now
appears in the Checklist of Great Britain and
Ireland (Coppins 2002: 47; see also http://
users.argonet.co.uk/users/jmgray/
checklist.html, visited on Dec. 29th, 2004,
no longer in use and replaced by http://
www.thebls.org.uk/checklist.html#P, visited
on July 31th, 2006). Besides the type collection and these other two specimens, P.
rhodostoma has never been cited elsewhere in
the literature (McCarthy 2003: 84).
The type collection of P. rhodostoma was
also examined (Cuba, C. Wright ‘‘Ser. II,
539, G—holotype). It grows on bark, has a
rugulose to verrucose thallus, large and
rather flattened perithecia immersed in
thallus verrucae (up to 0·8 mm diam.), with
a conspicuous, large, typically dark red
(dark red to red-brown) periostiolar region.
The original description states that the
ascospores are ‘‘fusiformes, 65–80 m
longae et 10–12 m latae, 7–11-septatae,
loculi 2 intermedii reliquis longiores’’. Two
perithecia have been sectioned and the
ascospores found outside the asci are rather
badly preserved; they are largely fusiform,
with rounded ends, not tapering towards
23
one end, 5–9-septate, 41–6413–17 m
(plus a 3 m thick perispore; Fig. 5B). We
have seen no collection from Macaronesia
that could match such characteristics.
A healthy, recently collected specimen
from Dominica, West Indies (Fig. 2D; see
below), is referred to the same species;
its thallus and perithecia are identical, its
ascospores have the same shape but are
11–13-septate and measure 74–8921–
26 m (plus a 2–3 m thick perispore;
Fig. 5C). If further collections can confirm
these observations, it would mean that the
ascospores studied in the type are not yet
fully mature. We suggest that these two
collections represent a different taxon from
P. atlantica, differing mainly by the typically
dark red and large periostiolar region and by
the size and septation of the ascospores.
Ecology and distribution. Porina atlantica
occurs in all stands of the laurisilva studied
in Madeira (where it can be common) and
the Canary Islands; so far, no specimens
have been collected in the Azores. It usually
grows on bark in rather shaded places and
has been recorded on basalt boulders in
La Palma, inside well-preserved laurisilva
where the species is rather common on trees.
It is also known from several localities
along the Atlantic coast of western Europe
where it seems to be very rare: SW Ireland
in two localities, France (Brittany and the
western Pyrenees), and Portugal (Sintra). In
Brittany, the thallus of the only collection
available is typical (isidioid outgrowths are
also present), but no mature perithecia
could be seen. It must also be noted that all
European localities are well-known for their
well-developed lichen flora comprising
several typical atlantic species.
Specimens examined. France: Dépt. Finistère: 26 km
W of Huelgoat, forêt de Cranou, near St-Conval, on old
Quercus, 175 m, 1996, A. M. Brand 34439 (hb Brand).
Dépt. Pyrénées Atlantiques: Ste Engrâce, near entrance of
Gorges of Kakouetta, shaded wood, on Buxus, 510 m,
1992, P. van den Boom 12821 (hb van den Boom).—
Ireland: V. C. H2, North Kerry: Cromaglown bridge,
on Quercus, 1982, P. M. Jørgensen 9126 (BG—L/48004,
mentioned as P. isidiata by Jørgensen 2000b); Killarney,
Meeting of the Waters, N of old Weir Bridge, on old
sheltered Quercus at shore, 2000, A. M. Brand 40241
24
THE LICHENOLOGIST
(hb Brand). V. C. H3, West Cork: Glengarriff woods,
shaded side of old Quercus, 20 ix 1982, P. W. James
s. n. (BM).—Portugal: W of Lisboa, Sintra, near
Capuchos monastery, on old Cupressus, 250 m, 2003,
A. M. Brand 49770 (hb Brand).—Canary Islands:
Gomera: Parque Nacional de Garajonay, El Cedro,
chemin depuis le village jusqu’à l’Ermita, le long du
barranco del Cedro, laurisylve de fond de vallée, avec
Persea indica, 900–950 m, 26 vii 1994, E. Sérusiaux s. n.
(LG, 2 collections). La Palma: 8·5 km N of Santa Cruz,
Bco la Galga, Cubo de la Galga, laurisilva forest in cleft,
550 m, 1999, P. van den Boom 22430 (hb van den
Boom); ibid., on basalt boulder inside the forest, 580 m,
1986, A. M. Brand 13313 (hb Brand); Los Tilos (W de
Las Lomados), laurisilve riche en Hedera canariensis
et en fougères (dont Woodwardia radicans), on bark,
800–850 m, vii 1997, E. Sérusiaux s. n. (LG); ibid.,
600–700 m, iv 2004, E. Sérusiaux s. n. (LG); ibid., near
picnic place, on old Octoea, 450 m, 1986, A. M. Brand
13558 (hb Brand); Barranco Gallegos, croisement
de la route Barlovento-Roque Faro par la laguna de
Barlovento, rochers de basalte en sous-bois, 900 m, iv
2004, E. Sérusiaux s. n. (LG). Tenerife: Laurisilva, head
of San Andres Valley near El Bailadero, Sierra de
Anaga, 1964, H. A. & F. H. Imshaug 35802 (LG);
Las Mercedes, on trees, 600–700 m, xii 1984, C.
Hdez Padrón (E).—Madeira: Ribeiro Frio, Hänge SW
oberhalb der Fischzuchtanstalt, in Lorbeerwald mit
einzelnen Kastanienbäumen, aus Laurus azorica,
950 m, 1990, K. & A. Kalb 23710 (hb Kalb); Ribeiro
Frio, le long de la Levada de Portela, laurisylve
dégradée, 850 m, ii 1988, E. Sérusiaux s. n. (LG); S de
Seixal, Chão da Ribeira, début du chemin montant vers
Fanal, laurisylve peu perturbée, 500 m, vii 2003, E.
Sérusiaux s. n. (LG); Rabaçal, entre les maisons de
Rabaçal et la Cascata do Risco, fourrés d’Erica en
bordure de la levada, 1050 m, vii 2003, E. Sérusiaux
s. n. (LG); Fanal (le long de la route Ribeira de Janela
vers Paul da Serra), vieux Ocotea foetens dans un pâturage, 1100 m, vii 2003, E. Sérusiaux s. n. (LG); Route
Ribeira Brava/São Vicente, un peu au N de Boca de
Encumeada, fourrés du Fayal-Brezal avec quelques
fûts, 900 m, vii 2003, E. Sérusiaux s. n. (LG); Casa das
Queimadas, promenade vers Caldeirão Verde, fourrés
d’Erica en bordure de la levada et arbres plantés, 900 m,
vii 2003, E. Sérusiaux s. n. (LG).
Selected specimens examined for other species mentioned.
Porina heterospora (Fink) R. C. Harris: USA: Florida:
along road no. 361 between Adams Beach and the
highway ALT27, Spring Warrio creek, on Cupressus
by a swamp, 5 m, 1976, E. Sérusiaux 1765 (LG).
Louisiana: East Feliciana Parish, Idlewild Experimental
Farm, c. 3 miles SE of Clinton, 30(48#N 90(45#W, on
Quercus in downed hardwoods, 1979, S. C. Tucker
18481 (E).—Guatemala: Dept. Alta Verapaz: NW of
Coban, ‘‘Las Victorias’’ forest in the national park,
15(28.4#N 90(22.8#W, on trunk of small tree, 1320 m,
2004, P. van den Boom 33491 (hb van den Boom, LG).
Porina nucula Ach.: Papua New Guinea: Madang
prov.: near Bogia, along road Bogia-Josephstaal, 4(27#S
144(56#E, on trunk of felled trees among gardens,
Vol. 39
330 m, 1992, P. Diederich 11489 (LG, hb Diederich);
Burbura logging site, c. 30 km NNW of Madang,
4(50#S 145(38#E, on tree in virgin rainforest on low
hills, 70 m, 1992, P. Diederich 11901 & 11909b (LG, hb
Diederich).
Porina mastoidea (Ach.) Müll. Arg.: Papua New
Guinea: Madang prov.: near Bogia, along road Bogia–
Josephstaal, 4(27#S 144(56#E, on trunk of felled trees
among gardens, 330 m, 1992, P. Diederich 11491 (LG,
hb Diederich); near Bogia, Laing Island in Hansa Bay,
4(10#S 144(52#E, corticolous in coastal forest on
coral island, 1 m, 1992, P. Diederich 11644 (LG, hb
Diederich).
Porina rhodostoma Müll. Arg.: West Indies:
Dominica: Cabrit National Park, disturbed dry forest,
on tree, c. 100 m, 26 iii 1998, E. Sérusiaux s. n. (LG).
Porina effilata Brand & Sérus. sp. nov.
A speciebus generis in Europa et Macaronesia differt a
sporis caudatibus, (7–)9–11(–13)-septatis, (65–)68–
83(–85)7–11 m.
Typus: Canary Islands, Gomera, Parque Nacional de
Garajonay, El Cedro, chemin depuis le village jusqu’à
l’Ermita, le long du barranco del Cedro, laurisilve de
fond de vallée, avec Persea indica, sur tronc, 900–950 m,
26 July 1994, E. Sérusiaux s. n. (LG—holotypus).
Porina atlantica auct. europ. p. p., non (Erichsen)
P. M. Jørg., Graphis Scripta 12: 1 (2000).
Porina guaranitica auct. europ. p. p., non Malme,
Ark. Bot. 23(A): 13 (1929).
Porina heterospora auct. europ. p. p., non (Fink) R. C.
Harris, in Tucker & Harris, Bryologist 83: 12 (1980).
Porina nucula auct. europ. p. p., non Ach., Syn. Meth.
Lich.: 112 (1814).
(Figs 1C & D, 6A–C, 7B)
Thallus mainly epiphloeodal, able to overgrow corticolous liverworts and mosses,
crustose and rather thin, green to dark green
or pale beige and pinkish, sometimes with
a tinge of blue, rather matt, rarely shiny, not
containing large oxalate crystals, without a
prothallus. Isidia very rare (seen only on
parts of thalli in a few collections from
Ireland), simple, almost globose, c. 0·1 mm
high, not glossy and usually almost powdery,
easily detached from the thallus.
Perithecia rare or abundant, at first
immersed in the thallus and occasionally
remaining so for a long time (and therefore
hardly visible), or protuberant and sessile,
not included in thallus-dominated verrucae,
sometimes remaining hidden in cracks of the
bark or under bryophytes, subglobose, 0·3–
0·45(–0·5) mm diam.; perithecia initially
2007
Porina in Macaronesia—Sérusiaux et al.
25
fusiform, slightly but distinctly inflated in
the upper half and tapering towards the
proximal end, straight or usually slightly
curved, (7–)9–11(–13)-septate, (65–)68–
83(–85)7–11 m, with a thin but distinct
perispore.
Pycnidia very rare, immersed in the
thallus with only the ostioles visible, 0·1–
0·2 mm diam., with a pale orange, K+
bright orange wall. Pycnidiospores cylindrical,
straight or slightly curved, with rounded
ends, hyaline, simple, 15–17(–18)c. 1 m.
F. 6. Ascospores of Porina species. A–C, P. effilata
[A, Gomera, El Cedro, 26 vii 1994, E. Sérusiaux s. n.
(LG—holotypus); B, La Palma, Bco la Galga, A. M.
Brand 13302 (hb Brand); C, Ireland, West Cork,
Glengarriff, 11 viii 1966, P. W. James s. n. (BM)]; D &
E, P. ocoteae [D, La Palma, Los Tilos, iv 2004, E.
Sérusiaux s. n. (LG—holotypus); E, La Palma, Los
Tilos, P. van den Boom 22174 (hb van den Boom)]; F,
P. africana [Guatemala, P. van den Boom 33423 (hb van
den Boom, LG)]; G, P. tetracerae [Rwanda, 30 iii 2005,
E. Sérusiaux s. n. (LG)]. Scale=10 m.
covered by a thin, greenish thallus layer that
can remain for a long time around the base,
pale orange to orange, sometimes pinkish,
rarely almost translucent, K+ bright orange
to deep red, sometimes distinctly and finely
greyish-whitish tomentose in the upper half,
rarely slightly verruculose, with a central
ostiole appearing as a pale or translucent dot
or as a hole. Involucrellum well-developed,
especially in upper parts, separated from the
excipulum in old and overmature perithecia,
especially near the base, c. 20–25 m thick.
Excipulum pale orange, c. 15–20 m thick.
Paraphyses usually abundant, simple or
rarely branched near the base, not inflated at
the apices, 1–1·5 m thick. Asci cylindricalclavate, up to 18020 m when fully
mature. Ascospores 8/ascus, narrowly
Notes. This new species is easily distinguished by its pale orange, sometimes
pinkish, and rather small perithecia, not
included in thallus warts (albeit sometimes
remaining immersed in the thallus and thus
hardly visible, or sometimes covered by a
thin thallus layer in young stages that can
remain around their base), sometimes with a
distinct tomentum in their upper half, and
by its long, distinctly tapering, mainly 9–11septate ascospores. Its thallus never contains
agglomerations of large oxalate crystals. It
does not belong to the same group as P.
atlantica with which it has been confused; we
suggest it comes close or even belongs to
Segestria Fr. sensu Harris (1995: 175–176) or
to the Porina rufula group sensu Lücking
(2004: 410, 416–417).
No close relative has been found in the
literature. Porina sylvatica McCarthy &
Kantvilas, known from SE Australia
(McCarthy & Kantvilas 1993: 144–145),
has larger (0·43–0·72 mm diam.), smooth
and orange-brown to reddish brown perithecia and its ascospores are narrower (6·5–
9·5 m), not tapering towards their proximal
ends. Porina speciosa McCarthy & Malcolm,
from New Zealand (McCarthy & Malcolm
1996), has larger (0·56–1·01 mm diam.),
smooth perithecia and much larger
ascospores (65–12215–22 m) with a
thick perispore.
As explained above under P. atlantica,
Porina effilata has been mentioned in the
European literature under many different
epithets: P. nucula (Swinscow 1962: 47–49),
P. heterospora (Purvis et al. 1992: 491), P.
guaranitica (McCarthy 1993: 49–54), and P.
26
THE LICHENOLOGIST
atlantica (Jørgensen 2000a). None of these
epithets match this species. The most obvious difference between all these species and
P. effilata is the presence of large oxalate
crystals (crystallocumuli) in their thalli and
in the thallus layer covering the perithecia;
indeed, all these species belong to Porina s.
str. sensu Harris (1995: 170, 172–175) or to
the Porina nucula group sensu Lücking
(2004: 410–412). However, the perithecia of
P. effilata can reach full maturity (i.e. producing mature ascospores) while still deeply
immersed in the thallus, and thus with only
their upper part visible; even when sessile
and subglobose, their bases can still be
covered by a thin thallus layer; these two
features can be confusing. Moreover, the
long and tapering ascospores of both P.
guaranitica and P. heterospora are quite similar to those of P. effilata and we suspect that
Swinscow (1962: 47–49) had been confused
by such a strong convergence.
In the field, Porina effilata can be confused
with Belonia lumbrispora Etayo (Etayo 1996)
with which it often grows in the Canary
Islands. The perithecia of the latter are
however slightly smaller, more yellowish or
orange and are not pilose. Examination
under the microscope immediately confirms
that Belonia has a hamathecium filled with
yellow oily droplets and ascospores that
are multiseptate (40–55 septa), filiform,
flexuose, very narrow, and measuring
125–1603–4 m.
The specimens from Macaronesia differ
somewhat from those from the British Isles,
mainly Ireland. All major characters that are
diagnostic for the species are shared by both
populations, but there are slight but significant differences in the size of perithecia and
ascospores. This matter has not been
studied in detail but can be demonstrated by
the following comparison: A. M. Brand
13302 (hb Brand) from La Palma (Canary
Islands) has perithecia always distinctly
pilose, 0·4–0·5 mm diam., and ascospores
(60–)62–70·8–81(–85) m long (n=22)
(Fig. 6B), while the collection by P. W.
James, coll. 11 viii 1966 (BM) from
Glengarriff, West Cork, Ireland, has smooth
or rarely pilose perithecia, 0·5–0·6 mm
Vol. 39
diam., and ascospores (76–)80–87·6–94
(–104) m (n=16) (Fig. 6C). Ascospores
have been measured in standard conditions
(mounted in water; only those expelled from
asci under gentle pressure measured). We
would not be surprised that future studies,
based on more detailed analysis of the variation (including molecular techniques),
might indicate two different taxa are
involved.
Ecology and distribution. Porina effilata has
been found in Madeira and in the Canary
Islands (Tenerife, Gomera and La Palma),
always in rather preserved stands of the
laurisilva where it can grow on bark as well
as on twigs. It is rare but, as it is an inconspicuous species, it may have been overlooked. It is also known from SW Ireland,
Wales, and N Devon in England, where it is
also quite rare and grows on old, mossy bark
of Quercus, usually at the base of the trees, in
highly preserved and humid localities. Both
collections from Wales and one from Ireland
are from bryophytes growing over rocks in
sheltered conditions, and thus represent the
only localities where P. effilata is not corticolous; one of them is the type locality of
the recently described Biatora britannica
Printzen, Lumbsch & Orange (Printzen et al.
2001). An additional locality is known in
Portugal in the famous locality at the
monastery of Sintra where the species is
found in much more artificial conditions
(bole of Cupressus in parkland).
Specimens examined. Great Britain: England: V.C. 4,
North Devon: Bideford, Clovelly, woodland below
(NW of) Gallantry Bower, on mossy base of large
Quercus, 45 m, 1994, B. J. Coppins 16465 & A. M.
O’Dare (E). Wales: V.C. 46, Cardiganshire: near
Cardigan, Coedmor National Nature Reserve, SE of
Coedmore house, on bryophytes on vertical rock face
sheltered by ivy, 1996, A. Orange 10998 (NMW). V.C.
48, Merioneth: Talsarnau, Bryn Bwbach, Ceunant
Coch, on slightly calcareous rock face in woodland,
2002, A. Orange 13840 (NMW).—Ireland: V.C. H2,
North Kerry: Killarney, Cromaglown, on trees, viii
1935, D. A. Jones s. n. (BM); Eagle’s Nest, shady ledge
on shale cliff, on bryophytes, 26 vi 1961, T. D. V.
Swinscow s. n. (BM, 2 specimens); near Killarney,
Eagle’s Nest Mountain, on Quercus in wood, 14 viii
1966, P. W. James s. n. (BM); Torc Mountain, oak in
shade on west-facing slope, 31 vii 1965, T. D. V.
Swinscow s. n. (BM); Killarney, Muckross, on Quercus,
2007
Porina in Macaronesia—Sérusiaux et al.
14 viii 1966, P. W. James s. n. (BM, E); Killarney Lake,
Dinish Island, Camillan wood, on oak, 17 ix 1982, P.
W. James s. n. (BM), P. M. Jørgensen 9121 (BG—L/
48007) and s. n. (E); Killarney, Meeting of the Waters,
N of old Weir Bridge, on old sheltered Quercus at shore,
2000, A. M. Brand 40242 (hb Brand). V. C. H3, West
Cork: Glengarriff, road to Barley Lake, on shaded bole
of Quercus, 11 viii 1966, P. W. James s. n. (BM).—
Portugal: W of Lisboa, Sintra, near Capuchos
monastery, on old Cupressus, 250 m, 2003, A. M. Brand
49782 (hb Brand).—Canary Islands: La Palma:
4·5 km WSW of Los Sauces, Los Tilos, laurisilva,
narrow cleft with path along N facing rock sheer, on
Laurus azorica and Ocotea foetens, 750–800 m, 1999, P.
van den Boom 22240 & 22273 (hb van den Boom); ibid.,
narrow cleft with path over Bco del Aqua, mixed trees
near bridge, on tree, 800 m, 1999, P. van den Boom
22285 (hb van den Boom); ibid., gorges profondes avec
laurisilve, 600–700 m, sur arbre, iv 2004, E. Sérusiaux s.
n. (LG, 5 collections); 8.5 km N of Santa Cruz, Bco la
Galga, Cubo de la Galga, laurisilva forest in cleft,
550 m, 1999, P. van den Boom 22457 (hb van den
Boom); ibid., on bark of young trees of Lauraceae in
wood, 580 m, 1986, A. M. Brand 13302 (hb Brand);
ibid., 500–600 m, on twigs, iv 2004, E. Sérusiaux s. n.
(LG). Tenerife: Laurisilve de Monte del Agua, chemin
au départ de Erjos, vers Las Portelas, laurisilve
en mosaïque avec des fourrés de Erica, sur troncs,
900 m, 27 ii et 2 iii 1997, E. Sérusiaux s. n. (LG, 3
collections).—Madeira: Casa das Queimadas, chemin
vers Caldeirâo Verde, laurisylvedégradée, 850–
900 m, v 1992, E. Sérusiaux s. n. (LG); S de Seixal,
Chão da Ribeira, le long du Riba da Seixal, plus haut
que l’élevage de truites, laurisilve en bord de torrent, sur
tronc, 500 m, vii 2003, E. Sérusiaux s. n. (LG, 2
collections).
Porina fortunata P. M. McCarthy &
Etayo
Lichenologist 34: 199 (2002); type: Canary Islands,
Gomera, La Meseta de Vallehermoso, cliff of the
‘‘Cueva Encantada’’, on shaded basalt in laurisilva,
720 m, 22 vii 2000, J. Etayo 17823 & A. Fernández
(TFC—holotypus, not seen; CANB, hb Etayo—isotypi,
not seen).
This saxicolous species has been recently
described from shaded basalt rocks in the
laurisilva of Gomera in the Canary Islands
(McCarthy & Etayo 2002), and is here
reported from similar habitats in La Palma
(collection checked by P. M. McCarthy).
Porina fortunata has hemispherical to subglobose, small [(0·22–)0·28(–0·36) mm
diam.], dark reddish brown to black
perithecia and ascospores 3(–7)-septate,
narrowly oblong to almost cylindrical with
rather rounded
3–4 m.
27
ends,
18–25–332·5–
Specimen examined. Canary Islands: La Palma:
8·5 km N of Santa Cruz, Barranco La Galga, laurisilva
forest in cleft with volcanic rockface, on shaded basalt
boulder, 500 m, 1999, P. van den Boom 22447 (LG, hb
van den Boom).
Porina leptospora (Nyl.) A. L. Sm.
Monogr. Brit. Lich. 2: 338 (1911).—Verrucaria
leptospora Nyl., Flora 47: 487 (1864).
Porina olivacea (Pers.) A. L. Sm. var. leptospora (Nyl.)
Keissler, Rabenhorst Kryptogamen Flora, Band 9, Abt. 1,
Teil 2: 318 (1938).
Porina borreri (Trevis.) D. Hawksw. & P. James var.
leptospora (Nyl.) D. Hawksw., Lichenologist 24: 367
(1992). Type: Ireland: Kerry, Killarney, Dinish, on Ilex,
I. Caroll (?BM—holotypus; not seen).
(Fig. 5D)
This taxon is either considered to be synonymous with Porina borreri (Trevis.) D.
Hawksw. & P. James (Santesson et al. 2004:
271, sub Pseudosagedia borreri), or a variety
[var. leptospora (Nyl.) D. Hawksw.; Purvis
et al. 1992: 490]. In corticolous material of
Porina with black perithecia and 7-septate
ascospores in Macaronesia, we encountered
only populations with narrow, almost cylindrical ascospores, rarely tapering towards
one end that measure 37–502·5–3·5
(–4·0) m. In material identified as P. borreri
in continental Europe, ascospores are fusiform or clavate-fusiform and measure 22–
33(–35)3–5 m (Fig. 5E & F). We did
not find any specimens with intermediate
ascospores and thus believe the species rank
is more appropriate for these taxa.
Porina leptospora occurs in the laurisilva
in Macaronesia (La Palma and Tenerife,
Canary Islands and São Miguel in the
Azores) and, to our knowledge, is known
with certainty only from SW Ireland and
Devon in Western Europe. Although we
have not seen the material, we assume that
the reports of P. borreri from Madeira by
Kalb & Hafellner (1992: 81–82) and Los
Tilos in La Palma by Etayo (1996: 156, sub
Pseudosagedia borreri) belong to P. leptospora.
Selected specimens examined. Canary Islands: La
Palma: Los Tilos, W de Las Lomados, laurisilve
28
THE LICHENOLOGIST
Vol. 39
dominée par Ocotea foetens, riche en Hedera canariensis,
sur tronc, 600–700 m, vii 1997, E. Sérusiaux s. n. (LG);
ibid., sur un vieux Ocotea, iv 2004, E. Sérusiaux s. n.
(LG). Tenerife: laurisilve de Monte del Agua, chemin au
départ de Erjos vers Las Portelas, sur tronc d’Erica, 27
ii 1997, E. Sérusiaux s. n. (LG).—Azores: São Miguel:
Praia, Caminho do Praia to Lagoa do Fogo, shady
hollow, on Myrica faya, 120 m, 2003, F. Berger 17801
(hb Berger); Lagoa Furnas, Uferweg an NW Ecke, auf
Alnus cordifolia & Quercus sp., 280 m, 2001 & 2003, F.
Berger 15974, 17813, 17814 (hb Berger).
Selected specimens examined for Porina borreri:
Belgium: prov. Namur: 1·2 km S of Furfooz, Parc
national de Haute Recène, S-exposed slope, on big
Acer, 160 m, 1993, P. van den Boom 13835 (LG).—
France: dépt. Pyrénées-Atlantiques: Ste Engrâce, gorges
de Kakouetta, fourrés de Buxus très humides à l’entrée
des gorges, sur Buxus, 10 vii 1989, E. Sérusiaux s. n.
[with P. W. James, F. Rose & J. Vivant] (LG).
Porina ocoteae Brand & Sérus. sp.
nov.
A speciebus generis in Europa et Macaronesia differt
a sporis elongatis-fusiformibus, 7–11(–13)-septate,
(42–)44–50(–53)(3·5–)4–5 m.
Typus: Canary Islands, La Palma, W de Los Sauces,
‘‘Los Tilos’’, gorges profondes avec laurisilve et
présence de vieux fûts d’Ocotea, base de tronc d’Ocotea,
alt. 600–700 m, April 2004, E. Sérusiaux s. n. (LG—
holotypus).
(Figs 1E & F, 6D & E, 7C)
Thallus epiphloeodal, able to overgrow
liverworts and mosses growing on the bark,
very rarely (one specimen seen) on basaltic
rock at the base of a tree on which the
species was abundant, crustose, shiny green
to grey-brown, or sometimes pale brownish
or olive-grey, usually maculate with black
irregular patches, adherent to the bark but
frequently forming bullate portions, bulges
or ridges, and thus appearing to peel off
from the bark, locally with oxalate crystals
deposited in globose or irregular masses
(crystallocumuli sensu Hafellner & Kalb
1995: 163–164); prothallus almost always
present, to 0·5 mm wide, black or bluish
black, sometimes covered by a thin, white
and conspicuous hyphal layer that can give
the prothallus a fibrous or cottony appearance. Isidia locally present, sometimes
common, especially on thalli without perithecia and on apparently stressed specimens,
F. 7. Cross-sections through perithecia of Porina
species. A, P. atlantica [Madeira, Chão da Ribeira, vii
2003, E. Sérusiaux s. n. (LG)]; B, Porina effilata,
perithecium still covered with a thallus layer on its right
side and with its left side emerged from the thallus and
slightly pilose on its upper part [Gomera, El Cedro, 26
vii 1994, E. Sérusiaux s. n. (LG—holotypus)]; C, P.
ocoteae [La Palma, Los Tilos, iv 2004, E. Sérusiaux s. n.
(LG—holotypus)]. Scale=100 m.
simple or coralloid, shiny or with an eroded
surface, sometimes globose and constricted
several times along their length, brittle and
easily removed, concolorous with the thallus
or paler, 0·1–1·0 mm high.
Perithecia abundant or absent, especially
absent on thalli with well-developed isidia,
hemispherical to subglobose, very rarely
constricted at the base, 0·5–0·6(–0·7) mm
diam., typically covered by a thallus layer
containing masses of oxalate crystals and
thus appearing as included in a large thallus
wart; ostiolar region slightly depressed or
not, concolorous with the thallus or pale
orange to reddish, sometimes brownish.
Involucrellum typically pale orange and K+
2007
Porina in Macaronesia—Sérusiaux et al.
bright orange to reddish, best developed in
the upper half of the perithecium, c. 25 m
thick. Excipulum pale yellow, c. 10 m thick.
Paraphyses abundant, simple or rarely
branched, especially near the base, not
inflated at apices, c. 1·5 m thick. Asci
cylindrical-clavate, 150–17010–15 m.
Ascospores 8/ascus, elongate-fusiform, sometimes slightly tapering towards one end,
straight or slightly curved, 7–11(–13)septate, (42–)44–50(–53)(3·5–)4–5 m,
without a perispore.
Pycnidia rare, immersed in the thallus,
c. 0·2 mm in diam., with a hyaline or
pale orange, K+ bright orange wall.
Pycnidiospores ellipsoid, hyaline, simple,
2·5–3·51·0–1·5 m.
Notes. In Macaronesia, this species is
easily distinguished from all other corticolous Porina with large perithecia by its
shiny green to grey-brown thallus, usually
maculate with black irregular patches, peeling off from the bark and frequently forming
bullate portions, bulges or ridges, a typical
black or bluish black prothallus, and narrow
ascospores (not exceeding 5 m wide).
Porina ocoteae belongs to the P. mastoidea
group (referred to the genus Clathroporina by
Harris 1995: 171). Closely related species
include the pantropical P. africana Müll.
Arg. and P. tetracerae (Ach.) Müll. Arg.
(Fig. 6F & G). The former has perithecia
with a typically black periostiolar region
and less septate ascospores [(5–)7(–9):
McCarthy 2001: 113] and the latter also has
perithecia with a dark periostiolar region and
regularly 7-septate ascospores 24–44 m
long (McCarthy 2001: 150). The material
from Florida (USA) identified by Harris
(1995: 171) as P. tetracerae has longer
ascospores (35–50 m). In any case, P.
ocoteae consistently has narrower ascospores
and most have more than 7 septa, making its
distinction from P. africana and P. tetraceae
easy. Dr P. M. McCarthy has studied several
of our collections and confirmed (in litt., 4.
2003) that they represent an undescribed
species.
29
Ecology and distribution. The new species is
named after its principal phorophyte, the
laurel tree Ocotea foetens (Ait.) Benth. &
Hook. f. (‘‘el til’’ in Spanish). On the island
of La Palma (Canary Islands) in the two
famous sites of Los Tilos and Cubo de La
Galga, Ocotea foetens forms a few spectacular
groves of old trees, becoming multistemmed by self-coppicing and with large
and complex bases, which develop wood
rolls expanding downwards and on which
this new species of Porina develops conspicuous populations. Porina ocoteae has
never been observed on other tree species,
even at Los Tilos; it is however able to grow
on basaltic stones embedded in the bases of
Ocotea trees on which it is present. Fully
developed and fertile collections are known
only from Los Tilos; mostly sterile specimens are known from the other localities in
La Palma and on São Jorge in the Azores.
Accompanying species in Los Tilos include
Porina aenea, P. leptospora and the very rare
Strigula brevis (Roux & Sérusiaux 2004:
53–55). Easily detected because of its shiny
green thallus with a conspicuous black prothallus, Porina ocoteae must be considered
as a very rare species as it has not been
observed elsewhere in the Canary Islands,
nor in Madeira. Its status in the Azores must
be further studied.
Specimens examined. Canary Islands: La Palma: Los
Tilos (W de Las Lomados), laurisilve dominée par
Ocotea foetens, riche en Hedera canariensis, on bark,
600–700 m, vii 1997, E. Sérusiaux s. n. (LG, 2 collections); ibid., 800–850 m, vii 1997, E. Sérusiaux s. n.
(LG); ibid. narrow cleft with path along N facing rock
sheer, on Ocotea, 700 m, 1999, P. van den Boom 22174
(hb van den Boom); 3·5 km WSW of Los Sauces, N
slope of Barranco del Agua, at the base of Ocotea in
laurel wood, 530 m, 1986, A. M. Brand 13599 (hb
Brand); 8·5 km N of Santa Cruz, Bco La Galga,
laurisilva forest in cleft, on Ocotea, 500 m, 1999, P. van
den Boom 22424 (hb van den Boom); La Galga, gorges
au lieu-dit ‘‘Cubo de La Galga’’, 500–600 m, laurisilve
sur flancs pentus, iv 2004, E. Sérusiaux s. n. (LG).—
Azores: São Jorge: Weg Ribiera da Cedro – Faja do
Alem, Hortensienhecke, auf abgestorbener Hydrangia,
450 m, 2001, F. Berger 15771 (hb Berger).
Selected specimens examined for other species mentioned.
Porina africana Müll. Arg.: Guatemala: Dept. Baja
Verapaz, SSE of Coban, SSE of Purulhá, Biotope
Mario Dary Rivera, 15(13.2#N 90(13.9#W, NE
exposed slope with tropical rain forest, on trunk of small
30
THE LICHENOLOGIST
tree, 1850 m, 2004, P. van den Boom 33423 (hb van den
Boom, LG).
Porina tetracerae (Ach.) Müll. Arg.: Rwanda: prov.
Cyangugu: Cyamudongo forest, 02(33#50·6$S,
28(58#94·9$E, lower montane forest with very tall trees
Vol. 39
of Newtonia buchananii, Entandrophragma excelsum, etc.,
on small trunks and lianas, 2000 m, 30 iii 2005, E.
Sérusiaux s. n. [Field trip to Rwanda with Damien Ertz,
Eberhard Fischer & Dorothee Killmann, 2005] (LG).
Key to the species of Porina in Macaronesia
1
On living leaves; ascospores 3-septate . . . . . . . . . . . . . . . . . . . . . . . 2
On bark; ascospores with 3 or more septa . . . . . . . . . . . . . . . . . . . . 4
On rocks; ascospores with 3 or more septa . . . . . . . . . . . . . . . . . . . . 9
2(1)
Perithecia black, usually lens-shaped, rarely hemispherical
. P. oxneri R. Sant.
[very rare: reported only once from Gomera, Canary Islands by Etayo (1998: 102), sub Pseudosagedia
obsoleta (Oksner) Hafellner & Kalb; not checked by us]
Perithecia never ‘‘pure’’ black, always with a reddish tinge, always constricted at the
base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3(2)
Perithecia 0·2–0·3 mm diam., subglobose or almost so, brownish red, at least when
young partly covered by hyphal tissue; algal cells typically present in a layer
between the inner and outer walls of the perithecium . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . P. hoehneliana (Jaap) R. Sant.
[very rare: known only from two localities in Madeira, i.e. the laurisilva of Riba da Seixal at
300–400 m, where it is abundant, and Chão de Louros where it was found in 1951 by C. Tavares but
not seen there since]
Perithecia 0·1–0·2 mm diam., globose or vertically elongated, dark red to dark
reddish brown; no hyphal tissue covering the perithecia and no algal cells present
between the perithecial walls . . . . . . . . . . . P. leptosperma Müll. Arg.
[known only from Madeira where it is abundant in the laurisilva, even when heavily disturbed]
4(1)
Thallus bluish grey or grey, rather pinkish when fresh, with abundant, simple to
coralloid isidia which usually have hyaline, apical ‘hairs’; perithecia rare or
absent; ascospores ellipsoid, 9–11-septate, c. 40–659–15 m . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. coralloidea P. James
[=Zamenhofia coralloidea (P. James) Clauzade & Cl. Roux]
[reported from Tenerife and Gomera in the Canary Islands, and the Azores archipelago; mostly
found on old trunks of arborescent Erica at the edge of laurisilva; never abundant]
Thallus never bluish grey (in P. ocoteae, a bluish grey prothallus is present, and in
P. effilata, the thallus can sometimes have a bluish tinge) and never with isidia
terminated with hyaline ‘hairs’ . . . . . . . . . . . . . . . . . . . . . . . . . 5
5(4)
Perithecia black or almost so . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Perithecia never black or almost so, ostiolar region sometimes dark brown or rarely
blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6(5)
Ascospores 3-septate, ellipsoid and rarely exceeding 20 m in length . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aenea (Wallr.) Zahlbr.
[on trees; probably rare in Macaronesia: only reported from Tenerife by Gil González et al. (1990:
106) and La Palma by Etayo (1996: 156); a single collection from La Palma, Los Tilos seen by us.
The reports of P. chlorotica on trees in Tenerife by Champion (1976: 30) and Gomera by Etayo
(1998: 102) may refer to this species]
Ascospores 7-septate, almost cylindrical and always exceeding 20 m in length .
. . . . . . . . . . . . . . . . . . . . . . . . . P. leptospora (Nyl.) A. L. Sm.
[on trees; specimens seen by us from La Palma and Tenerife in the Canary Islands and São Miguel
in the Azores; reports of P. borreri from Madeira (Kalb & Hafellner 1992: 81–82) and La Palma
(Etayo 1996: 156) not checked but assumed to represent this species]
2007
7(5)
Porina in Macaronesia—Sérusiaux et al.
31
Perithecia subglobose, 0·3–0·45(–0·5) mm diam., pale orange to orange, sometimes pinkish, sometimes distinctly and finely greyish whitish tomentose in the
upper half, rarely slightly verruculose, but never containing agglomerations of
oxalate crystals; ascospores narrowly fusiform, slightly but distinctly inflated in
the upper half and tapering towards the proximal end, straight or most usually
slightly curved, (7–)9–11(–13)-septate, (65–)68–83(–85)7–11 m, with a thin
but distinct perispore . . . . . . . . . . . . . . . . P. effilata Brand & Sérus.
[on bark and twigs in the laurisilva of Madeira and Gomera, La Palma and Tenerife in the Canary
Islands; rare but possibly overlooked]
8(7)
Perithecia hemispherical to subglobose, containing agglomerations of oxalate
crystals in their outer walls, always larger when mature . . . . . . . . . . . . 8
Thallus rugose to verrucose, sometimes wrinkled, sometimes with isidioid outgrowths, without a prothallus; perithecia 0·5–0·9(–1·1) mm diam.; ascospores
fusiform, 7–9(–13)-septate, 44–807–13 m, plus a distinct, 2–3 m thick
perispore . . . . . . . . . . . . . . . . . . P. atlantica (Erichsen) P. M. Jørg.
[on bark in the laurisilva of Madeira (locally very abundant) and Gomera, La Palma and Tenerife in
the Canary Islands]
Thallus usually maculate with black irregular patches, adherent to the bark but
peeling off and forming bullate portions, bulges or ridges, locally isidioid
outgrowths present, sometimes common, especially on thalli without perithecia,
with a black or bluish black prothallus; perithecia 0·5–0·6(–0·7) mm diam.;
ascospores elongate-fusiform, 7–11(–13)-septate, (42–)44–50(–53)(3·5–)4–
5 m, without a perispore . . . . . . . . . . . . . P. ocoteae Brand & Sérus.
[known only at the bases of old boles of Ocotea foetens, in localities with well-preserved laurisilva in
La Palma, Canary Islands—the most important being the most famous laurisilva of ‘‘Los Tilos’’, and
from São Jorge in the Azores—Endemic]
9(1)
10(9)
Perithecia either yellow-orange or covered by a thallus layer but with the ostiole
usually pale orange, pinkish to brownish; ascospores at least 7-septate . . . 10
Perithecia black or dark reddish brown; ascospores 3 or 7-septate . . . . . . 11
Perithecia 0·3–0·4 mm diam., yellow-orange to brownish red, not covered by a
thallus layer containing oxalate crystals; thallus thin and smooth; ascospores
7-septate, 42–5212–15 m, plus a c. 2 m thick perispore . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . P. ahlesiana (Körb.) Zahlbr.
[known from shaded basaltic boulders in laurisilva; reported from Gomera, Canary Islands by
McCarthy & Etayo (2002) and Graciosa in the Azores by Aptroot & Rodrigues (2005), from La
Palma and São Jorge in the Azores (LG, hb Berger); specimens from Gomera and Graciosa not seen
by us]
Perithecia 0·5–0·9(–1·1) mm diam., usually distinctly constricted at their base,
typically covered by a thallus layer containing agglomerations of oxalate crystals
and thus concolorous with the thallus; ostiole usually pale orange, pinkish to
brownish; thallus rather thick and rugulose-verruculose; ascospores fusiform,
7–9(–13)-septate, 44–807–13 m, plus a distinct, 2–3 m thick perispore .
. . . . . . . . . . . . . . . . . . . . . . . P. atlantica (Erichsen) P. M. Jørg.
[normally a corticolous species but found once on basaltic boulders inside laurisilva at La Palma,
Canary Islands]
11(9)
Perithecia hemispherical to subglobose, small [(0·22–0·28(–0·36) mm diam.], dark
reddish brown to black; ascospores 3(–7)-septate, narrowly oblong to almost
cylindrical with rather rounded ends, 18–332·5–4 m . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . P. fortunata P. M. McCarthy & Etayo
[known from shaded basaltic boulders in laurisilva; only two localities, in Gomera and La Palma,
Canary Islands—Endemic]
Perithecia usually hemispherical and rather immersed in the thallus, rarely
subglobose, never dark reddish brown, always black . . . . . . . . . . . . 12
32
THE LICHENOLOGIST
Vol. 39
12(11) Perithecia 0·2–0·3 mm diam., very rarely aggregated; ascospores 3-septate, 16–
25(–32)4–6 m . . . . . . . . . . . . . . P. chlorotica (Ach.) Müll. Arg.
[reported from the Azores (Degelius 1941: 8), Madeira (Tavares 1952: 319), Gomera (McCarthy &
Etayo 2002), and from La Palma and Tenerife, at: www.gobcan.es/medioambiente/biodiversidad/
ceplam/bancodatos/libro.html]
Perithecia 0·4–0·6 mm diam., sometimes aggregated into clusters of 2–4; ascospores 7-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13(12) Ascospores fusiform to oblong-cylindrical with rounded ends, (26–)30–455–
6 m; thick (up to 5 m) perispore present but not easily seen, usually associated
with immature ascospores only . . . . . . . . . P. guentheri (Flot.) Zahlbr.
[known from shaded basaltic boulders, usually in laurisilva, in Gomera and La Palma, Canary
Islands; only specimens from La Palma seen by us—report from La Gomera at: www.gobcan.es/
medioambiente/biodiversidad/ceplam/bancodatos/libro.html; recently reported from Graciosa in the
Azores by Aptroot & Rodrigues (2005)]
Ascospores narrow-fusiform to cylindrical, with rounded ends but usually slightly
tapering towards one, 42–553–4 m; perispore not seen . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. curnowii A. L. Sm.
[reported here from coastal localities, at low elevations in São Jorge and Flores, Azores; coll. in hb
Berger]
Excluded species
Porina glabra (A. Massal.) Zahlbr.: this
species is included in the checklist of
lichens and lichenicolous fungi in the
Canary Islands, available at the address
below. The name refers to Strigula glabra
(A. Massal.) V. Wirth, a species for
which no material has been seen from
Macaronesia (Roux & Sérusiaux 2004).
www.gobcan.es/medioambiente/biodiversidad/
ceplam/bancodatos/libro.html
Porina rosei Sérus.: the doubtful report of a
sterile collection of that species (Sérusiaux
1991) has been assumed to be actually
one of Porina isidiata Kalb & Hafellner
(Kalb & Hafellner 1992). We here confirm that the specimen (LG!) is close to
but not unequivocally conspecific with P.
rosei and is not an isidiate form of Porina
atlantica.
P. semecarpi Vain.: the only collection of that
species mentioned from Madeira belongs
to P. hoehneliana (Sérusiaux 1996: 225);
the world distribution map of the species
published by McCarthy (2003: 89) should
thus be corrected.
We thank the curators of the following herbaria for the
loan of material in their care: BG, BM, E, G, NMW, S
and hb Kalb. Dr P. Diederich kindly made his material
of Porina from Papua New Guinea available to us and
Dr Robert Lücking kindly provided relevant data on the
type collection of Porina nucula Ach. in H and NY. Dr
P. M. McCarthy examined several specimens of ours
from Macaronesia and made available his valuable
notes and observations: we owe him a great debt; he
further reviewed carefully the ms as a referee and
provided additional useful comments. We also thank
Alan Orange, the second referee, for his interesting
comments and suggestions.
R
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Accepted for publication 25 August 2006