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New Halecania species (Catillariaceae) from Europe and
South America
PIETER P. G. VAN DEN BOOM
Arafura 16, NL-5691JA Son, The Netherlands
e-mail: pvdboom@zonnet.nl
ABSTRACT.Four new species of Halecania,H. etayoana (saxicolous) and H. fuscopannariae
(lichenicolous) both from South America, and H. laevis and H. pannarica, both saxicolous
from Europe, are described as new to science. Their ecology, distribution and chemistry are
discussed.
KEYWORDS.British Isles, central Europe, Chile, Ecuador, taxonomy, saxicolous,
lichenicolous, Catillariaceae, Halecania.
¤¤¤
The genus Halecania was originally described by M.
Mayrhofer (1987) to accommodate six species (five
saxicolous, one bryicolous) from Europe, formerly
included in Lecania s. lat. A key to the species occurring
in the British Isles was given by Coppins (1992) who
treated four saxicolous and one corticolous species and
two more species were described from the British Isles
by Fryday and Coppins (1996). In a recent publication
(van den Boom & Elix 2005) some new species from
Asia were described and further information was given,
including a world-wide key to the known species of the
genus. Van den Boom and Elix (2005) regarded
Halecania as a subcosmopolitan genus, being widely
distributed in Europe from the most northern boreal
and arctic areas to the Mediterranean region, as well as
occuring in Asia, Australia, New Zealand and North
and South America. Several species are lichenicolous,
at least in the juvenile state. In the present paper three
new saxicolous and one new lichenicolous species are
described from Europe and South America
MATERIALS AND METHODS
Measurements of ascospores and conidia were
made in water and 10% KOH at 4003or 10003
magnification. Amyloid reactions were tested using
Lugol’s iodine solution (I/KI). Specimens were
identified by TLC to check the content of secondary
metabolites (Culberson & Ammann 1979; Culberson
& Johnson 1982; Elix & Ernst-Russell 1993) and/or
by HPLC (Elix et al. 2003).
THE NEW SPECIES
Halecania etayoana Palice, van den Boom & Elix
sp. nov.Fig. 1
Thallus parvus, plus-minusve arcte affixus, ad 1
mm crassus, marginaliter areolis angularibus vel
rotundis, indistincte lobatus, lobis ad 0.5(–0.6) mm
longis; superficies fulva vel griseo-fuscescens, sine
corticem. Apothecia adnata, lecanorina vel lecideina,
0.2–0.6 mm diam., discus fuscus vel fusco-niger,
epruinosus; margo thallinus vel biatorinus, planus,
tenuis, ad 0.05 mm crassus; apices paraphysarum
plerumque fusco-capitatae. Ascosporae hyalinae,
uniseptatae, ellipsoideae vel oblongo-ellipsoideae
(7–)8–11(–12) 32.5–3(–3.5) mm (sine strato
perisporico), stratum perisporicum gelatinosum,
1.5–2 mm crassum in KOH. Conidia bacilliformia vel
ellipsoidea, 3–4(–4.5) 30.9–1.1 mm. Substantiae
lichenicae absentes. Habitat ad saxa montana.
TYPE:ECUADOR.PROV.TUNGURAHUA: Ban
˜os–Rı
´o
Pastaza, near the cascade Bascu
´n close to the bridge
The Bryologist bryo-112-04-02.3d 23/4/09 16:27:52 1
The Bryologist 112(4), pp. 000–000 0007-2745/09/$0.00/0
Copyright E2009 by The American Bryological and Lichenological Society, Inc.
‘‘Puente San Francisco,’’ on damp schistose boulders
on the bank of the river, 1800 m, 31 Jul 1999, Z.
Palice 2571 &J. Etayo (PRM, holotype; HB.VAN DEN
BOOM, isotype).
Description. Thallus crustose, effuse, areolate to
knobby-warted, becoming squamulose; warts 0.2–0.6
mm wide; areoles 0.3–1 mm wide, to 0.5(–0.6) mm
long, 0.3 mm high, angular to roundish, sometimes
with upturned to erected, weakly lobulate margins,
incised, sometimes imbricate; upper surface pale
brown, gray-brown, dark gray to rarely greenish gray;
upper cortex not differentiated or occasionally
paraplectenchymatous, cells 2–5mm, without
epinecral layer. Prothallus sometimes visible, thin
and pale to medium brown. Photobiont
chlorococcoid, algal cells 5–12 mm diam., scattered
throughout the thallus. Apothecia 0.2–0.6 mm wide,
roundish, frequent, semi-immersed, adnate to
slightly sessile, scattered to grouped, sometimes
crowded; disc red-brown to dark brown or almost
black, paler (reddish brown) when wet, plane,
sometimes weakly convex, epruinose; thalline exciple
rarely present, mainly developed at the underside of
the apothecia, concolorous with the thallus, swollen
when young, narrower when old, sometimes
crenulate, often excluded, without cortex, with a
colorless, amorphous surface layer, intermixed with
photobiont cells, with algae filling the entire margin
and abundant at the underside of the hypothecium,
to 0.05 mm wide; true exciple well developed and
appearing as a small (to 0.05 mm) dark rim, giving
the apothecia mostly a lecideine appearance, hyaline
at inner part, dark (red) brownish towards the outer
rim in which hyphae tips are well developed, dark
brown-pigmented and widened apically to 5 mm;
hymenium 35–45 mm high, hyaline; epihymenium
brown, medium- to dark-brown, without granules,
6uniformly colored, K-, N-; hypothecium to 50 mm
high, hyaline; paraphyses slightly swollen (clavate)
and short-celled in the upper third, mid-hymenium
cells 1.5–2 mm wide, apical cells to ca. 5 mm wide and
dark brown-pigmented, simple to occasionally
branched; asci narrowly clavate, ca. 40–50 312–20
mm, Catillaria-type, 8-spored; ascospores 6ellipsoid,
slightly clavate or ovoid, (7–)8–11(–12) 32.5–3
(–3.5) mm, 1-septate, the septum sometimes central,
but often situated toward the lower end, thin-walled,
halonate, perispore swelling to ca. 2 mm in KOH,
often not well developed. Pycnidia occasionally
present, inconspicuous, immersed in the thallus, ca.
50–80 mm in diam., dark brown around the ostiole,
hyaline below; conidia simple, short bacilliform to
oblong or ellipsoid, 3–4(–4.5) 30.9–1.1 mm.
Chemistry. Thallus K-, C-, KC-, P-; no chemical
compounds detected by TLC or HPLC.
Etymology. Halecania etayoana is named in
honor of our friend and colleague Javier Etayo for his
contributions to lichenology in Central and South
America.
Distribution and ecology. This new species
occurs on acidic rocks, including granite and
sandstone, in the supralittoral zone, in open inland
areas at elevation of 1800 m. It is currently known
only from the type locality.
Notes. Halecania etayoana is characterized by
having the smallest ascospores of any other known
Halecania species. Among the other known Halecania
species growing on acidic rocks, only H. pannarica and
H. subsquamosa (Mu
¨ll. Arg.) van den Boom & H.
Mayrhofer have a pale brown to grayish thallus that
could be considered squamulose, but this latter species
has some unknown chemical compounds and the
ascospores are longer and wider (van den Boom & Elix
2005). Halecania pannarica is a sorediate species.
Another Halecania species with biatorine-looking
apothecia is H. elaeiza (Nyl.) M. Mayrhofer, but that
species has a thin, smooth and rimose thallus and
occurs on calcareous rocks. In habit, H. etayoana
resembles Lecania inundata (Hepp ex Ko
¨rb.) M.
Mayrhofer, especially the thallus, but this latter species
The Bryologist bryo-112-04-02.3d 23/4/09 16:28:25 2
Figure 1. Habit of Halecania etayoana (holotype). Warted-
squamulose-areolate thallus with apothecia. Scale 50.5 mm.
0THE BRYOLOGIST 112(4): 2009
has different asci, ascospores, paraphyses and
epihymenium, and has a relative well-developed
thalline excipulum.
Halecania fuscopannariae Etayo & van den Boom
sp. nov.Fig. 2
Thallus absens. Apothecia biatorina, ad 0.1–
0.25(–0.3) mm lata, in thallo Fuscopannariae
corticolae crescens. Discus convexus, fuscus. Ascosporae
10–12(–15) 35.5–8.5 mm, 1-septatae, perisporae tenui
ad crassiore, 0.5–2mm. Lichenicola.
TYPE:CHILE.PROV.ANTA
´RTICA CHILENA: Isla
Navarino, Puerto Williams, senda que atraviesa
Virgen de Lourdes hacia Barranca Guarriaco por
zona militar, 54u569460S, 67u34952.20W, 90 m, 14 Jan
2005, Go
´mez, Søchting, Vilches & J. Etayo 22503 (MAF,
holotype; HB.ETAYO, isotype).
Description. Thallus and prothallus absent.
Apothecia scattered on thallus of Fuscopannaria sp.,
lecideine in appearance, rounded, 0.1–0.25(–0.3) mm
diam.; margin thin to lacking; disc flat to slightly
convex, dark brown to nearly black; thalline
excipulum to 50 mm wide, to 80 mm high, composed
of pseudoparaplectenchymatous cells; algal cells few,
5–10 mm diam.; hymenium hyaline, to 100 mm high;
epihymenium moderately to dark brown, N-;
paraphyses simple to branched near the tips, septate,
apical cells to ca. 3.5 mm wide and 6dark brown-
pigmented, mid-hymenium cells 1–1.5 mm wide;
hypothecium hyaline; asci clavate, 30–37 311–16
mm, Catillaria-type, 8-spored; ascospores ellipsoid, or
with one pointed end, 10–12(–15) 35.5–8.5 mm, 1-
septate, not or slightly constricted at septum, septum
often situated toward the lower end, thin-walled,
halonate, perispore thin to relative wide, 0.5–2 mm.
Pycnidia unknown.
Chemistry. Not tested by TLC (apothecia very
small), spot test on apothecia (in sections) K-, C-, P-.
Ecology. Only known on an unidentified
Fuscopannaria growing on Nothofagus pumilio. The
host is a squamulose, abundantly fertile species.
Notes. Several species of Halecania are
lichenicolous but all have at least small thalli. For
example, although the thallus of H. alpivaga (Th. Fr.)
M. Mayrhofer can be strongly reduced, the thalline
margin of the apothecia is always clearly present.
Halecania fuscopannariae is the first species in the
genus which has been found with apothecia only; no
thallus is developed. However in the thalline
excipulum algal cells are present.
Halecania laevis M. Brand & van den Boom sp.
nov.Fig. 3
Thallus saxicola tenuis, crustaceus, Apothecia
parva lecanorina, margine thallino 0.3–0.35 mm
diam., late appressa ad sessilia, plana, disco fusco-
nigro, margine tenui 0.03–0.05 mm
paraplectenchymatico. Asci octospori, sporae
ellipsoideae ad late fusiformes, 8.6–10.7 33.1–3.4 mm,
1-septatae, perisporio tenui (0.5 mm). Pycnidia semper
abundantes, conidia breve ellipsoidea 2.8–2.9 31.1
mm. Thallus sine substantias lichenicas. Habitat ad
saxa acida litoralia.
TYPE:FRANCE.DEPT.FINISTE
`RE: Goulien-Cap,
Sizun, cove 0.4 km N of Kerisit, N-exposed coastal
The Bryologist bryo-112-04-02.3d 23/4/09 16:28:29 3
Figure 2. Habit of Halecania fuscopannariae (holotype). One
apothecium in the center among three apothecia of Fuscopannaria
and one apothecium, below, at the right corner. Scale 50.5 mm.
Figure 3. Habit of Halecania leavis (holotype). Scale 5
0.5 mm.
van den Boom: New species of Halecania 0
gneiss rocks, at sheltered bay, base of high cliff,
supralittoral zone, 48u4.39N, 4u35.39W, 22 Jul 1997,
M. Brand 36859 (L, holotype; HB.BRAND,HB.VAN DEN
BOOM, isotypes).
Description. Thallus crustose, small to sometimes
rather large, to 6 cm wide, 0.13–0.25 mm thick,
sometimes rimose to areolate; areoles occasionally
ascendant at the edge and subsquamulose, upper
surface smooth, matte grayish brown; epinecral layer
mostly present, to 20 mm, of loose issue, filled with air;
cortex inconspicuous, cells at the outer rim globose, to
5mm diam., brown-pigmented, N-. Prothallus
inconspicuous or absent, brownish gray. Apothecia
abundant, small, to 0.3(–0.35) mm diam., broadly
appressed to sessile, plane, disc brownish black; margin
thin 0.03–0.05 mm, to evanescent, concolorous with
the disc, algae in under part only, composed of
paraplectenchyma, cells 3 35mm, outer rim of
margin, especially at top, with dark brown-pigmented
cells; hymenium 40–50 mm; epihymenium brown
caused by the pigmented top of the paraphyses, N-;
paraphyses not or slightly conglutinated, widened at
apices, to 5(–6) mm wide, in mid-hymenium 1.2–1.8
mm diam.; asci 8-spored, 25–35 38–10 mm, Catillaria-
type; ascospores 8.6–10.7 33.1–3.4 mm, 1-septate,
ellipsoid to broadly fusiform, halonate, perispore to 1.5
mm, sometimes indistinct.
Pycnidia always abundant, immersed, ca. 60 mm
diam., hyaline at base, top dark brown pigmented.
Conidia short ellipsoid, 2.8–2.9 31.1 mm.
Chemistry. No chemical compounds found by
TLC.
Distribution and habitat. Halecania laevis is
known from western Ireland and in western Bretagne
(France). It is a maritime species, occurring in the
supralittoral zone on acidic stones of sheltered, N-
exposed coastal areas. Accompanying species include
Lecanora helicopis,Solenopsora holophaea,Halecania
ralfsii,Verrucaria maura,Catillaria chalybeia,
Pertusaria pseudocorallina,Lecidella cf. asema,
Lecanora dispersa and Diplotomma chlorophaea.
Notes. This new species differs from Halecania
ralfsii (Salwey) M. Mayrhofer, which occurs in the
same habitat, in chemistry, smaller apothecia, smaller
ascospores and the lack of bluish green N+red
pigment in surface tissues. Halecania spodomela
(Nyl.) M. Mayrhofer differs in having a granular
thallus and occurring in habitats above the
supralittoral zone. Halecania elaeiza may also be a
related species but that species is known only from a
different substrate (limestone in Romania) and has
much wider ascospores.
Halecania laevis is easily mistaken for other
species of Catillariaceae, especially Catillaria chalybeia.
The seemingly biatorine black apothecia and the rather
thin and smooth thallus resembles this latter species
and even the swollen dark paraphyses tips and the
small ascospores. Catillaria chalybeia can be
distinguished by its smaller, non-halonate ascospores
and its brown hypothecium. Specimens of Catillaria
subviridis (HB.BRAND) have been compared with
Halecania laevis.Catillaria subviridis has black
apothecia with a persistent black margin, non-halonate
ascospores that are clearly larger, 10–16 34.5–6 mm,
paraphysis tips that are blackish pigmented, conidia
that are 2.2–2.5 31.3 mm, and it occurs on nutrient-
enriched rocks, in a different community. The
difficulty to distinguish it could be the reason why H.
laevis has been overlooked for so long.
Additional specimens examined. IRELAND.
CONNEMARA: 3 km SW of mouth of Killary Harbour,
coastal area S of mouth of Culfin River, 53u36.3N,
9u54.49W, on schistose rocks above beach, 19 May
2000, M. Brand 40696 (HB.BRAND,HB.VAN DEN BOOM);
KERRY: Dingle peninsula, Clogher Head, N-side, steep
N-facing cliff, protruding rocky point, 52u99N,
10u289W, on conglomerate rocks, 11 May 2000, M.
Brand 40343 (HB.BRAND). FRANCE.FINISTE
`RE: Crozon
Peninsula, Cap de La Che
`vre, SE side, Men Coz,
48u10.29N, 4u339W, 18 Jul 1997, M. Brand 36712 (HB.
BRAND,HB.VAN DEN BOOM).
Halecania pannarica M. Brand & van den Boom
sp. nov. Figs. 4,5
Halecania giraltiae similis sed apothecia ad 0.4
mm lata, ascospores, (9–)10–14(–18) 34.5–5.5(–6)
mm, 1-septae, haud constricta at septum, in statu
juvenili perisporio ad 1.5 mm lato. Conidia ellipsoidea,
ca. 2.2–2.8 31–1.2 mm. Thallusacidum pannaricum
continens. Habitat ad saxa acida in Alpium.
TYPE:SWITZERLAND.DEPT.VALAIS: Val d’He
´rens, 0.5
km WNW of Evole
`ne, shore of Borgne, just N of
bridge, 46u6.99N, 7u29.49E, on large boulders of schist
in stream, 1340 m, 6 Aug 1990, M. Brand 24858 (L,
holotype).
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0THE BRYOLOGIST 112(4): 2009
Description. Thallus of small squamules,
appressed to the substrate, on Aspicilia cf.
caesiocinerea or A. candida, partly directly on acidic
rock, with many fine crystals; squamules 0.3–1.3 mm
wide, the smaller ones roundish, the larger ones more
lobulate, to 0.17 mm thick; upper surface plane,
matte, pale gray to grayish brown; cortex and
epinecral layer absent; algal cells scattered
throughout the thallus, roundish, ca. 10 mm, with
many haustoria; underside with loose hyphae; soralia
always present, laminal on squamules, roundish,
plane, when young weakly crateriform, bluish black,
soredia dark gray to black, ca. 22 mm wide, of algae
mainly enclosed in a outer layer of hyphae, brownish
or obscure bluish green, partly pigmented at the
outer rim. Prothallus absent. Apothecia sometimes
present, scattered to crowded, to ca. 0.4 mm wide,
initially immersed, becoming widely adnate, with a
lecanorine margin; disc moderately brown, matte;
excipulum hyaline at inner part, brownish at rim,
abundantly filled with algae; hymenium ca. 60 mm;
epihymenium brown, N-; paraphyses not or slightly
conglutinate, septate, widened at apices, to 5(–6) mm
and dark brown-pigmented; asci 8-spored, 35–50 3
10–18 mm, Catillaria-type; ascospores (9–)10–14
(–18) 34.5–5.5(–6) mm, 1-septate, not or rarely
constricted at septum, halonate, mainly when young
with perispore; perispore to 1.5 mm wide. Pycnidia
immersed to semi-immersed, ca. 70 mm diam.;
conidiogenous cells pyriform, ca. 5 32.5 mm;
conidia ellipsoid, 2.2–2.8 31–1.2 mm.
Chemistry. Thallus with pannaric acid by TLC.
Thallus extract P+orange.
Distribution and ecology. So far this new species
is known only from central Europe, from France and
Switzerland. Halecania pannarica is known from large
acidic boulders in streams and meadows in the Alps
(1150–1820 m), in open situations. It has been found
on Acarospora badiofusca,Aspicilia caesiocinerea and
Rhizocarpon geminatum, in communities with further
species such as Acarospora impressula,Aspicilia
candida,Lecanora garovaglii and Protoparmelia
cupreobadia. On sandstone it has been found on ridges,
at 2300 m, on a sterile crust (cf. Aspicilia candida), in a
community with Acarospora impressula,Lecidella
stigmatea and Verrucaria sphaerospora.
Notes. Morphologically, H. pannarica is easily
overlooked for H. giraltiae van den Boom & Etayo,
but this latter species contains argopsin and
norargopsin and occurs mainly in Spain and
Portugal. Halecania giraltiae has a darker thallus,
larger apothecia, to 0.6 mm, somewhat shorter and
wider ascospores (9–12 35.5–6.5 mm) and conidia
that are more bacilliform and 3–4.5 31mm.
However H. giraltiae is also known lichenicolous on
genera such as Rhizocarpon and Aspicilia (van den
Boom & Etayo 2001). Both species have a P+
yellowish-orange spot test reaction, so they are best
separated by TLC. However the extract in acetone is
also different, argopsin +norargopsin is dry, white,
with amorphous, nonrefractive crystals; pannaric
acid extract in acetone is dry, white but with many
small polarized parallellogram-form crystals.
Additional specimens examined. SWITZERLAND.
URI: 9 km SW of Altdorf, 0.3 km SE of Surenenpass,
sandstone outcrops at top of ridge, 46u50.29N,
The Bryologist bryo-112-04-02.3d 23/4/09 16:28:37 5
Figure 4. Habit of Halecania pannarica (M. Brand 45312).
Thallus with soralia. Scale 51 mm.
Figure 5. Habit of Halecania pannarica (holotype). Crowded
apothecia. Scale 50.5 mm.
van den Boom: New species of Halecania 0
8u339E, 2340 m, 23 Sep 1991, M. Brand 26904 (HB.
BRAND); VALAIS: Val d’He
´rens, S of Evole
`ne, between
Les Haude
`res and Arolla, S of La Gouille, SE slope
with boulders in meadow, grid ref. 603.9-99.9, 1820
m, 21 Jul 1990, P. van den Boom 9715 (HB.VAN DEN
BOOM). FRANCE.ALPES MARITIMES: 2.5 km NW of St.
Martin d’Entraunes, La Berarde, schistose rock in a
small valley, 44u9.69N, 6u45.19E, 1150 m, 20 Jun
2002, M Brand 45312 (HB.BRAND).
ACKNOWLEDGMENTS
I am grateful to Harrie Sipman for correcting the text,
especially the Latin descriptions and diagnosis, and to Alan
Fryday and Brian Coppins who reviewed the manuscript.
LITERATURE CITED
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B. J. Coppins, D. L. Hawksworth, P. W. James & D. M.
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Culberson, C. F. & K. Ammann. 1979. Standardmethode zur
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¨nnschichtchromatographie von Flechtensubstanzen.
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——— & A. Johnson. 1982. Substitution of methyl tert.-buthyl
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Elix, J. A. & K. D. Ernst-Russell. 1993. A Catalogue of
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¨ber die saxicolen Arten der
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ms. received August 13, 2008; accepted April 1, 2009.
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