MYCOTAXON
Volume 109, pp. 171–179
July–September 2009
Notes on the lichen genus Bacidia s.l.
(lichenized Ascomycota) in the Cape Verde Islands
and new lichen records for the archipelago
Esteve Llop1, 2* & Pieter van den Boom3
*ellop@ub.edu
Departament Biologia Vegetal-Botànica, Universitat de Barcelona
Avda. Diagonal 645, 08028 Barcelona, Spain
2
Centro de Biologia Ambiental, Faculdade de Ciências da Universidade de Lisboa
Campo Grande, Bloco C2, 5º Piso, sala 37, 1749-016 Lisboa, Portugal
1
pvdboom@zonnet.nl
Arafura 16, 5691JA Son, the Netherlands
3
Abstract — Five species belonging in Bacidia s. l. are newly reported from Cape
Verde Islands and discussed. Four species belong to Bacidia: B. atlantica, B. polychroa,
B. subincompta, and B. trichosperma; and one species to Bacidina: B. pallidocarnea. Four
other species are also newly reported: Buellia dispersa, Cresponea flava, Fellhaneropsis
vezdae, and Toninia submexicana. he new combination Bactrospora thyrsodes is
proposed for Bacidia thyrsodes.
Key words — Atlantic islands, biogeography, Macaronesia, new records, taxonomy
Introduction
Mies (1993) published a critical compilation of species belonging to Bacidia
De Not., considered in a wide sense, found in the Cape Verde Islands. In his
checklist, Mies cited five species without further comment, reporting three
species (Bacidia effusa, Bacidia sp. A, and Bacidia sp. B) from just one island,
B. laurocerasi from two islands, and B. thyrsodes from five islands.
he aim of our current paper is to revisit Bacidia s. l. in the Cape Verde
archipelago in greater depth and to compare the distribution of the treated
species in the Macaronesian archipelagos.
Material and methods
he Cape Verde Islands are situated in the Atlantic Ocean located near the west
coast of Africa between 14°48’ and 17°12’ N, and 25°42’ to 22°41’W.
172 ... Llop & van den Boom
his study is based on specimens identified as Bacidia collected on Cape Verdean
islands and borrowed from the herbaria BM and M, as well as on samples
collected by P. & B. v. d. B. in July 2006, from Santo Antão, São Tiago and São
Vicente of the archipelago. hese have been examined by stereomicroscope and
light microscope using the standard techniques.
Results
he study of the available collections has shown that five species belonging in
Bacidia s. l. occur in the Cape Verde Archipelago. One species, Bacidia thyrsodes,
is excluded from Bacidia and a new combination is made. In addition, four
species misidentified as Bacidia are reported as new for the lichen flora of Cape
Verde.
Bacidia atlantica (Müll. Arg.) Zahlbr.
hallus grey green, granular to areolate, areoles sometimes resembling
squamulose when margins rise; surface disaggregating into goniocysts 30–60
µm wide; upper cortex prosoplectenchymatous. Algae chlorococcoid, 5–8 µm
diam. Apothecia orange to reddish, 0.35–0.65 mm wide; with a well developed
proper margin, paler than the disc; disc flat, finally slightly swollen, with a
darker rim. Exciple colourless, prosoplectenchymatous, made of branched
and anastomosed hyphae with cell lumina of 12 × 2–3.5 µm. Hypothecium
prosoplectenchymatous, yellowish, reacting K+ intensifying, rubella-orange
pigment present. Hymenium colourless, 60–70 µm thick; upper hymenium not
differentiated, sometimes small crystals present that solve in KOH. Paraphyses
not or slightly agglutinated, not anastomosed, apices not swollen, 1.5 µm thick.
Asci clavate, 8-spored, 40–60 × 7–10 µm, Bacidia-type. Ascospores colourless,
long bacilliform, not or slightly tapering to one end, (20–)22–39 × 2–3(–3.5)
µm, with 3–5 septa. Pycnidia not seen.
Because of the thallus morphology, B. atlantica resembles the pantropical B.
medialis (Tuck.) B. de Lesd. However, the rim of the exciple is pigmented pale
orange-brown and ascospores are shorter and narrower in the latter.
his taxon was previously known only from Ascension Island, from where
it was described. his is the first report of this species for the Cape Verde
archipelago. Its current range suggests a tropical Middle Atlantic distribution,
although it could have been overlooked elsewhere.
Specimens examined - Santo Antão: SW of Vila das Pombas, Figueiral de Paúl,
SW part of the valley, Chã de Padre, small coffee plantation, scattered mixed trees, acidic
outcrops, 25º 03.0’ W, 17º 07.0’ N, 195 m, on Coffea, 21.VII.2006. P. & B. v. d. Boom
36857 (hb. v. d. Boom). São Tiago: W of São Domingos, WNW of Rui Vaz, along path
to “Monte Tchopa”, E of telecommunication station, hilly area with mixed trees, 1035 m,
on Mangifera, 8.VII.2006, P. & B. v. d. Boom 36424 (hb. v. d. Boom); ibid. on Hibiscus, P.
& B. v. d. Boom 36418 (hb. v. d. Boom).
Bactrospora thyrsodes nom. nov. (Cape Verde) ... 173
Bacidia polychroa (h. Fr.) Körb.
Most of the specimens identified as B. polychroa have been previously identified
under B. effusa (Sm.) Trev., a synonym for Bacidina assulata (Körb.) S. Ekman,
and Bacidia laurocerasi (Delise ex Duby) Zahlbr. Bacidia polychroa differs
from Bacidina assulata in the size of ascospores, which are (30–)40–60(–75)
× (2–)2.5–4 µm in the former, and (28–)32–49(–54) × 1–2 µm in the latter;
the structure of the exciple, prosoplectenchymatous and paraplectenchymatous
respectively; and asci. he differences between B. polychroa and B. laurocerasi
are based on the non-soluble pigments and the size of ascospores. he former
has a red pigment (polychroa-red according to Meyer & Printzen, 2000) in
the exciple, hypothecium, and subhymenium, which reacts K+ purplish.
his pigment is also present, mixed with a low concentration of bagliettoanagreen pigment, in the epihymenium. Bacidia laurocerasi lacks the polychroared pigment in exciple, hypothecium, and subhymenium. However, a certain
amount of rubella-orange pigment is found, which reacts K+ intensifying
the yellow or orange colour. In the epihymenium, the pigment is a mixture
of laurocerasi-brown and bagliettoana-green. Ascospores of B. polychroa are
slightly shorter and wider than those of B. laurocerasi, although this is not the
best character by which to distinguish these taxa.
Bacidia polychroa has a boreal to temperate distribution, mainly in areas
with humid climate. his species has not been recorded from the Macaronesia,
but a careful study of samples determined as B. laurocerasi would result in
misidentifications of B. polychroa, basically because some authors have included
the concept of the latter into B. laurocerasi.
Specimens examined - Santo Antão: S of Ribeira Grande, Corda, centre of village,
outcrops and roadside trees along main road, 1060 m, on Pinus, 17.VII.2006, P. & B. v.
d. Boom 36660 (hb. v. d. Boom); ibid., on Acacia sp., P. & B. v. d. Boom 36701 (hb. v. d.
Boom). São Nicolau: Mt. Gordo, NW-Hang, an Säumen von Euphorbia tuckeyana,
24’21º W, 16’38º N, 940 m, Expos. NW, 29.IX.1988, B. Mies 960i (M-0142068). São
Tiago: W of São Domingos, Rui Vaz, centre of village, mixed trees and outcrops along
small road, 825 m, on Casuarina, 07.VII.2006, P. & B. v. d. Boom 36336 (hb. v. d. Boom);
ibid., on unidentified roadside tree, P. & B. v. d. Boom 36349 (hb. v. d. Boom). W of São
Domingos, WNW of Rui Vaz, along path to “Monte Tchopa”, E of telecommunication
station, hilly area with mixed trees, 1035 m, on Mimosaceae, 8.VII.2006, P. & B. v. d.
Boom 36440 (hb. v. d. Boom); ibid. on shrub, P. & B. v. d. Boom 36445 (hb. v. d. Boom);
ibid. on unidentified big tree, P. & B. v. d. Boom 36448 (hb. v. d. Boom). W of São
Domingos, N of Rui Vaz, along village, on rocky mountain, with some shrubs, 855 m,
on shrub, 9.VII.2006, P. & B. v. d. Boom 36463, 36498, 36504 (hb. v. d. Boom). Monte
Verde, just below top of the mountain, NW slope with acidic outcrops, shrubs and
±scattered small trees, 700 m, on shrub, 15.VII.2006, P. & B. v. d. Boom 36616 (hb. v. d.
Boom); ibid. on rotting trunk of Agave, P. & B. v. d. Boom 36606 (hb. v. d. Boom). São
Jorge das Orgãos, am Staum von Ceratonia siliqua, 630 m, Expos. NE, 16.IX.1988, B.
Mies 840n (M-0142052, 0142053).
174 ... Llop & van den Boom
Bacidia subincompta (Nyl.) Arnold
Features of the examined samples, such as the presence of bagliettoana-green
pigment in the outer part of the exciple and upper hymenium, and reddish
brown hypothecium, agree with Bacidia subincompta. However, the ascospores
are slightly longer than European material; they measure 32–45 µm for 20–36
(–40) µm in European samples (Purvis et al. 1992, Llop 2007). he number
of septa is also higher, 7 to 11 in the Cape Verdean samples for 5 to 7 in the
European collections.
his species has a boreal to temperate distribution, and is known from
Madeira (Hafellner 1995) and Canary Islands (van den Boom & Etayo 2006).
Specimens examined - Santo Antão: S of Ribeira Grande, SE of Corda, N of trail
203, from Chã de Mato to Losnã, small (secondary) trail to Fajã de Baixo, outcrops,
bouldres and walls along trail, 25º 04.5’ W, 17º 08.1’ N, 975 m, on Eucalyptus, 18.VII.2006,
P. & B. v. d. Boom 36774 (hb. v. d. Boom); ibid., Corda, centre of village, outcrops and
roadside trees along main road, 25º 05.3’ W, 17º 07.9’ N, 1060 m, on Acacia, 18.VII.2006,
P. & B. v. d. Boom 36941 (hb. v. d. Boom).
Bacidia trichosperma (Müll. Arg.) Zahlbr.
hallus pale greyish to green, granulose; granules up to 50 µm wide;
hypothallus byssoid. Apothecia flesh to yellowish cream, 0.25–0.50 mm diam.;
proper margin slightly thick, disc flat to rather swollen. Exciple colourless to
pale yellowish at the basis, prosoplectenchymatous, made of branched and
anastomosed hyphae with cell lumina of 5–10 × 2–3 µm; margin with a byssoid
aspect. Hypothecium prosoplectenchymatous, colourless to pale yellowish,
rubella-orange pigment present. Hymenium colourless, 40–45 µm thick;
upper hymenium not differentiated. Paraphyses not to slightly agglutinate,
not branched, not anastomosed; apical cells not swollen, 1–1.5 µm thick. Asci
clavate, 8-spored, 30–35 × 7 µm, Bacidia-type. Ascospores colourless, acicular,
20–27 × 1–2 µm, 3–5 septa. Pycnidia not seen.
he features of our sample do not fit any hitherto known European or North
American species (Ekman 1996, Llop 2007). Its characteristics appear closest
to B. trichosperma, as compared to the available information by Dodge (1953).
Some characters of thallus and apothecia resemble those of Bapalmuia Sérus.,
although the ascus structure and ascospores are completely different.
his African species is known elsewhere only from the Usambara Mountains
(Tanzania) in the west regions of the African continent (Dodge 1953).
Specimens examined - São Tiago: W of São Domingos, Rui Vaz, centre of village,
mixed trees and outcrops along small road, 23º 36.0 W, 15º 02.1’ N, 825 m, on Eucalyptus,
07.VII.2006, P. & B. v. d. Boom 36451 (hb. v. d. Boom).
Bactrospora thyrsodes nom. nov. (Cape Verde) ... 175
Bacidina pallidocarnea (Nyl.) Vězda
his taxon is pantropical and foliicolous, but can also be found in subtropical
and even wet-temperate areas (Lücking 2008). It has been found on the remains
of Agave leaves in Cape Verde, which are ecologically similar to twigs.
Specimens examined - São Vicente: Monte Verde, just below top of the mountain,
NW slope with acidic outcrops, shrubs and ±scattered small trees, 24º 56.0’W, 16º 52.2’
N, 700 m, on rotting leaf of Agave, 15.VII.2006, P. & B. v. d. Boom 36600 (hb. v. d.
Boom).
he next five species were misidentified as Bacidia but a careful examination
has shown that the specimens represent very different genera from Bacidia s. l.
In addition, most of them are new for the lichen flora of the Cape Verde Islands.
We first propose a new combination for Bacidia thyrsodes:
Bactrospora thyrsodes (Stirt.) Llop & van den Boom comb. nov.
MycoBank 513119
≡ Lecidea thyrsodes Stirt., J. Linn. Soc. London, Bot. 14: 368, 1874;
Bacidia thyrsodes (Stirt.) Zahlbr., Cat. Lich. Univ. 4: 245, 1926.
Type: CAPE VERDE. Sant Vincent. 1987 (holotype-bm !)
= Lecidea heterobola Cromb., J. Linn. Soc. London, Bot. 16: 214, 1877
= Bactrospora carneopallida Egea & Torrente, Lichenologist 25: 226, 1993.
Type: SPAIN. Islas Canarias: Lanzarote, Mirador del Rio, c. 400 m, rocas
volcánicas, 13 January 1990, J. M. Egea (holotype-mub, isotype-gzu)
Some samples identified as Bacidia thyrsodes or Bacidia cf. from B. Mies’
collection in M appear to be conspecific with Bactrospora carneopallida. he
study of the type material of Bacidia thyrsodes showed that the specimen was
also conspecific with Bactrospora carneopallida. Because the epithet thyrsodes
has priority over carneopallida, we propose the new combination Bactrospora
thyrsodes as the correct name for the taxon (McNeill et al. 2006: Art. 11.4).
his species was previously cited from Cape Verde by Egea & Torrente
(1993b) as Bactrospora carneopallida. Its distribution ranges from Macaronesia
to the European Atlantic coast (Paz-Bermúdez & López de Silanes 1998).
Specimens examined - Sal: Algudeiro, N Santa Maria, an Basaltblöcken, Substrat:
Fels, Basalt, 22’56º W, 16’37º N, 10 m, Expos. N, 17.X.1988, B. Mies 1040b, 1040c,
1040d1 (M-0142072, 0142080, 0142076). Santo Antão: Fontainhas, N-Küste, an
Basalt, Substrat: Fels, Basalt, 25’06º W, 17’12º N, 260 m, Expos. N, 21.X.1988, B. Mies
1191n (M-0142070). Ibid., S side above village, N exposed slope with acidic outcrops
along trail, 25’06º W, 17’11º N, 245 m, 20.VII.2006, P. & B. van den Boom 36811 (hb.
v.d. Boom). São Vicente: NW-Hang des Mt. Verde, Punta Antonio Gomes, an
Tublöcken, 50 m, 11.IX.1986, B. Mies 14d2 (M-0142069).
176 ... Llop & van den Boom
Buellia dispersa A. Massal.
his taxon has a disjunct distribution; it is known from the Mediterranean area
and drier inner alpine valleys (Scheidegger 1993), Canary Islands (Hafellner
1995), and southwest North America (Bungartz et al. 2002).
Specimens examined - Sal: Mt. Grande, S unterhalb der Spitze, an Tuffit, Substrat:
Fels, 22’54º W, 16’49º N, 370 m, Expos. E, 08.XI.1988, B. Mies 1059d (M-0142074,
0142079); ibid., NE-Hang, an Tuffit, Substrat: Fels, Tuffit, 22’54º W, 16’49º N, 350 m,
Expos. NE, 08.X.1988, B. Mies 1063g1 (M-0142078).
Cresponea flava (Vain.) Egea & Torrente
he examined specimens were misidentified as Bacidia thyrsodes, but the
type collection of the latter proves to belong to the related genus Bactrospora
A. Massal. (see above). Cresponea differs from Bactrospora in having
conglutinated and frequently anastomosed paraphysoids (Grube 1998).
Cresponea flava has a disjunct distribution in the Tropics, being known from
southeast Asia, South America, and both African coasts (Egea & Torrente
1993a).
Specimens examined - Brava: Vinagre, Bewässerunsgebiet der W-Küste,
Strassenbäume und Totholz, Substrat: Baum, 24’41º W, 14’52º N, 100 m, Expos. E,
10.XI.1988, A. Kalnins 1274c (M-0142077). São Nicolau: NW des Mt. Bissau, Ribeira
zum Rib. Madeira Vermelha, an Basalt des Flussbettrands, Substrat: Fels, Basalt, 24’15º
W, 16’37º N, 120 m, Expos. N, 11.XI.1988, B. Mies 1077d1 (M-0140271).
Fellhaneropsis vezdae (Coppins & P. James) Sérus. & Coppins
his is the first report of this taxon from the Cape Verde archipelago. Foliicolous
and corticolous collections were previously reported from Madeira (Sérusiaux
1996) and Topham & Walker (1982) found it on bark in the Canary Islands.
Specimens examined - São Vicente: Monte Verde, just below the top of the
mountain, NW slope with acidic outcrops, shrubs and ±scattered small trees, 24º 56.0’
W, 16º 52.2’ N, 700 m, on Casuarina, 15.II.2006, P. & B. v. d. Boom 36598 (hb. v. d.
Boom).
Toninia submexicana B. de Lesd.
his taxon was hitherto known only from North and South America (Timdal
1992). It differs from all previously reported species of Toninia in Cape Verde
by its grey epithecium reacting K + violet and ascospore morphology.
Specimens examined - Fogo: Cha das Caldeiras, N-Hang des Pico Novo, an
Phonolith und über Feinerde, Substrat: Felse, Phonolith, Erde, 24’21º W, 14’57º N, 2675
m, Expos. N, 02.XI.1988, B. Mies 1241f1 (M-0142073).
Bactrospora thyrsodes nom. nov. (Cape Verde) ... 177
Key to the species of Bacidia s. l. and allied species from Cape Verde
1a. Exciple carbonaceous, asci Arthonia-type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
1b. Exciple not carbonaceous, asci of different type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
2a. Paraphysoids branched, ascospores (30–)35–60(–65) × 3–4(–4.5) µm,
3–9 septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bactrospora thyrsodes
2b. Paraphysoids not branched, sometimes anastomosed, ascospores
15–22(–24) × (4–)4.5–5.5 µm, 3(–5) septate . . . . . . . . . . . . . . . . . .Cresponea flava
3a. Ascus Byssoloma-type, paraphyses branched and anastomosed,
exciple red brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fellhaneropsis vezdae
3b. Ascus type different, paraphyses not branched and anastomosed. . . . . . . . . . . . . . . .4
4a. Ascus Lecanora-type, exciple paraplectenchymatous, paraphyses with
a swollen apical cell, 4–5 µm thick . . . . . . . . . . . . . . . . . . Bacidina pallidocarnea
4b. Ascus Bacidia-type, exciple prosoplectenchymatous, paraphyses without
a swollen apical cell. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5a. Margin of exciple and upper hymenium olivaceous to blackish green,
K+ green and N+ violet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bacidia subincompta
5b. Margin of exciple and upper hymenium not green . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6a. Exciple and hypothecium reddish purple, K+ purple, ascospores
(30–)40–60(–75) × (2–)2.5–4 µm, (5–)7–15 septate . . . . . . . . . .Bacidia polychroa
6b. Exciple and hypothecium colourless to yellowish, ascospores smaller . . . . . . . . . . . .7
7a. Ascospores (20–)22–39 × 2–3(–3.5) µm, 3–7 septate, apothecia
orange to reddish, disc with a darker rim . . . . . . . . . . . . . . . . . . . Bacidia atlantica
7b. Ascospores 20–27 × 1–2 µm, 3–5 septate, apothecia
fleshy to yellow cream, evenly pigmented . . . . . . . . . . . . . . . .Bacidia trichosperma
Discussion
he genera Bacidia and Bacidina are represented by five species, all new for
the lichen flora of Cape Verde. he number of species representing Bacidia s.
l. varies considerably across the Macaronesian archipelagos: Canary Islands
16 species, Madeira 10 species and Azores 5 species (Hafellner 1995, 1999,
2002, 2005; van den Boom & Etayo 2006, Llop et al. 2007). Although Cape
Verde is sometimes not included in Macaronesia (Vanderpoorten et al. 2007),
despite the distance between the archipelagos, the number from Cape Verde is
similar to Azores, but the composition of species is completely different, even
though there are few coincidences with the other islands (Canary Islands and
Madeira).
Biogeographically, the species of Bacidia s. l. show three distributional
patterns. A Macaronesian-Mediterranean distribution is shown by Bacidia
polychroa, B. subincompta, Bacidina pallidocarnea, and Fellhaneropsis vezdae.
hese species grow on the Mediterranean pluviseasonal oceanic bioclimate belt.
178 ... Llop & van den Boom
his belt is occupied by laurisilva or bush communities that replace the forest
ater alteration (Duarte et al. 2005). Bacidia atlantica represents a Mid-Atlantic
endemism (Mies & Lösch 1995), as it occurs only in Ascension Island and Cape
Verde. he third pattern (as shown by Bacidia trichosperma) corresponds to a
tropical distribution, basically including mainland Africa. Other species from
the archipelago with a similar distribution include Heterodermia isidiophora
(Nyl.) D.D. Awasthi (Mies & Lösch 1995). Vanderpoorten et al. (2007) also
suggests an affinity of the Cape Verde cryptogamous flora with that of
continental Africa.
hose aforementioned species not belonging to Bacidia s. l. show a different
distribution pattern, except for Toninia submexicana. hey are growing on
the Mediterranean xeric or desert belt, showing a tropical to subtropical arid
distribution. In addition, these species are saxicolous. his pattern occurs
equally among bryophytes from the driest Macaronesian islands (GonzálezMancebo et al. 2008). Toninia submexicana has a disjunct distribution, as it
was known from North and South America (Timdal 1992). his species grows
above the Mediterranean pluviseasonal oceanic belt, which has a drier climate.
Acknowledgements
We thank the curators of BM and M for loans of specimens. A. Sánchez-Cuxart (BCN)
is thanked for his kind help with loans, and H. Sipman for providing information on
material from Cape Verde from B. he authors are grateful to reviewers B. J. Coppins
and H. J. M. Sipman, and S. Pennycook, whose comments and suggestions helped to
improve the manuscript.
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