Three overlooked species of Bacidia from insular Laurimacaronesia

Abstract

We discuss the taxonomy of three species of Bacidia occurring in insular Laurimacaronesia. Two of them, B. amylothelia (Vain.) Vain. and B. endoleucoides (Nyl.) Zahlbr., which were previously described from Angola and Madeira, respectively, are found here to belong in Bacidia s. str. (Ramalinaceae). Modern descriptions and illustrations are provided for the first time. Bacidia amylothelia is similar to B. areolata Gerasimova & A. Beck, B. campalea (Tuck.) S. Ekman & Kalb, B. fusconigrescens (Nyl.) Zahlbr., B. heteroloma (Vain.) Zahlbr., B. millegrana (Taylor) Zahlbr. and B. suffusa (Fr.) A. Schneid. and is reported here from the Canary Islands. Bacidia endoleucoides is most likely to be confused with B. absistens (Nyl.) Arnold, B. friesiana (Hepp) Körb., B. salazarensis B. de Lesd. and B. caesiovirens S. Ekman & Holien and was found to be widely distributed in the Canary Islands and Azores in addition to Madeira. The third species, Bacidia deludens S. Ekman, Tønsberg & van den Boom, is described here as new to science. Bacidia deludens is characterised by a greyish, crustose thallus with whitish soralia, pale apothecia with crystals in the hymenium and proper exciple, acicular ascospores with 3–19 septa, and the production of fumarprotocetraric acid as the consistently present major substance. It is described here from the Canary Islands and Madeira. Although conservatively treated here in Bacidia, we argue that it is likely to belong in the Malmideaceae. An identification key to all known species of Bacidia s. str. in insular Macaronesia is provided.

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NORDIC JOURNAL OF
BOTANY
Nordic Journal of Botany
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© 2021 e Authors. Nordic Journal of Botany published by John Wiley & Sons Ltd on behalf of
Nordic Society Oikos
is is an open access article under the terms of the Creative Commons
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Accepted 12 January 2021
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doi: 10.1111/njb.03055
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Published 12 March 2021
We discuss the taxonomy of three species of Bacidia occurring in insular
Laurimacaronesia. Two of them, B. amylothelia (Vain.) Vain. and B. endoleucoides
(Nyl.) Zahlbr., which were previously described from Angola and Madeira, respec-
tively, are found here to belong in Bacidia s. str. (Ramalinaceae). Modern descriptions
and illustrations are provided for the rst time. Bacidia amylothelia is similar to B.
areolata Gerasimova & A. Beck, B. campalea (Tuck.) S. Ekman & Kalb, B. fusconi-
grescens (Nyl.) Zahlbr., B. heteroloma (Vain.) Zahlbr., B. millegrana (Taylor) Zahlbr.
and B. suusa (Fr.) A. Schneid. and is reported here from the Canary Islands. Bacidia
endoleucoides is most likely to be confused with B. absistens (Nyl.) Arnold, B. friesiana
(Hepp) Körb., B. salazarensis B. de Lesd. and B. caesiovirens S. Ekman & Holien and
was found to be widely distributed in the Canary Islands and Azores in addition to
Madeira. e third species, Bacidia deludens S. Ekman, Tønsberg & van den Boom, is
described here as new to science. Bacidia deludens is characterised by a greyish, crustose
thallus with whitish soralia, pale apothecia with crystals in the hymenium and proper
exciple, acicular ascospores with 3–19 septa, and the production of fumarprotocetraric
acid as the consistently present major substance. It is described here from the Canary
Islands and Madeira. Although conservatively treated here in Bacidia, we argue that it
is likely to belong in the Malmideaceae. An identication key to all known species of
Bacidia s. str. in insular Macaronesia is provided.
Keywords: Bacidia, determination key, Malmideaceae, new species, Ramalinaceae
Introduction
During ongoing investigations of the lichen ora of Macaronesia (van den Boom and
Magain 2020 and references therein), the junior author came across three Bacidia-like
species. Further studies of that material as well as a number of types of previously pub-
lished names indicated that two of them belong in Bacidia s. str. (Ramalinaceae) and
correspond to the named but poorly understood species B. amylothelia (Vain.) Vain.
and B. endoleucoides (Nyl.) Zahlbr. We provide here the rst modern descriptions and
illustrations of these species and provide an identication key to all currently known
members of Bacidia s. str. in insular Laurimacaronesia. e third species posed more of
Three overlooked species of Bacidia from
insular Laurimacaronesia
StefanEkman, TorTønsberg and Pieter P. G.van den Boom
S. Ekman (https://orcid.org/0000-0003-3021-1821) ✉ (stefan.ekman@em.uu.se), Museum of Evolution, Uppsala Univ., Uppsala, Sweden. – T. Tønsberg,
Dept of Natural History, Univ. Museum of Bergen, Univ. of Bergen, Bergen, Norway. – P. P. G. van den Boom (https://orcid.org/0000-0002-1929-2088),
Arafura 16, Son, the Netherlands.
Research

2
an enigma, displaying supercial similarities with Bacidia s. str.
in the acicular ascospores (Ekman 1996, Kistenichetal. 2018).
However, it nally became clear from ascus studies that simi-
larities with the Ramalinaceae were only supercial and that
the third species represents an undescribed species that is likely
to belong in the Malmideaceae. It is described here but provi-
sionally placed in the genus Bacidia as B. deludens.
Material and methods
Microscopic characters were investigated either in a 10% aque-
ous solution of KOH (ascospores, paraphyses) or in pure water
(all other characters). e nomenclature and identication of
apothecial and pycnidial pigments follow Meyer and Printzen
(2000). Descriptions are based on the cited Macaronesian
material only. Colour reactions of pigments were observed in K
(a 10% aqueous solution of KOH), N (a 35% aqueous solution
of HNO3) and a 15% aqueous solution of HCl. Measurements
of quantitative characters are given either as ‘minimum value –
maximum value’ or ‘minimum value – arithmetic mean value
– maximum value (s = sample standard deviation, n = sample
size)’. Asci were stained with a 0.3% w/w aqueous solution of
IKI for the study of tholus structures. Lichen substances were
screened using high performance thin layer chromatography
(HPTLC) according to Arupetal. (1993) in system A (Bacidia
amylothelia, B. endoleucoides) or thin layer chromatography
(TLC) according to Orangeetal. (2010) in system A, B and C
(Bacidia deludens). Coordinates are provided as latitude-longi-
tude in the WGS84 reference system, either as decimal degrees
or degrees + decimal minutes.
Taxonomy
Bacidia amylothelia (Vain.) Vainio (1926, p. 18),
MB377918 (Fig. 1)
Basionym: Lecidea laurocerasi var. amylothelia Vainio (1901,
p. 420), MB607922.
Type: Angola, Cuanza Norte: Golunga Alta, hab. in truncos
juniores in sylvis prope Sange, undated, F. Welwitsch: Iter
Angolense 184 (BM 001096042–holotype, seen by SE).
Figure1. Bacidia amylothelia. (A) allus with apothecia. (B) part of hymenium showing asci with ascospores, (C) section through apothe-
cium in bright-eld showing pigmentation, (D) section through apothecium in polarized light showing minute crystals appearing as white
shine in proper exciple, (E) section through proper exciple, showing enlarged cell lumina of terminal excipular hyphae. (A–E): van den
Boom 45747. Scales: 0.5 mm (A), 25 µm (B), 50 µm (C–E).

3
Nomenclature
When Vainio (1901) described Lecidea laurocerasi var. amylo-
thelia, he cited the Welwitsch gathering Iter Angolense 184,
which may be represented by duplicates in other herbaria.
However, the preface by George Murray in Hiern (1896, p.
v–vi) makes it clear that the entire catalogue project dealing
with the Welwitsch collections, to which Vainio contributed
the lichens, is based on the specimens housed in BM. ere
is no type material in Vainio’s lichen herbarium at TUR.
Consequently, the name appears to be based on the single
specimen in BM, which we refer to as the holotype.
Description
allus crustose, thin to medium thick, almost white to
light greenish grey, ranging from discontinuous, of scat-
tered, ± convex and often slightly egurate areoles, to con-
tinuous, ± cracked, with warted surface. Prothallus lacking
or present, forming thin black lines in competition with
other lichens. Photobiont a member of Trebouxiophyceae,
unicellular, cells globose or short-ellipsoidal, 4.5–11.0 μm
long.
Apothecia biatorine, 0.4–0.6–1.0 mm diam. (s = 0.1,
n = 20), at rst at, becoming more or less convex with age.
Disc pale brownish yellow to beige to grey-brown, with ±
thin, white pruina. Margin concolorous with disc or slightly
paler or darker, distinct, slightly raised above disc in young
apothecia, soon level with the disc, persistent or becoming
excluded in convex apothecia, ± with white pruina, partic-
ularly close to the disc. Proper exciple 49–56–61 µm wide
(s = 5, n = 10), in lower part with numerous and diusely
distributed, short-bacilliform, tiny (ca 1 µm long) crystals
that are soluble in K, ± diusely brown-orange (K+ intensi-
fying) along edge and sometimes also in inner part closest to
hypothecium, otherwise ± unpigmented; excipular hyphae
dichotomously branched, in inner part of exciple with long
and narrow lumina (0.8–1.3 µm wide) and very thick and
gelatinized walls; terminal 5–6 cells with cell lumina gradu-
ally larger towards the edge, up to 8 µm wide. Hypothecium
± brown-orange (K+ intensifying). Hymenium 66–73–81
µm tall (s = 5, n = 10), colourless except for very pale orange
to pale brown-orange (K+ intensifying) epihymenium with
crystals (soluble in KOH). Paraphyses 1.2–1.7–2.0 µm wide
in mid-hymenium (s = 0.3, n = 20), unbranched or moder-
ately branched in upper part; apices ± clavate, 2.0–3.4–5.1
µm wide (s = 0.9, n = 20), without pigment. Asci clavate;
young spore mass forming a bluntly and broadly conical ocu-
lar chamber; tholus staining dark blue in I with a paler blue,
narrowly conical axial body, the zone closest to the axial body
concolorous with rest of the tholus. Ascospores 8 per ascus,
colourless, without perispore or ornamentation, acicular,
straight or shallowly helical, straight or somewhat coiled in
young asci, 39–47–57 µm long (s = 4, n = 20), 2.3–3.0–3.1
µm wide (s = 0.2, n = 20), 12.5–15.8–24.7 times as long
as wide (s = 2.5, n = 20), with 3.0–7.5–13.0 septa (s = 2.2,
n = 20).
Conidiomata not seen.
Chemistry and pigmentation
allus without acetone-soluble lichen substances or with
atranorin in trace amounts, K–, C–, KC–, PD–. Rubella-
orange in proper exciple, hypothecium and epihymenium.
Distribution and habitat
Bacidia amylothelia was described from northwestern Angola
by Vainio (1901) and is reported here as new to insular
Laurimacaronesia, where it is known from the two Canary
Islands Tenerife and La Palma. It has also been reported from
South Africa by Vainio (1926), but we were unable to locate
that material. As the species was originally discovered on
‘young trees’, it might turn out to be weedy and substantially
overlooked in at least the Old World tropics and subtrop-
ics. Two Tenerife collections are from the phorophyte Laurus
novocanariensis, growing on branches. One La Palma collec-
tion was found on a trunk of a small Laurus tree (9 cm diam.)
and one on a medium-sized trunk of Ocotea foetens.
Remarks
Bacidia amylothelia is a member of Bacidia in a strict sense
on account of the heavily gelatinized excipular cells with
long and narrow cell lumina (Kistenichet al. 2018). It can
be identied by its distinctly orange-brown hypothecium,
supercial white pruina, presence of minute crystals predom-
inantly in the lower part of the exciple near the edge, no radi-
ating clusters of large crystals in the proper exciple, and an
indistinctly delimited 5–6-layer zone of enlarged cell lumina
along the excipular edge. Apothecia in the Macaronesian
specimens are, however, on average somewhat paler than in
the type material, although apothecial colour in the holotype
varies from grey-brown to orange-brown to purplish black.
Similar and possibly closely related species include B. areolata
Gerasimova & A. Beck (colourless or pale yellow hypothe-
cium, no crystals or radiating crystal clusters in the proper
exciple, distinct 3–4-layer zone of enlarged cell lumina
along excipular edge; Gerasimovaetal. 2018), B. millegrana
(Taylor) Zahlbr. (colourless or pale yellow hypothecium, no
crystals or radiating crystal clusters in the proper exciple,
distinct 2-layer zone of enlarged cell lumina along excipu-
lar edge; Ekman 1996, p. 69), B. suusa (Fr.) A. Schneid.
(yellowish hypothecium, usually radiating crystal clusters in
the proper exciple, distinct 4–6-layer zone of enlarged cell
lumina along excipular edge; Ekman 1996, p. 108–109), B.
fusconigrescens (Nyl.) Zahlbr. (similar to B. suusa but with
minute crystals evenly distributed throughout proper exciple;
Ekman 1996, p. 69), B. campalea (Tuck.) S. Ekman & Kalb
(brown-orange to brown hypothecium, proper exciple with
large crystal clusters in lower part and with evenly dispersed
and minute crystals in upper part, cell lumina along excipular
edge as in B. amylothelia; Ekman 1996, p. 68–69), as well
as B. heteroloma (Vain.) Zahlbr., which is known only from
the Angolan type material and is characterized by the pale
brown hypothecium, evenly distributed minute crystals in
the proper exciple, and absence of enlarged cell lumina along
the excipular edge (based on observations in the syntypes BM
001107917 and BM 001107919 by SE).

4
Additional specimens examined
Canary Islands, Tenerife, Las Montanas de Anaga, SW of
Chamorga, E of Las Piedras, small open areas along trail
from start at road TF 123, through a small mirador to end
of trail, laurisilva with E- to N-exposed vertical outcrops,
28°33'63''N, 16°10'11''W, elev. 780 m a.s.l., 02 Mar 2011,
P. & B. van den Boom 45747, 45759 (herb. van den Boom).
La Palma, 3.5 km WSW of Los Sauces, Los Tilos, laurisilva,
narrow cleft with path along N facing volcanic outcrops,
between tunnel and mirador, 28°47'10''N, 17°48'60''W,
elev. 750 m a.s.l., 27 Oct 2012, P. & B. van den Boom 48327
(herb. van den Boom). La Palma, N of Santa Cruz, W of La
Galga, Cubo de La Galga, laurisilva in big valley with steep
shaded volcanic outcrops, 28°45'60''N, 17°47'00''W, elev.
650 m a.s.l., 31 Oct 2012, P. & B. van den Boom 48664
(herb. van den Boom).
Bacidia endoleucoides (Nyl.) Zahlbruckner (1926,
p. 193), MB378074 (Fig. 2)
Basionym: Lecidea endoleucoides Nyl. in von Krempelhuber
(1868, p. 234), MB390568.
Type: Portugal. Madeira, 1867, A. da Costa de Paiva s.n.
[= Barão de Castelo de Paiva] (H-NYL 17072–syntype, seen
by SE).
Figure2. Bacidia endoleucoides. (A) allus and apothecia in normally pigmented specimen, (B) allus with partially pigment-decient
apothecia, (C) Section through apothecium, (D) Ascospore. (A, C, D): van den Boom 48374; (B): van den Boom 48513. Scales: 0.5 mm
(A, B), 50 µm (C), 10 µm (D).

5
Nomenclature
Species described as new in von Krempelhuber (1868) are all
suxed ‘Nyl. spec. nov.’, and in the text there is a reference to
a letter from Nylander (either as ‘Nyl. in litt.’ or ‘Nyl. l. c.’).
We interpret this to mean that the descriptions were worded
by Nylander in this letter and used by von Krempelhuber.
erefore, we cite the basionym as ‘Nyl. in Kremp.’ as
opposed to ‘Nyl. ex Kremp’, which would have been cor-
rect if descriptions were worded by von Krempelhuber
(ICN Art. 46.5). Krempelhuber (1868) describes how the
relatively small Madeiran lichen collection by Costa de Paiva
was sent to Germany and made available to him. ere are
no duplicates or other gatherings of Lecidea endoleucoides in
M (where the Madeiran Costa de Paiva lichen collection is
housed) and the only specimen appears to have been sent to
Nylander. However, as Nylander was living in Paris at the
time, the specimen may have been divided and a duplicate
may be present in PC (which did not respond to a request for
material). erefore, we refer to the material in H-NYL as a
syntype for the time being.
Description
allus crustose, thin, almost white to light grey, ranging
from discontinuous, of scattered, convex areoles, to continu-
ous, cracked or areolate and with a warted surface. Prothallus
lacking. Photobiont a member of Trebouxiophyceae, unicel-
lular, cells globose or short-ellipsoidal, 4.5–9.5 μm long.
Apothecia biatorine, 0.3–0.4–0.7 mm diam. (s = 0.1,
n = 40), at rst at, becoming more or less convex with age.
Disc mostly bluish grey to bluish black, sometimes entirely or
partially paler, ± beige to purplish brown. Margin in upper
part concolorous with disc or paler, paler in lower part, dis-
tinct, slightly raised above disc in young apothecia, soon level
with disc, becoming excluded in convex apothecia. Proper
exciple 37–53–71 µm wide (s = 7, n = 20), without crystals,
blue-green (K+ intensifying) in uppermost part close to the
hymenium (except in pigment decient apothecia), ± brown
(K+ purplish) along the edge, colourless inside or with pale
brownish yellow pigment (K–) extending from the hypothe-
cium; excipular hyphae dichotomously branched, with long
and narrow lumina (0.6–1.0 µm wide) and very thick and
gelatinized walls; terminal 1–2 cells with cell lumina larger,
up to 5 µm wide. Hypothecium ± brownish yellow (K–) in
upper part, colourless below. Hymenium 68–76–85 µm tall
(s = 5, n = 20), colourless except for blue-green (K+ intensify-
ing) epihymenium without crystals. Paraphyses 0.9–1.3–1.6
µm wide in mid-hymenium (s = 0.2, n = 40), unbranched or
moderately branched in upper part; apices ± narrowly cla-
vate or not at all thickened, 1.2–2.7–5.4 µm wide (s = 0.9,
n = 40), sometimes with a diuse external hood of blue-green
pigment. Asci clavate; young spore mass forming a bluntly
and broadly conical ocular chamber; tholus staining dark
blue in I with a paler blue, narrowly conical axial body, the
zone closest to the axial body concolorous with rest of the
tholus or slightly darker. Ascospores 8 per ascus, colour-
less, without perispore or ornamentation, acicular, straight
or shallowly helical, straight or somewhat coiled in young
asci, 40–53–70 µm long (s = 6, n = 40), 2.3–2.8–3.6 µm
wide (s = 0.3, n = 40), 14.0–19.2–25.3 times as long as wide
(s = 2.5, n = 40), with 3–7.3–11 septa (s = 1.8, n = 40).
Conidiomata not seen.
Chemistry and pigmentation
allus without acetone-soluble lichen substances or with
atranorin in trace amounts, K–, C–, KC–, PD–. Bagliettoana-
green in epihymenium and uppermost part of proper exciple.
Laurocerasi-brown along excipular edge. Arceutina-yellow in
hypothecium.
Distribution and habitat
Bacidia endoleucoides was described from Madeira and has so
far never been reported from anywhere else (Carvalhoetal.
2008). It appears to be quite widespread in insular
Macaronesia, as we have seen specimens from the Canary
Islands (Tenerife and La Palma) and the Azores (São Jorge),
in addition to Madeira. In addition, we have recently come
across a few specimens from South Hampshire in southern-
most England, previously identied as B. friesiana, suggesting
that B. endoleucoides may have a wider distribution in coastal
Europe. In the Macaronesian sites studied by us, B. endo-
leucoides inhabits smooth as well as rough bark of trees and
shrubs. It seems to prefer light conditions in laurisilva as well
as habitats shaped by human activity (e.g. forest edges in the
cultural landscape). Known phorophytes include Apollonias
barbujana, Laurus novocanariensis, Ocotea foetens and Prunus
lusitancia.
Remarks
Bacidia endoleucoides is a member of Bacidia in a strict sense
on account of the heavily gelatinized excipular cells with long
and narrow cell lumina (Kistenichet al. 2018). Apart from
the original and quite vague diagnosis and brief mentions in
checklists (Tavares 1952, Hafellner 1992, 1995), this spe-
cies has not been discussed in the literature and no modern
description is available. Bacidia endoleucoides can be confused
with B. absistens (Nyl.) Arnold, B. friesiana (Hepp) Körb., B.
salazarensis B. de Lesd. and B. caesiovirens S. Ekman & Holien.
Forms of Bacidia absistens with a blue-green epihymenium
are similar to B. endoleucoides and are primarily distinguished
by the paler hypothecium and abundance of minute crystals
throughout the proper exciple (Ekman 1996, Coppins and
Aptroot 2009). Bacidia friesiana, unlike B. endoleucoides, has
a hyaline or pale straw hypothecium, wider and less gela-
tinized excipular hyphae, as well as a thallus that becomes
minutely granular (own observations, accurate descriptions
lacking, the one by Wirthet al. 2013 coming closest). We
have not come across any correctly identied material of
B. friesiana from Macaronesia. B. salazarensis, a widespread
tropical species, can be separated from B. endoleucoides by
the colourless or pale straw hypothecium, warmer red-brown
proper exciple, and most ascospores being 7–13 times as long
as wide (Ekman 2004). Bacidia caesiovirens, a European oce-
anic species not known from Macaronesia, can be recognized

6
by its pale yellowish hypothecium and granular thallus with
blue-green pigment (Ekman and Holien 1995). In addition,
there is a seemingly undescribed species of Bacidia occurring
in Macaronesia with which B. endoleucoides can be confused.
It is mentioned and briey characterized in the comments to
the identication key below (as B. a. salazarensis).
Additional specimens examined
Azores, São Jorge, NW of Velas, WNW of Rosais, trail to
Farol dos Rosais, near Cha do Areeiro, small forest with
Erica and Pittosporum and stones of walls, 38°44'82''N,
28°17'92''W, elev. 290 m a.s.l., 07 Sep 2017, P. & B. van den
Boom 57182 (herb. van den Boom). Madeira, Ribeiro Frio,
along Levada do Furado, 32°73'67.0''N, 16°88'59.6''W, elev.
600 m a.s.l., 22 Jan 1999, S. Ekman 3521 (UPS L-945335).
S of Ilha, 32°80'18.1''N, 16°91'41.3''W, elev. 600 m a.s.l.,
20 Jan 1999, S. Ekman 3485 (UPS L-945301). 0.9 km NW
of Ribeiro Frio, Balcões, 32°74'06''N, 16°89'19''W, elev.
890 m a.s.l., 10 Jan 2008, L. Tibell (UPS L-173341). Canary
Islands, Tenerife (NW), N of Santiago del Teide, Bco. de
Cuevas Megras o del Agu, path from Erjos to Los Silos, cen-
tral part, near the houses of Las Cuevas Negras, laurisilva,
with Erica arborea, Laurus novocanariensis and Apollonias
barbujana, outcrops and walls of stones, 28°20'53''N,
16°48'61''W, elev. 590 m a.s.l., 15 May 2007, P. & B. van
den Boom 37940 (herb. van den Boom). La Palma, 3.5 km
WSW of Los Sauces, Los Tilos, laurisilva, narrow cleft with
path along N facing volcanic outcrops, between tunnel and
mirador, 28°47'10''N, 17°48'60''W, elev. 750 m a.s.l., 27
Oct 2012, P. & B. van den Boom 48374, 48382 (herb. van
den Boom). La Palma, 3.5 km WSW of Los Sauces, Los
Tilos, laurisilva, steep trail from visitor centre to mirador de
las Barandas, 28°47'70''N, 17°47'0''W, elev. 750 m a.s.l.,
29 Oct 2012, P. & B. van den Boom 48513 (herb. van den
Boom). La Palma, N of Santa Cruz, W of La Galga, Cubo
de La Galga, laurisilva in big valley with shaded steep volca-
nic outcrops, 28°45'60''N, 17°47'00''W, elev. 650 m a.s.l.,
31 Oct 2012, P. & B. van den Boom 48570 (herb. van den
Boom). Great Britain, England, south Hampshire (V.C. 11),
Hants, Roydon, Mill Copse, 50°79'36.0''N, 01°54'62.9''W,
10 May 1998, N. A. Sanderson 206 (UPS L-984795);
New Forest, Mark Ash Wood, Pond Hill, 50°86'42.4''N,
01°65'36.7''W, 21 Jan 2005, N. A. Sanderson 850 (UPS
L-984796); New Forest, Busketts Wood, Great Stubby
Hat, 50°89'47.5''N, 01°56'72.0''W, 27 Sep 2016, N. A.
Sanderson 2207 (UPS L-984798).
Bacidia deludens S. Ekman, Tønsberg & van den
Boom sp. nov. (Fig. 3, 4)
MycoBank: MB836877
allus crustose, greyish, with whitish, circular to ellipsoi-
dal soralia, producing fumarprotocetraric acid as consistently
present major substance. Photobiont a Trebouxiophyceae, uni-
cellular, small. Apothecia biatorine, ± convex when mature,
pale with a margin that often becomes ± brown on the out-
side, internally with no or small amounts of pigment but with
crystals near the edges. Ascus mostly more or less Micarea-
type. Ascospores acicular, up to 90 µm long, with 3–19 septa.
Type: Spain, Tenerife, N of Santiago del Teide, 1.5 km WSW
of Erjos, path to Las Portelas, laurisilva, path in forest, rather
shaded, with mainly Laurus novocanariensis and Erica arborea,
on Erica arborea, 28°19'70''N, 16°48'70''W, elev. 1000 m a.s.l.,
8 May 2007, P. & B. van den Boom 37646 (UPS L-972106–
holotype, herb. van den Boom–isotype, BG–isotype).
Etymology
e epithet deludens means ‘deceiving’, the present active par-
ticiple of deludo, and refers to the fact that, morphologically,
B. deludens may be taken for a member of the Ramalinaceae
even though it is a member of another family.
Description
allus crustose, thin, yellow-grey to pale brown-grey, sore-
diate, composed of scattered, irregular, convex areoles that
coalesce to form a continuous and ± cracked crust with smooth
or rough surface, on furrowed bark forming small, indeter-
minate patches of areoles up to 0.08 mm diam. with rough
surface, on smooth bark usually well delimited, to 20 mm
Figure3. Bacidia deludens sp. nov. (A, B) Sorediate thallus with apo-
thecia. Note younger and paler apothecia with dark-pigmented
outer rim in (B) (A): van den Boom 37646 (holotype); (B): van den
Boom 47804. Scales: 0.5 mm (A, B).

7
diam., areolate (areoles up to 0.16 mm diam.) or more often
continuous, then cracked but with otherwise ± smooth sur-
face. Soralia ± tinged pale brownish in outer part, whitish in
the centre where soredia have been shed, on furrowed bark
eorescent, irregularly rounded or (following the ridges of
the bark) ellipsoidal, at to convex, to 0.8 mm when elon-
gate; on smooth bark bursting through the thallus, mostly
rounded and to 0.5 mm diam., at and level with the sur-
rounding thallus, often forming a ± vertical, discontinuous
rim along the edge of the soralia. Soredia globose to ellipsoi-
dal, 20–28–41 μm long (s = 7, n = 50). Prothallus, when pres-
ent, blackish, particularly prominent where several specimens
form a mosaic. Photobiont a member of Trebouxiophyceae,
unicellular, cells globose or short-ellipsoidal, 5–8 μm long.
Apothecia biatorine, 0.2–0.4–0.8 mm diam. (s = 0.2,
n = 30), at rst at, soon becoming more or less convex. Disc
pale yellow, pale pink or pale beige. Margin in upper part
concolorous with disc or paler or darker, particularly outer
part sometimes dark brown, level with disc in young apothe-
cia, becoming excluded in convex apothecia. Proper exciple
34–46–56 µm wide (s = 8, n = 15), with a layer of minute
crystals (≤ 1 µm long) along the edge (soluble in K, insoluble
in N), colourless to very pale orange (K–), sometimes with
brown-orange (K–) in diuse and ± wide zone along edge;
excipular hyphae dichotomously branched, ± radiating,
with long and narrow lumina (ca 1 µm wide) and very thick
and gelatinized walls; terminal cell lumina not expanded.
Hypothecium colourless. Hymenium 44–61–73 µm tall
(s = 9, n = 15), colourless or with pale yellowish (K–) epi-
hymenium containing a thick layer of minute crystals (same
as in proper exciple). Paraphyses 1.2–1.3–1.5 µm wide in
mid-hymenium (s = 0.1, n = 30), abundantly branched; api-
ces narrowly clavate or not at all thickened, 1.2–1.7–2.3 µm
wide (s = 0.3, n = 30), without pigment. Asci clavate; young
spore mass forming an indistinct ocular chamber; axial body
when stained with IKI (in some asci) narrowly conical and
not reaching all through d-layer, or (in most asci) cylindrical
and reaching all through d-layer, the zone closest to the axial
body darker than rest of tholus, forming a thick, dark tube
in asci with a cylindrical axial body (approximately Micarea-
type sensu Hafellner 1984). Ascospores 8 per ascus, colour-
less, without perispore or ornamentation, acicular, straight or
slightly curved or very shallowly helical, straight or somewhat
coiled in young asci, 31–51–90 µm long (s = 13, n = 30),
2.6–3.1–3.6 µm wide (s = 0.2, n = 30), 10.0–16.5–29.0
times as long as wide (s = 4.5, n = 30), with 3–11.3–19 septa
(s = 4.0, n = 30), not constricted at septa.
Conidiomata pycnidia, rare, globose, 30–50 µm diam.,
semi-immersed, unilocular, with blue-green (K+ intensify-
ing, N+ purple) pigment around ostiole. Conidia 1-celled,
long-ellipsoidal, 4–6 µm long and ca 1.2 µm wide, formed
terminally from cylindrical, 1.2–1.5 µm wide conidiophores.
Chemistry and pigmentation
allus with fumarprotocetraric acid (major), protocetraric
acid (trace) and ± two unidentied substances (faint traces),
K–, C–, KC–, PD+ bright orange-red. Apothecia without
lichen substances in any detectable amounts. Small amounts
of an unidentied, yellowish or pale orange (K–) pigment
(Arceutina-yellow or perhaps Rubella-orange) in minute
Figure4. Bacidia deludens sp. nov. (A) Section through apothecium. Note that the epihymenial layer that appears brown is actually a
dense, colourless and mostly opaque layer of crystals. (B) Ascus stained with 0.3% IKI in water (after pretreatment with 10% KOH).
Note darker tube-like structure in apex. (C–E) Ascospores. (A, C–E): van den Boom 45954; (B): van den Boom 47804. Scales: 50 µm
(A), 10 µm (B–E).

8
quantities in proper exciple, hypothecium and hymenium.
Bagliettoana-green in pycnidial wall around ostiole.
Distribution and habitat
Bacidia deludens is currently known from the Canary Islands
(Tenerife and La Gomera) as well as Madeira at altitudes
between 600 and 1200 m a.s.l. We have observed it on bark
of Erica, Vaccinium, an unidentied shrub in laurisilva and
on Cupressus in a mixed forest. Localities vary from disturbed
and poor in lichens to fairly species-rich. In the type local-
ity, where B. deludens inhabits bark of Erica, accompanying
microlichens include Byssoloma marginatum, Coenogonium
luteum, Endohyalina ericina, Fellhaneropsis vezdae, Jamesiella
anastomosans, Micarea alabastrites, M. doliiformis, M. pyc-
nidiophora, Porina coralloidea and Scoliciosporum pruinosum.
One specimen lacks apothecia, which opens the possibility
that B. deludens may occur as an overlooked ‘sterile, sorediate
crust’ in insular Laurimacaronesia.
Remarks
In a fertile state, B. deludens may remind of a member of
Bacidia, Bacidina or Toniniopsis on account of the com-
bination of acicular and transversely septate ascospores,
biatorine apothecia and the Trebouxiophyceae photobiont
(Kistenich et al. 2018). e prominent, whitish or pale
brownish soralia, combined with the somewhat Cliostomum-
like apothecia with sparse amounts of pigment and pres-
ence of minute crystals only along the edge of the proper
exciple is unprecedented in the Ramalinaceae, however. In
a sterile state, on the other hand, Bacidia deludens bears a
supercial resemblance to Biatora britannica Printzen et al.,
B. eorescens (Hedl.) Räsänen and Lecanora jamesii J. R.
Laundon, in which argopsin is dominant in the two rst and
atranorin, usnic acid and 2-O-methylsulphurellin is usually
present in the latter (Tønsberg 1992, Lumbschetal. 1995,
Printzenetal. 2001).
Bacidia deludens does not t in any known genus in the
Ramalinaceae. e ascus structure instead suggests ani-
ties to the Malmideaceae or Pilocarpaceae. Based on the
well developed and strongly gelatinized proper exciple with
radiating hyphae, lack of constrictions at the ascospore
septa and the abundance of crystals in the hymenium, we
suggest that B. deludens is a member of the Malmideaceae.
Among the genera currently classied in the Malmideaceae
(Spribilleetal. 2020, Wijayawardeneetal. 2020), Malmidea,
Sprucidea and Zhurbenkoa possess mainly non-septate asco-
spores and a brown hypothecium, Malmidea also having
halonate ascospores, Sprucidea producing sporodochia and
Zhurbenkoa having a parasitic life-style on other lichens
(Kalbet al. 2011, Cáceres etal. 2017, Flakus et al. 2019).
Puttea includes species with non-septate ascospores, min-
ute apothecia with a poorly developed proper exciple, and
(when pigmented) brown pigment caps on the terminal cells
of the excipular hyphae (Dillmanetal. 2012). Ascomata are
unknown in Cheiromycina and Savoronala, which are instead
recognised by their prominent sporodochia (Ertzetal. 2013,
Muggiaetal. 2017). Ascospores with up to 3 septa are known
in Crustospathula and Kalbionora, but Crustospathula is also
characterised by stalked soralia, Kalbionora by a brown hypo-
thecium, and both genera by a variety of lichen substances
not involving fumarprotocetraric acid (Kalb et al. 2012,
Sodamuk et al. 2017). All genera except Malmidea cur-
rently include 1–4 species, while Malmidea has 52 species
(Wijayawardeneetal. 2020). e circumscription and generic
classication of the Malmideaceae is poorly known and it has
been suggested that several additional taxa may belong in that
family, e.g. the genus Porpidinia as well as couple of species
formerly referred to Phyllopsora in the Ramalinaceae or the
large and distantly related Lecidea, the type of which belongs
in the Lecideaceae (Breuss and Lücking 2015, Kistenichetal.
2018, 2019, Paliceet al. 2018, Flakuset al. 2019). At the
moment, we see three options to classify our new species: 1) a
new genus in the Malmideaceae could have been erected. We
do not favour this option, partly because the genus to which
our species belongs may turn out to be already described
but misclassied in another family, partly because we would
prematurely create a monotypic genus based on fragmentary
data in a phylogenetically and taxonomically poorly known
family. In the latter case, typication of the genus may turn
out to be suboptimal if several species are later shown to
belong to the genus. 2) We could have provisionally recog-
nised the species in an already described genus classied in
the Malmideaceae. As outlined above, all currently recog-
nised genera are morphologically homogeneous and match
poorly with our species. e two genera Crustospathula and
Kalbionora may seem as the least bad alternatives because of
the presence of 3-septate ascospores, but our species deviates
substantially from these genera in characters outlined above.
Adding our species to any of the genera would extend the
morphological variation and make the genus problematic to
characterise, which is why we decided to avoid this option.
3) e third option, which we settled for, is to provision-
ally classify our new species in the genus Bacidia awaiting
a reasonable overview of the phylogeny and classication of
the Malmideaceae. Bacidia has historically been the home to
basically all crustose lichens with a chlorococcoid photobi-
ont, biatorine apothecia and ascospores with three or more
transverse but no longitudinal septa (Ekman 1996). Most
of those species are not congeneric with the type species
B. rosella (Pers.) De Not., but a more natural classication
was recently proposed (Kistenichet al. 2018). Provisionally
accepting another species not congeneric with the type in
an already heterogeneous assemblage is in line with recent
history and would minimally impact the endeavour to
achieve a monophyletic Bacidia. In addition, there is prec-
edent for provisionally treating species as members of his-
torically heterogeneous genera awaiting phylogenetic and
taxonomic clarication, recent examples being, e.g. Lecidea
coriacea Holien & Palice (Holienetal. 2016), Bacidia gullah-
geechee Lendemer (Lendemer 2018), Bacidia pruinata Fryday
(Fryday 2019) and Lecidea streveleri T. Sprib. (Spribilleetal.
2020). e latter was explicitly placed in the Malmideaceae,
yet described in Lecidea for the same reason we described our
species in Bacidia.

9
Additional specimens examined (paratypes)
Madeira, Ribeiro Frio, at the head of Levada do Furado,
32°73'52.5''N, 16°88'59.0''W, elev. 600 m a.s.l., 22 Jan 1999,
S. Ekman 3520 (UPS L-945318, sterile). NW of Funchal,
road (ER228) from Ribeira Brava to São Vicente, ca 1 km N of
Boca da Encumeada, trail PR22 ‘Vereda do Chao dos Louros’,
laurisilva, 32°45'50''N, 17°01'10''W, elev. 880 m a.s.l., 30 Apr
2012, P. & B. van den Boom 47804 (herb. van den Boom).
S of Santiago, S of Redondo, Pico das Pedras, picnic area at
the edge of a mixed forest, scattered trees, including Camellia
and conifer trees such as Cupressus, 32°46'66''N, 16°53'85''W,
elev. 880 m a.s.l., 09 Apr 2019, P. & B. van den Boom 58422
(herb. van den Boom). Canary Islands, Tenerife, N of Santiago
del Teide, 1.5 km WSW of Erjos, path to Las Portelas, lau-
risilva, path in forest, rather shaded, with mainly Laurus novo-
canariensis and Erica arborea, 28°19'70''N, 16°48'70''W, elev.
1000 m a.s.l., 08 May 2007, P. & B. van den Boom 37594,
37641 (herb. van den Boom). La Gomera, NE of Valle Gran
Rey, NE of Arure, Garajonay N. P., S of road TF-713, trail
from Montaña de los Mamantiales to Raso de Don Pedro, lau-
risilva, 28°08'88''N, 17°17'55''W, elev. 1185 m a.s.l., 31 Aug
2011, P. & B. van den Boom 45954 (herb. van den Boom).
A provisional key to Bacidia sensu stricto in insular
Laurimacaronesia
e circumscription of Bacidia s. str. used for this key follows
Kistenichetal. (2018), which means that we include members
of the Ramalinaceae with ‘acicular ascospores, pycnidia with
liform and curved conidia, and a proper exciple consisting
of furcate hyphae with very thin cell lumina and thick, heavily
gelatinized cell walls (terminal cells sometimes excepted)’. We
include species listed as members of Bacidia from the area by
Hafellner (1995, 1999, 2005, 2008), Carvalhoetal. (2008),
Aptroot et al. (2010), Hernández Padrón and Pérez-Vargas
(2010), Breuss (2018) and van den Boom and Alvarado (2019)
with the following exceptions: Bacidia acclinoides (Nyl.) Zahlbr,
B. avida (Hepp) Tav. and B. fritzei (Stein) Zahlbr. are highly
unlikely to belong in Bacidia s. str. according to their origi-
nal descriptions, although we have not seen the types of these
names. Bacidia albonigricans (Nyl.) Zahlbr. is a member of the
Arthoniales according to studies of type material (M0101870,
seen by SE). Bacidia auerswaldii (Stizenb.) Mig. belongs in
Scutula as S. eusa (Rabenh.) Kistenichetal. (Kistenichetal.
2018). Bacidia arnoldiana Körb., B. caligans (Nyl.) A.L. Sm., B.
delicata (Leight.) Coppins, B. egenula (Nyl.) Arnold, B. inun-
data (Fr.) Körb. and B. phacodes Körb. all belong in Bacidina;
Wirthetal. 2013). Bacidia friesiana (Hepp) Körb. is probably
incorrectly reported for Macaronesia, as all investigated speci-
mens have turned out to be misidentications of B. endoleucoi-
des or B. heterochroa. Bacidia incompta (Borrer) Anzi belongs
in Bellicidia (Kistenichetal. 2018). Bacidia propinqua (Hepp)
Arnold belongs in Bilimbia (Ekman 1996, although not con-
specic with Bilimbia sabuletorum (Schreb.) Arnold as stated
there). Bacidia scopulicola (Nyl.) A.L. Sm. probably represents
misidentications of B. sipmanii (Brandet al. 2009). Bacidia
subacerina Vain. is a synonym of B. laurocerasi (Delise ex Duby)
Zahlbr. (Ekman 1996). Bacidia subilludens (Harm.) Zahlbr. is
a member of Bactrospora according to studies of type material
(DUKE-169907, seen by SE). Bacidia bagliettoana (A. Massal.
& De Not.) Jatta and B. subincompta belong in Toniniopsis
(Kistenich et al. 2018). Bacidia trachona (Ach.) Lettau and
B. viridifarinosa Coppins & P. James belong in Aquacidia in
the Pilocarpaceae (Aptrootet al. 2018). e newly described
Bacidia deludens is included in the key even though it does not
belong to Bacidia (nor Ramalinaceae).
e species referred to in the key as Bacidia a. salazarensis
is most likely not that species but an undescribed one. Like
B. salazarensis, it contains red-brown pigment in the exciple
and green pigment in the epihymenium. However, whereas
true B. salazarensis has straight and quite stout ascospores like
B. heterochroa (Müll. Arg.) Zahlbr. (Ekman 2004), B. a.
salazarensis has long and helically twisted ascospores like B.
laurocerasi. In addition, it has modest amounts of crystals in
the exciple unlike B. salazarensis, B. heterochroa and B. lauroc-
erasi. It is known for sure only from two relatively small speci-
mens (UPS L-945301 and as an immixture in a specimen of
B. endoleucoides cited above, UPS L-945335).
For the use of this key, one should study thin sections of
the darkest apothecia available. To be able to observe spe-
cic hues, it is recommended to use a light microscope with
strong light (daylight temperature) and keep the condenser
aperture as open as possible (maximum 1/3 closed). In ordi-
nary light (bright-eld), crystals may be mistaken for brown
pigmentation. Crystals are best studied between two crossed
polarization lters.
1 allus with distinct, circular or ellipsoidal, whitish or
pale brownish soralia. Uppermost part of hymenium and
uppermost part of proper exciple with a continuous layer
of crystals……………………………………B. deludens
1 allus without soralia. Uppermost part of hymenium
and uppermost part of proper exciple mostly without a
continuous layer of crystals………………….…………2
2 Apothecial sections at least in K with green hues in epi-
hymenium and/or proper exciple of at least the darkest
apothecia………………………………………………3
2 Apothecial sections in K without green hues, unpigmented
or with orange to purplish brown pigmentation…………5
3 Epihymenium purple (or sometimes green) in water, turn-
ing green in K; proper exciple densely and evenly set with
minute crystals………………………………B. absistens
3 Epihymenium at least partly green in water; proper exciple
without or with modest amounts of crystals……………4
4 Epihymenium purely blue-green, intensifying in K; with-
out crystals in proper exciple……………B. endoleucoides
4 Epihymenium with a mixture of blue-green and red-brown
pigment (intensifying and turning purplish, respectively);
proper exciple with modest amounts of minute crystals…
……………………………….………B. a. salazarensis
5 Darkest apothecia orange, bronze-brown or pale brown
to orange-brown to red-brown to black; epihymenium,
hypothecium, and/or inner part of proper exciple with
yellow, orange or brown pigment………………………6

10
5 Darkest apothecia ± pink to pale pink-orange; epihy-
menium, hypothecium and inner part of proper exciple
unpigmented or with tiny amounts of yellowish pig-
ment (although sometimes with crystals that may appear
brownish in bright-eld microscopy)…………………14
6 Hypothecium brown-orange to dark brown, conspicu-
ously K+ purple-red…………………………B. polychroa
6 Hypothecium pale yellowish to dark brown (K–, K+ inten-
sifying or K+ purplish), never conspicuously K+ purple-
red…………………………………………………….7
7 Apothecia with thin white pruina and/or at least epihyme-
nium with crystals………………………………………8
7 Apothecia without pruina and epihymenium without
crystals…………………………………………………9
8 Hypothecium and interior of proper exciple evenly dark
brown, K–; habit similar to Lecidella elaeochroma………
……………………………………………B. canariensis
8 Hypothecium ± brown-orange, K+ intensifying; interior
of proper exciple mostly pale but often somewhat brown-
orange near hypothecium; habit otherwise………………
…………………………………………...B. amylothelia
9 Epihymenium in water with red-brown, K+ purplish pig-
ment in distinct layer…………………………………10
9 Epihymenium brown-yellow (K–), ± orange (K± intensi-
fying) or unpigmented………………………………..11
10 Red-brown pigment forming distinct hoods in the walls
of the paraphysis apices……………………B. heterochroa
10 Red-brown pigment present as irregular grains or dis-
solved in the gelatinous matrix surrounding the paraphy-
ses, not forming distinct hoods……………B. laurocerasi
11 Epihymenium with pigment in distinct layer, brown-yel-
low, K–……………………………………..B. arceutina
11 Epihymenium unpigmented or vaguely and diusely pig-
mented, pigment not in a distinct layer………………12
12 allus granular……………………………….B. rubella
12 allus smooth or ± warted continuous, never
granular………………………………………………13
13 Spores distinctly helically twisted; on bark (Erica)………
……………………………………………B. sigmospora
13 Spores straight or slightly curved; on rock…………
……………………………………………...B. sipmanii
14 Ascospores bacilliform to fusiform; epihymenium with-
out crystals………………………………B. paramedialis
14 Ascospores acicular; epihymenium with crystals……
………………………………………………………15
15 Ascospores > 65 µm long, with up to 15 septa; proper
exciple thick, laterally > 80 µm wide, composed of nar-
row, not distinctly radiating cell lumina with gelatinized
walls thicker than the lumina; epihymenial crystals form-
ing sharply delimited layer………………...B. rosella
15 Ascospores ≤ 60 µm, with up to 7 septa; proper exciple
thinner, laterally < 80 µm wide, in outer part composed
of stout, distinctly radiating cell lumina with gelatinized
walls thinner than the lumina; epihymenial crystal layer
diuse, extending downwards between paraphyses………
…………………………………….………B. thyrrenica
Acknowledgements – We thank herbarium BM, H, as well as DUKE,
HBG, LY, M and TUR for sending type material of candidate
names for species treated in this paper. We are grateful to Pawel
Czarnota and Maarten Brand for commenting on the new species.
Author contributions
Stefan Ekman: Conceptualization (supporting); Data cura-
tion (supporting); Formal analysis (lead); Investigation
(lead); Methodology (lead); Project administration (equal);
Resources (equal); Validation (equal); Visualization (lead);
Writing – original draft (lead); Writing – review and editing
(lead). Tor Tønsberg: Conceptualization (supporting); Data
curation (supporting); Formal analysis (lead); Investigation
(supporting); Methodology (supporting); Project admin-
istration (equal); Resources (equal); Validation (equal);
Visualization (supporting); Writing – original draft (support-
ing); Writing – review and editing (supporting). Pieter P.
G. van den Boom: Conceptualization (lead); Data curation
(lead); Formal analysis (supporting); Investigation (lead);
Methodology (supporting); Project administration (equal);
Resources (equal); Validation (equal); Visualization (support-
ing); Writing – original draft (supporting); Writing – review
and editing (supporting).
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