Verrucaria mundula P.M. McCarthy (Verrucariaceae,
Ascomycota) – a new record for the Northern Hemisphere
Carlos Pérez Valcárcel1, M. Eugenia López de Silanes2,3, and
Graciela Paz-Bermúdez2
1
I.E.S. Adormideras, Juan Sebastián Elcano 28 A Coruña, España; 2 E.U.E.T.
Forestal, Campus A Xunqueira, Universidade de Vigo, 36005 Pontevedra, Spain
ABSTRACT. Verrucaria mundula P.M. McCarthy var. mundula is recorded for the first time for the
Northern Hemisphere. Differences between Iberian specimens and the Australian type material are
indicated, and the species is compared to other freshwater Verrucaria species found in Europe.
KEYWORDS. Freshwater lichens, saxicolous, Iberian Peninsula.
¤
In the continental waters of Europe, lichens that
withstand immersion have, in general, little studied.
Significant treatments include those by Zschacke
(1927, 1933–34), Wirth (1972), Keller (2000), Thüs
(2002), and Thüs and Schultz (2009) for Central
Europe, by Swinscow (1968), Hawksworth (1989),
Gilbert (1996), Gilbert and Giavarini (1997), and
Orange (2004) for the British Isles and by Asta et al.
(1973), Ménard (1997), and Coste (2005) for France.
The Iberian species were studied by Pereira (1992),
Pereira and Llimona (1986–87, 1992) who focused
on NE part of the Peninsula, or by Valcárcel et al.
(1999) and Valcárcel & Carballal (2002) that dealt
with the taxa occurring in the NW.
The genus Verrucaria is frequent in freshwater
habitats; 11 aquatic species belonging to this genus
sensu lato have so far been collected in the Iberian
Peninsula, namely Verrucaria aethiobola, V. aquatilis,
V. elaeomelaena, V. funckii, V. hydrela, V. latebrosa,
V. margacea, V. pachyderma, V. praetermissa, V.
rheitrophila (Hydropunctaria rheitrophila by Gueidan
3
Corresponding autor e-mail: esilanes@uvigo.es
DOI: 10.1639/0007-2745-113.2.267
The Bryologist 113(2), pp. 267–271
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et al. 2009) and V. submersella. All of them are
widespread throughout Europe.
Most of the authors cited above regarded
ascospore size as an important character to
distinguish these species. To date, species in Europe
with the smallest ascospores (less than 15 mm)
include, in freshwater habitats, V. aquatilis, V.
madida, V. rheitrophila and V. scabra. During work
for the ‘‘Flora liquenológica Ibérica’’ project, the first
author sampled many freshwater sites in all parts of
the Iberian Peninsula. An additional species with
small ascospores was collected in seven Spanish
provinces; it is reported here as V. mundula
McCarthy, the first record of this species from the
Northern Hemisphere.
MATERIAL AND METHODS
Specimens in SANT (recently collected material of
Verrucaria mundula, and comparative material of
other species) and MEL were studied.
Microscopical measurements were made on
hand-cut sections, which were mounted in water or
KOH. An optical microscope was used to study
samples by bright field, dark field or phase contrast.
Dark field was considered a suitable method to
0007-2745/10/$0.65/0
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113(2): 2010
Figure 1. Verrucaria mundula: A. vertical section of perithecium; B. asci and subcylindrical ascospores. C. V. rheitrophila, vertical
section of thallus with carbonaceous spots. D. V. aquatilis vertical section of perithecium. Scale bars: A, B, D 5 10 mm and C 5
100 mm.
photograph and observe involucrellum growth, while
phase contrast proved useful for observation of asci,
ascospores, excipulum and periphyses.
THE SPECIES
Verrucaria mundula P.M. McCarthy Fig. 1A&B
TYPE: AUSTRALIA. NEW SOUTH WALES: Chichester State
Forest, 19 km W of Stratford, Karuah R., by a
ford on the Karuah Valley Road, off Wards
River-Berrico Trig. Road, on moderately
exposed aquatic slate, 9 Feb 1991, P.M.
McCarthy 517 (MEL 1055304 - holotype!).
Description of specimens from Spain. Thallus,
crust-like, epilithic, subgelatinous, light to dark
brown, sometimes almost black, smooth, 24.5–60
(–80) mm thick, ecorticate. Most of the specimens
with abundant colorless crystals on the thallus of
substratum. Photobiont green, 5–7 mm diam.;
cyanobacteria often present in addition. Perithecia
black, 0.14–0.18 mm in diameter, solitary, scattered,
hemispheric, varying from half-immersed in the
thallus to more or less superficial, covered by a thin
layer of thallus or naked. Excipulum colourless.
Involucrellum dark, dark brown in thin section, 25–
40 mm thick, contiguous with the exciple, extending
up to two-thirds of the distance from apex to base of
the perithecium. Hymenial gel I+ brown-orange.
Paraphyses absent. Asci fissitunicate, clavate, (25–)
35–40(–50) 3 12–15 mm. Ascospores simple,
colorless, narrowly ellipsoid to subcylindrical, 7.5–
12.5(–14) 3 4–6(–7.5) mm (45 measured).
Ecology. On granite and schist rocks in river
banks, from 0 to 30 cm above minimum water level,
thus totally submerged for most or all of the year. It
belongs to the Submerged to the Fluvial mesic zone
of Gilbert (1996). It is always found in shady sites,
and grows on surfaces of various inclinations. This
species has been found accompanied only by
Verrucaria aquatilis Mudd and also sometimes by a
poorly developed lichen material, belonging in
Verrucaria.
Observations. The specimens from the Iberian
Peninsula coincide with those described by McCarthy
(1995) to a great extent, although some of our
specimens have a thicker thallus, but in any case
correspond to the V. mundula var. deminuta also
described by McCarthy (1995) characterized by
smaller asci (28–35 3 10–15 mm) and broadly
ellipsoid or subglobose ascospores (6–)8.5(–10.5) 3
(4.5–)5.5(–6.5) mm. All the Iberian Peninsula samples
are light to dark brown in color with a smooth
Valcárcel et al.: Verrucaria mundula new to Northern Hemisphere
269
Table 1. Differences between the holotype material of Verrucaria mundula and the Iberian specimens.
Verrucaria mundula
Thallus
Perithecium
Excipulum
Involucrellum
Ascospores
Type material
Iberian Peninsula
Green to greenish black
(15–)20–30(–40) mm thick
Not covered by a thallus layer
0.12–0.18(–0.2) mm diam.
Colorless to light brown
25–35 mm thick
(8.5–)11(–14.5) 3 (3 .5–)5(–7.5) mm
Light to dark brown
24.5–60(–80) mm thick
Covered or not by a fine thallus layer
0.14–0.18 mm diam.
Colorless
25–40 mm thick
7.5–12.5(–14) 3 4–6(–7.5) mm
thallus surface, whereas the Australian specimens are
medium green to greenish black and some of them
have a sparingly rimose thallus. Some perithecia are
covered by a fine thallus layer, a feature that is not
seen in the type material (Table 1). However, we do
not believe that these minor differences justify the
description of the Iberian specimens as a new species.
The ecology of the specimens of both
hemispheres is similar as McCarthy (1995) indicates
that this is a saxicolous, aquatic or semi-aquatic
taxon that grows in shaded or very shaded places on
schists, slate, sandstone, basalt and rhyolites. The
same can be said of the specimens from the Iberian
Peninsula, but they are located only on acid rocks.
Only four species of Verrucaria s.l. with
ascospores less than 15 mm long are known from
freshwater habitats in Europe; these are readily
distinguishable from V. mundula (Table 2).
Verrucaria aquatilis differs in the black thallus, larger
perithecia and more immersed in the thallus,
involucrellum diverging from the excipulum below
(Fig. 1D), and smaller, broadly ellipsoid ascospores.
Verrucaria madida Orange differs in the consistently
dark green-coloured thallus, with green pigment in
the cortex, larger perithecia, and in the asci with 4
(–5) ascospores are apparently unique in the genus
Verrucaria s.l. (Orange 2004). Verrucaria rheitrophila
Zsch. (5Hydropunctaria rheitrophila (Zsch.) Keller,
Gueidan & Thüs) is distinguished by its greenish
thallus (pigment in the cortex, if present, is brown)
with carbonaceous spots (Fig. 1C), typical of the
genus Hydropunctaria (Gueidan et al. 2009), which
are only visible microscopically at times, but usually
present in section, by its irregular involucrellum that
Table 2. Differences between Verrucaria mundula and similar European species.
V. aquatilis
Thallus
Perithecium
Involucrellum
Ascospores
V. madida
V. mundula
Black without
Grey green to dark Light to dark brown
without
carbonaceous spots
green without
carbonaceous
carbonaceous
spots
spots
36–50 (–76) mm thick 40–60 mm thick
24.5–60(–80) mm
thick
Immersed to 2/3
Superficial
1/3 immersed to
immersed
superficial
0.15–0.35 mm diam.
0.2–0.42 mm diam 0.14–0.18 mm diam
Contiguous with
Contiguous with
Contiguous with
excipulum and
excipulum and
excipulum, not
spreading at the
spreading at the
spreading at the
base
base
base
Subspherical (8/ascus) Ellipsoid (4/ascus) Subcylindrical
(8/ascus)
(9–)10.5–13.5(–15) 7.5–12.5(–14) 3
(5–)10(–12) 3
3 6–7(–7.5) mm
4–6(–7.5) mm
4–6(–8.5) mm
V. rheitrophila
V. scabra
Green to greenish
black with
carbonaceous
spots
54–142 mm thick
Greenish to blackish
with carbonaceous
spots
Immersed
(110–)120(–140) mm
thick
Immersed
0.12–0.3 mm diam 0.3–0.65 mm diam
Only around the
Contiguous with
ostiole
excipulum and
spreading at the
base
Ellipsoid (8/ascus) Subcylindrical
(8/ascus)
8.5–12(–15) 3
(9.7–)14.2(–15) 3
5–10 mm
(5–)7.2 (–9.4) mm
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113(2): 2010
generally grows only around the ostiole and by the
more broadly ellipsoid ascospores. Verrucaria scabra
Vzda (Hydropunctaria scabra (Vzda) Keller, Gueidan &
Thüs), has a thicker thallus with internal carbonaceous
spots and a dark basal layer; the perithecia immersed in
the thallus and much broader.
It is likely that V. mundula has been overlooked
previously in the Iberian Peninsula, due to its thin
thallus, small perithecia, and occurrence in shady,
often submerged habitats. It has previously been
recorded from Australia and the South Pacific
(McCarthy 1995, 2000).
Selected specimens examined of V. mundula
P.M. McCarthy var. mundula. AUSTRALIA. NEW SOUTH
WALES: Eden District, Yurammie State Forest,
Chalkhill, Rats Valley Road, off Chalkshills Road,
11 km WNW of Pambula, 55HGV4618713669, on
aquatic shale, [?shaded], 1991, P. M. McCarthy (MEL
1055260); 13 km WNW of Kiama, Minnamurra Falls
Park, Minnamurra River, 56HKE3534790651, on
deeply shaded semi-aquatic sandstone, P.M.
McCarthy 488 (MEL 1055275); Chichester State Forest,
Tributary of Karuah River, 1.5 km SE of Red Cedar
Creek, by Karuah Valley Road, off Wards RiverBerrico Trig. Road, 56HLK8269841560, on deeply
shaded aquatic basalt, 1991, P.M. McCarhy 537 (MEL
1055324). QUEENSLAND: S of Ingham, Mount Spec
National Park, Little Crystal Creek under bridge on
Paluma Road, 55KDU2261597141, on aquatic and
semi-aquatic basalt, 1993, P.M. McCarthy 809 (MEL
1057463). TASMANIA: 4 Km E of Little Swanport, Little
Swanport River, on semi-aquatic dolerite, 1993, P.M.
McCarthy (MEL 2280718). SPAIN. CORUÑA: Vimianzo,
Calo, Rı́o Grande, on granite, 120 m, 1999, C. Pérez
Valcárcel (SANT 11326). HUELVA: Sierra de Aracena,
Alájar, Rı́o Alájar, on schist, 500 m, 2006, C. Pérez
Valcárcel (SANT 11327). LUGO: Rábade, Rı́o Miño, on
schist, 440 m, 1999, C. Pérez Valcárcel (SANT 11628).
OURENSE: Barco de Valdeorras, Entoma, Rı́o Galir, on
schist, 360 m, 2000, C. Pérez Valcárcel (SANT 11329);
Barco de Valdeorras, Xagoaza, Rı́o Mariñán, on
schist, 460 m, 2000, C. Pérez Valcárcel (SANT 11330).
PONTEVEDRA: A Golada, Seixo, Rı́o Ulla, on granite,
300 m, 2000, C. Pérez Valcárcel (SANT 11331). ZAMORA:
Parque Natural de Sanabria, Sotillo de Sanabria, Rı́o
Truchas, on granite, 1000 m, 2005, C. Pérez Valcárcel
(SANT 11332).
ACKNOWLEDGMENTS
This study has been financed by the ‘‘Flora liquenologica
Ibérica’’ project DGI CGL2004-04795-C04-03. The authors are
most grateful to the curators at the SANT and MEL herbaria for
the material loaned.
LITERATURE CITED
Aptroot, A. & M. R. D. Seaward. 2003. Freshwater lichens.
Fungal Diversity Research Series 10: 101–110.
Asta, J., G. Clauzade & C. Roux. 1973. Étude de quelques
groupements lichéniques saxicoles-calcicoles du parc
national de la Vanoise. Travaux Scientifiques du Parc
National de la Vanoise 3: 73–104.
Coste, C. 2005. Lichens et communautés lichéniques
hydrophiles et subhydrophiles saxicoles-calcifuges du
département du Tarn (France). Bulletin de la Société
Botanique du Centre-Ouest, Nouvelle série 36:
577–596.
Gilbert, O. L. 1996. The lichen vegetation of chalk and
limestone streams in Britain. Lichenologist 28: 145–159.
——— & V. J. Giavarini. 1997. The lichen vegetation of acid
watercourses in England. Lichenologist 29: 347–367.
Gueidan, C., S. Savic, H. Thüs, C. Roux, C. Keller, L. Tibell,
M. Prieto, S. Heidmarsson, O. Breuss, A. Orange, L.
Fröberg, A. Amtoft Wynns, P. Navarro-Rosinés, B.
Krzewicka, J. Pykälä, M. Grube & F. Lutzoni. 2009.
Generic classification of the Verrucariaceae
(Ascomycota) based on molecular and morphological
evidence: recent progress and remaining challenges.
Taxon 58: 184–208.
Hawksworth, D. L. 1989. Notes on aquatic species of Verrucaria
in the British Isles. Lichenologist 21: 23–28.
Keller, C. 2000. Die Wasserflechten der Teigitsch zwischen der
Langmannsperre und dem Kraftwerk Arnstein (Steiermark,
Österreich). Herzogia 14: 49–58.
McCarthy, P. M. 1995. Aquatic species of Verrucaria in Eastern
Australia. Lichenologist 27: 105–126.
———. 2000. The lichens of Rarotonga, Cook Islands, South
Pacific Ocean I: Pyrenocarpous taxa. Lichenologist 32:
15–47.
Ménard, T. 1997. Étude phytosociologique et écologique des
peuplements lichéniques saxicoles calcifuges du sud-est de la
France. Thèse en sciences, Univ. Aix-Marseille III. Aix-enProvence.
Orange, A. 2004. A remarkable new freshwater Verrucaria from
Europe. Lichenologist 36: 349–354.
Pereira, I. 1992. Flora, vegetación y ecologı́a de los lı́quenes
acuáticos de España. Tesis Doctoral (Inéd.). Universidad de
Barcelona.
——— & X. Llimona. 1986–1987. Un aspecte poc conegut del
Montseny: Les comunitats de lı́quens hidròfils. Primeres
dades. Jornadas de Recerca Naturalista at Montseny 33–35.
Diputació de Barcelona.
Valcárcel et al.: Verrucaria mundula new to Northern Hemisphere
——— & ———. 1992. Algunos lı́quenes hidrófilos de la
Sierra del Moncayo. Folia Botanica Miscellanea 8: 55–59.
Swinscow, T. D. V. 1968. Pyrenocarpous lichens: 13. Freshwater species of Verrucaria in the British Isles. Lichenologist
4: 34–54.
Thüs, H. 2002. Taxonomie, Verbreitung und Ökologie silicoler
Süßwasserflecheten im außeralpinen Mitteleuropa.
Bibliotheca Lichenologica 83: 1–214.
——— & M. Schultz. 2009. Fungi: Lichens. Pages xxx–xxx. In
B. Büdel, G. Gärtner, L. Krienitz, H. R. Preisig & M.
Schagerl (eds.), Süßwasserflora von Mitteleuropa Freshwater Flora of Central Europe. Vol 21/1: 1–224.
Valcárcel, C. P., M. J. Sánchez-Biezma & R. Carballal. 1999.
Lı́quenes de los cursos de agua de la provincia de Lugo
(Galicia, España). I: Especies con peritecios. Nova Acta
Cientı́fica Compostelana (Bioloxı́a) 9: 21–31.
271
——— & R. Carballal. 2002. Lı́quenes pirenocárpicos de los
rı́os y arroyos de Galicia (España). Cryptogamie, Mycologie
23: 245–271.
Wirth, V. 1972. Die Silikatflechten-Gemeinschaften im
ausseralpinen Zentraleuropa. Dissertationes Botanicae 17:
1–325.
Zschacke, H. 1927. Die mitteleuropäischen Verrucariaceen V.
Hedwigia 67: 45–85.
———. 1933–34. Epigloeaceae, Verrucariaceae und
Dermatocarpaceae. Pages 44–480 (1933) and 481–695
(1934). In A. Zahlbruckner (ed.), Dr. L. Rabenhorst’s
Kryptogamen-Flora von Deutschland, Österreich und der
Schweiz. 2. Aufl., Bd. 9, Abt. 1, Teil 1. Akademische
Verlagsanstalt, Leipzig.
ms received June 15, 2009; accepted November 12, 2009.