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Phytotaxa 191 (1): 066–080
www.mapress.com/phytotaxa/
Copyright © 2014 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
66 Accepted by Thorsten Lumbsch: 26 Nov. 2014; published: 30 Dec. 2014
http://dx.doi.org/10.11646/phytotaxa.191.1.4
Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0
The Bacidia coprodes group (Ramalinaceae, Lecanoromycetes, Ascomycota), with
special reference to the species in Europe and North America
STEFAN EKMAN1
1 Museum of Evolution, Uppsala University, Norbyvägen 16, SE-75236 Uppsala, Sweden. Email: stefan.ekman@em.uu.se
Abstract
This paper aims to clarify taxonomy and nomenclature of the lichen Bacidia coprodes and its closest relatives in Europe
and North America. Bacidia coprodes as treated in recent works was found to consist of three distinct species in Europe and
North America, the correct names of which are B. coprodes (s. str.), B. notarisiana, and B. granosa. In addition, a fourth
species, B. inornata (Nyl.) Blomb. & Forssell, was found to be closely related to this group of lichens and is also treated
here. The four species can be separated by apothecial and pycnidial pigmentation as well as conidium shape and size. They
are closely related to species currently treated in Bacidia, Toninia, or Arthrosporum. Six basionyms, viz. Bilimbia coprodes,
Patellaria salevensis, Bilimbia subtrachona, Lecidea chytrina, L. chytrina var. hormospora, and L. inornata, are lectotypi-
fied. Detailed descriptions and a determination key are provided.
Key words: Toninia, Bacidia trachona, Bacidia subcoprodes, Bacidia chrysocolla, Bacidia clavigera, Bilimbia intercedens,
Bacidia verecundula, Catillaria aphana, Fellhaneropsis.
Introduction
When Körber first described Bilimbia coprodes for a crustose lichen with black apothecia, brown hypothecium, green
epihymenium, and 3-septate ascospores (Arnold 1858), he also noted the remarkable similarity with B. notarisiana A.
Massal., which had been validly described a few years earlier (Massalongo 1855). In a later treatment, Körber provided
a more detailed morphological assessment of B. coprodes and clearly distinguished between this lichen and Biatora
trachona (Ach.) Körb. (originally described as Verrucaria trachona Ach.), another lichen with superficially similar
morphological characteristics (Körber 1860). Although B. trachona always seems to form conspicuous pycnidia, it
seldom produces apothecia. However, rare observations of apothecia (e.g., Körber 1855; Zwackh-Holzhausen 1883;
Arnold 1884) seem to have caused confusion and led Stizenberger (1868) to unite the two species under the name
Lecidea trachona (Ach.) Nyl. This collective treatment was followed by most authors for a very long period of
time, one notable exception being Lettau (1912), who listed them as separate species and also transferred them to
Bacidia in accordance with the artificial generic circumscription suggested by Zahlbruckner (1905). Bacidia coprodes
consequently fell into disuse as a mere synonym of B. trachona until Llop & Ekman (2007) pointed out the distinction
between them and that they even belong in different families, B. trachona in the Pilocarpaceae and B. coprodes in the
Ramalinaceae. Meanwhile, several additional names of lichens related to Bacidia coprodes were published, starting
with Patellaria salevensis (Müller Argoviensis 1862), soon synonymized with Lecidea trachona by Stizenberger
(1868), and ending with Bilimbia pammellii (Hedrick 1934).
Although Bacidia coprodes was recently resurrected and discussed by Llop & Ekman (2007), that treatment did
not address the full extent of the taxonomical challenges in the group and was fraught with nomenclatural shortcomings
that need attention. The Antarctic material was later investigated by Olech & Czarnota (2009), who recognized two
species, B. subcoprodes Olech & Czarnota and B. chrysocolla Olech et al. The purpose of this paper is to further clarify
taxonomy and nomenclature of the European and North American members of the Bacidia coprodes group and discuss
their relationships.
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 67
Material and Methods
This study is primarily based on material in the following herbaria: C, E, LD, MA, MSC, TSB, and UPS. In addition,
I have examined type material in FH-TUCK, G, H-NYL, L, M, MICH, MU, S, US, VER, W, and ZT. Microscopic
characters were investigated either in a 10% aqueous solution of KOH (ascospores, paraphyses) or in pure water (all
other characters). Colour reactions of pigments and crystal solubility were observed in K (a 10% aqueous solution
of KOH) and N (a 50% aqueous solution of HNO3). 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)’. Lichen substances were screened using High Performance Thin Layer Chromatography
according to Arup et al. (1993).
I follow the recommendations of McNeill (2014) concerning the indication of holotypes. When it remains unclear
whether a specimen should be considered a holotype or just one among several syntypes (the locations of which may
currently be unknown), and lectotypification on the available specimen seems premature, such a specimen is simply
referred to as a ‘type’.
Results
Bacidia coprodes as treated by Llop & Ekman (2007) was found to consist of three distinct species in Europe and
North America, the correct names of which are B. coprodes (Körb.) Lettau, B. notarisiana (A. Massal.) Zahlbr., and B.
granosa (Tuck.) Zahlbr. In addition, a fourth species, B. inornata (Nyl.) Blomb. & Forssell, was found to be closely
related to this group of lichens and is treated here. The pigmentation of the proper exciple, hypothecium, hymenium,
shape and size of ascospores, pigmentation of the pycnidial wall, and shape and size of the conidia were found to be
essential characters for distinguishing among species. Two different and omnipresent pigments seem to be present in
the apothecia and pycnidia of the B. coprodes group, one greenish (K+ intensifying, N+ purple with the deposition
of blue crystals) and one reddish brown (K+ purplish, N+ orange-red) pigment. In addition, an orange-brown (K+
intensifying, N-) pigment is (at least sometimes) present in the hypothecium of B. notarisiana and B. granosa. These
three pigments correspond, respectively, to Bagliettoana-green and Laurocerasi-brown of Meyer & Printzen (2000),
and Rubella-orange of Ekman (1996). The latter was included in Arceutina-yellow by Meyer & Printzen (2000).
Discussion
None of the four species treated here belong to Bacidia s. str. (the type of which is B. rosella (Pers.) De Not.), but
are instead closely related to other species currently treated in Bacidia (e.g., B. bagliettoana (A. Massal. & De Not.)
Jatta, B. illudens (Nyl.) H. Olivier, B. subincompta (Nyl.) Arnold), in Toninia (e.g. T. aromatica (Sm.) A. Massal., T.
coelestina (Anzi) Vězda, T. plumbina (Anzi) Hafellner & Timdal, T. verrucarioides (Nyl.) Timdal), or in Arthrosporum
(with its single species A. populorum A. Massal.) (Ekman 2001). A comprehensive phylogenetic investigation of the
Ramalinaceae is needed, however, before the currently used circumscriptions of Bacidia and Toninia (Timdal 1991,
Ekman 1996) can be finally abandoned.
Apart from the four species treated here, the Bacidia coprodes group also includes B. subcoprodes Olech &
Czarnota and B. chrysocolla Olech et al., both of which were described from the Antarctica (Olech & Czarnota 2009).
I did not investigate the single Antarctic specimen of ‘B. coprodes’ listed by Llop & Ekman (2007). I did, however,
examine three specimens listed as ‘Bacidia trachona’ by Øvstedal (1986) and ‘Bacidia sp. A’ by Øvstedal & Lewis
Smith (2001), which were found to belong to Bacidia subcoprodes Olech & Czarnota (UPS L-016062, L-016063,
and L-016064). The latter name is illegitimate, being a younger homonym of B. subcoprodes (de Lesd.) Zahlbr., a
combination based on Bilimbia subcoprodes de Lesd. in Crozals (1924). B. subcoprodes Olech & Czarnota differs
from the species treated here in having a finely granular thallus, apothecia that become convex and often tuberculate in
early stages of development, a thin proper exciple that is often poorly pigmented in the outer parts, a thin hypothecium
that is more or less homogeneously red-brown, and pycnidia that consistently produce filiform and curved conidia. B.
chrysocolla is said to differ from all species treated here in having more or less curved ascospores that are commonly
5-septate. I have not seen material of that species.
EKMAN
68 • Phytotaxa 191 (1) © 2014 Magnolia Press
Taxonomy
Bacidia coprodes (Körb.) Lettau (1912: 132) (Fig. 1, 4A–C)
Bilimbia coprodes Körber in Arnold (1858: 503). Lecidea trachona (Ach.) Nyl. var. coprodes (Körb.) Stizenberger
(1868: 60). Lecidea coprodes (Körb.) Vainio (1883: 12). Type: GERMANY. Bayern: “Ober Wasserzell” [= short for
“oberhalb Wasserzell”], 1857, F. Arnold 652 (lectotype, designated here: M 0157923!; isolectotypes: L 0938376!, UPS
L-649434!, UPS L-649435!).
Patellaria salevensis Müller Argoviensis (1862: 400). Gyalecta salevensis (Müll. Arg.) Olivier (1911: 192).
Bacidia salevensis (Müll. Arg.) Zahlbruckner (1926a: 145). Bilimbia salevensis (Müll. Arg.) M. Choisy (1953: 177).
Type: FRANCE. Rhône-Alpes: Haute-Savoie, ‘‘Längs des Baches oberhalb Les Châbles gegen den Fuss des Piton”, 6
June 1856, J. A. P. Hepp (lectotype, designated here: G 00053997!; isolectotype: BM 000731134).
Bilimbia subtrachona Arnold (1870: 122). Bacidia subtrachona (Arnold) Lettau (1912: 133). Type: AUSTRIA.
Tirol: “Dolomit des grossen Rettenstein bei Kitzbühel”, 2 August 1869, F. Arnold (lectotype, designated here: M
0156514!; isolectotype: M 0012391!).
Lecidea chytrina Stizenberger (1868: 56). Bilimbia chytrina (Stizenb.) Arnold (1869: 643). Bacidia chytrina
(Stizenb.) Zahlbruckner (1926a: 105). Type: GERMANY. Bayern: “Nebelhorn in Allgäu”, undated, H. Rehm (lectotype,
designated here: S L4898!).
Lecidea chytrina var. hormospora Stizenberger (1868: 57). Bilimbia hormospora (Stizenb.) Arnold (1884: 575).
Bacidia hormospora (Stizenb.) Lettau (1912: 132). Type: GERMANY. Bayern: “Kalkstein im Rosenthal bei Eichstätt”,
27 April 1866, F. Arnold (lectotype, designated here: ZT Myc-3656!).
Biatorina seposita Th. Fries (1867: 151). Bilimbia coprodes var. seposita (Th. Fr.) Th. Fries (1874: 385). Bacidia
coprodes var. seposita (Th. Fr.) Zahlbruckner (1926a: 108). Bacidia trachona var. seposita (Th. Fr.) H. Magnusson
(1945: 312). Type: NORWAY. Troms: “Tromsöe”, 17 June 1857, T. M. Fries (holotype: UPS L-138582!).
Nomenclature:—Both the name and the diagnosis of Bilimbia coprodes were ascribed to Körber by Arnold
(1858), the diagnosis being taken directly (with citation marks) from a letter from Körber to Arnold dated 28 Aug 1857.
Therefore, the correct author citation is Bilimbia coprodes Körber in Arnold (ICN Art. 46.2). The attempt to lectotypify
B. coprodes by Llop & Ekman (2007) was invalid, as the selected specimen (Arnold, Lich. exs. 333 in M) is not part
of the original material, being collected several years after the publication of the name. The lectotype was selected
here from among the duplicates of the gathering numbered “652” by Arnold, this number being explicitly cited in the
protologue (“oberhalb Wasserzell (652 !)”).
The original material of Biatorina seposita Th. Fr. consists of a single specimen in UPS, which is considered
here as the holotype (ICN Art. 9.N.1). This specimen also bears an annotation by Stizenberger, the wording of which
(translated into Swedish) was partially used by Fries in the discussion on relationships with other taxa. A second
specimen of B. coprodes (cited below) from the type locality of Biatorina seposita was collected by Fries in 1864 (i.e.,
prior to the publication of the name in 1867) and was annotated by him as “Bilimbia coprodes Körb. α”, bearing no
trace of having been associated with the name Biatorina seposita.
Thallus crustose, thin, pale green to grey-green to brownish green, finely areolate with warted areoles to more or
less granular (particularly when overgrowing bryophytes). Granules, when present, 0.06–0.25 mm diam. Prothallus
lacking or thin, arachnoid, white (between discrete granules).
Apothecia lecideine, 0.3–0.5–1.2 mm diam. (s=0.2, N=70), rounded to irregularly lobed, at first flat, becoming
more or less convex with age, rarely with thin whitish pruina on margin or disc. Disc dark purplish brown to black.
Margin concolorous with disc, often shiny, distinct, slightly raised above disc in young apothecia, soon level with disc,
becoming excluded in convex apothecia.
Proper exciple 47–78 μm thick, usually without crystals but pruinose apothecia sometimes with internal, radiating
clusters of small crystals that dissolve in N but not K, composed of radiating, mainly dichotomously branched hyphae
with thick gelatinous walls and cell lumina not or only slightly constricted at septa; cell lumina 7–12 × 1.5–3 μm in
inner part of exciple, gradually wider towards edge, terminal cells up to 5 μm wide; laterally red-brown to greenish
black, containing a mixture of a red-brown (K+ purplish, N+ orange-red) and a sordid green pigment (K+ intensifying
green, N+ purple with the deposition of blue crystals), rim sometimes paler than inner exciple. Hypothecium in upper
part dark red-brown to greenish black (with a red-brown, K+ purplish and N+ orange-red, and sometimes also a green,
K+ intensifying green and N+ purple pigment), concolorous with upper part of exciple, lower part of hymenium
gradually merging into paler medulla (visible at least in thin sections). Hymenium 50–58–64 μm tall (s=5, N=15),
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 69
FIGURE 1. Bacidia coprodes, thalli with apothecia. A, B. Sweden (UPS L-647807). C. Sweden (UPS L-103910).—Scales = 0.5 mm.
EKMAN
70 • Phytotaxa 191 (1) © 2014 Magnolia Press
colourless below, upper part with varying amounts of green pigment, sometimes as a distinct epihymenial layer,
sometimes diffusely and unevenly dispersed in upper part. Paraphyses abundant, 1.5–2.0–2.8 μm wide in mid-
hymenium (s=0.3, N=27), unbranched or sparingly branched in upper part, infrequently anastomosed; apices ± clavate
or some not thickened at all, 2.4–3.5–4.9 μm wide (s=0.7, N=48), with or without surrounding gelatinous cap with
green pigment. Asci clavate, 8-spored, Bacidia-type sensu Hafellner (1984). Ascospores colourless, without perispore
or ornamentation, straight or slightly curved, bacilliform or narrowly ellipsoid, sometimes slightly tapering towards
one end, with blunt ends, 10–13–20 μm long (s=2, N=40), 2.5–3.1–4.4 μm wide (s=0.4, N=39), with 3 (or exceptionally
with 5) septa at maturity.
Pycnidia often present, semi-immersed in thallus except for a dark brown to blackish ostiolar region, 50–100
μm diam., unilocular, with a brown (K+ purplish, N+ orange-red) pigment (but no green pigment) in upper part;
inside of cavity lined with conidiophores terminated by cylindrical conidiogenous cells, 4–8 × 1.5–2.5 μm. Conidia
acrogenously formed, usually (1) short-bacilliform, narrowly ellipsoid, or tear-shaped, straight, non-septate, 3–10 ×
1.5–2.8 μm, or sometimes (2) filiform, curved, non-septate, 6–20 × 0.8–1.0 μm.
Chemistry:—No lichen substances detected.
Habitat and distribution:—On limestone or other base-rich types of rock, usually in shade, such as crevices or
overhangs. Some mountain specimens (judging from associated species) seem to be collected in moist habitats in more
or less full sun. The species is known from highlands and mountains throughout much of Europe: Italy, Montenegro,
France, Austria, Germany, Scotland, Norway (incl. Bear Island in the Svalbard Archipelago), Sweden, and Finland.
Reports from Hungary, Romania, and northern Spain by Llop & Ekman (2007) have not been re-investigated.
Occurrences in central, western, and northern Europe span from high to fairly low elevations, whereas the few known
finds from southern Europe always seem to be at high elevations. In addition, there is a single known find from the
Black Hills of South Dakota in the United States. B. coprodes is likely to be overlooked in other parts of the northern
United States and Canada. Wirth et al. (2013) pointed out that the species has not been observed in Germany for over
a century.
Notes:—B. coprodes is characterized by the hypothecium being as dark as or darker than the surrounding
proper exciple (Fig. 4A–C). Pycnidia with the straight and wide type of condia are by far the most common. In some
specimens, however, pycnidia contain narrow, filiform and curved conidia. These specimens do not seem to have any
other distinguishing features, but rather represent a random sample from the variation within B. coprodes. Therefore,
they are treated here as part of the variation within B. coprodes.
Bacidia coprodes can be confused with B. notarisiana and B. granosa, both of which constantly have filiform
conidia and a hypothecium that is paler than the surrounding exciple. It can also be confused with poorly developed
forms of Toninia aromatica, which, however, tends to produce rounded, sometimes pruinose squamules with irregular
maculae.
There are a few deviating specimens that I have excluded from B. coprodes as treated above and which may
represent undescribed species. Two rock-inhabiting specimens from Ben Lawers in Scotland (E 00425707 and E
00425708) are very similar to B. coprodes in most respects, except that ascospores are longer (19–26 um) and frequently
5- or 7-septate at maturity. A specimen overgrowing dying cyanophilic lichens and collected in Härjedalen, Sweden
(UPS L-649189), is also similar to B. coprodes, except that ascospores are narrowly bacilliform (15–25 × 1.5–2.5 um)
and the proper exciple dominated by green pigment. Finally, a corticolous specimen from Toledo in Spain (MA-Lichen
7145) deviates from B. coprodes in having a microsquamulose thallus, apothecia with a tall hymenium (60–75 um) and
producing mainly 1-septate ascospores (but some 3-septate).
Selected specimens examined:—AUSTRIA. Niederösterreich: “In monte Sonntagsberg prope Rosenau”,
undated, P. P. Strasser in Krypt. exs. (Vindob.) 657 (LD, UPS L-649467). FINLAND. Tavastia australis: Tammela
par., “Mustiala”, 1870, A. Kullhem (UPS L-648011). GERMANY. Bayern: “Am Wege zum Bahnhof bei Eichstätt”,
November 1873, F. Arnold (LD). “In der Waldschlucht des Rosenthales bei Eichstätt”, 6 December 1866, F. Arnold
in Arnold: Lich. exs. 333 (LD, UPS L-168564). ITALY. Abruzzi: L’Aquila, “Gran Sasso, above Rifugio Duca degli
Abruzzi, 2200 m,” 9 August 1996, P. L. Nimis & M. Tretiach (TSB 24544). MONTENEGRO. Plav: “In the mountain
pass Čakor, W of Peć, alt. 1850 m”, 8 July 1979, S. Svane 4229-1 (C). NORWAY. Troms: “Tromsö, Flöjfjeldet”, 23
June 1864, T. M. Fries (UPS L-138583). SVALBARD AND JAN MAYEN. Bjørnøya, “Mt. Misery”, 24 July 1868, T.
M. Fries (UPS L-650772). SWEDEN. Västergötland: Österplana par., “Kinnekulle, nedanför Högkullen, åt söder”, 20
September 1859, F. Graewe (LD, UPS L-647788). Närke: Lerbäck par., “Ödeskärr”, 1869, P. J. Hellbom (UPS L-647778).
Östergötland: Väversunda par., “Omberg, Väfversunda”, 1873, P. G. E. Theorin (LD, UPS L-647807). Värmland:
Filipstad par., “Pajsberg vid Yngen”, 1867, J. G. Lagergren (UPS L-647811). Hälsingland: Skog par., Holmsveden,
Sagberg kalkbrott”, 12 August 1991, Å. Ågren 268 (UPS L-54892). Härjedalen: Tännäs par., “Funnesdalen”, 1867, P. J.
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 71
Hellbom (UPS L-171930). Jämtland: Kall par., “Skutån”, undated, S. Almquist (UPS L-647836). Åre par., “Rennberg”,
August 1873, S. Almquist (UPS L-171901). Lule lappmark: Gällivare par., “Vastenjaure, Puoltaketjejauratj”, 9 August
1965, G. Gilenstam 1218a (UPS L-103910). Lycksele lappmark: Tärna par., “Björkfors, Mortsbäcken”, 19 July 1924,
A. H. Magnusson 8413 (UPS L-171922). Torne lappmark: Jukkasjärvi par., ”Nuolja, 1000 m”, 2 August 1921, A. H.
Magnusson 6200a (UPS L-171926). UNITED KINGDOM. Angus (VC 90): “Caenlochan Glen (N side): valley of
Glasallt Burn”, 7 August 1989, B. J. Coppins 13370 & O. L. Gilbert (E 00425705). UNITED STATES OF AMERICA.
South Dakota: Lawrence Co., “Black Hills, near Bridalveil Falls in Spearfish Canyon (5 mi south of Spearfish), in
deeply shaded east-west gulch, 4300 ft”, 12 August 1960, C. M. Wetmore 9377A (LD, MSC 0097749).
Bacidia notarisiana (A. Massal.) Zahlbruckner (1926a: 131) (Fig. 2, 4E)
Bilimbia notarisiana A. Massalongo (1855: 46). Type: ITALY. Liguria: Genova, “Genova, nella salita di Mt Cucco”,
undated, F. Baglietto (holotype: VER!; isotype: ZT!).
Nomenclature:—The invalidly published herbarium name Coniangium notarisianum was coined by Francesco
Baglietto and appears on the label of the type material of Bilimbia notarisiana. The duplicate specimen in ZT was
probably sent directly to Stizenberger by Baglietto and bears no sign of having been studied by Massalongo. The only
specimen seen by Massalongo, in VER, is considered here as the holotype.
Thallus crustose, thin, pale green to grey-green, finely areolate with warted areoles. Prothallus lacking.
Apothecia lecideine, 0.2–0.3–0.5 mm diam. (s=0.1, N=50), rounded to irregularly lobate, at first flat, becoming
more or less convex with age, without pruina. Disc beige-brown to black, often piebald. Margin concolorous with disc
or darker than disc, often shiny, distinct, slightly raised above disc in young apothecia, soon level with disc, becoming
excluded in convex apothecia.
Proper exciple 34–60 μm thick, without crystals; composed of radiating, mainly dichotomously branched hyphae
with thick gelatinous walls and cell lumina not or only slightly constricted at the septa; cell lumina 10–17 × 1.5–2.5
μm in inner part of exciple, gradually wider towards edge, terminal cells up to 5 μm wide; dark red-brown or blackish
green, with a mixture of a red-brown (K+ purplish, N+ orange-red) and a sordid green pigment (K+ intensifying green,
N+ purple with the deposition of blue crystals), rim sometimes paler than inner exciple. Hypothecium pale to medium
orange-brown throughout (with a red-brown, K+ purplish and N+ orange-red pigment and, at least in paler versions, an
orange-brown, K+ intensifying, N- pigment), distinctly paler than and contrasting with exciple. Hymenium 47–55–62
μm tall (s=5, N=15), colourless below, upper part with varying amounts of green pigment, which is diffusely and
unevenly dispersed in upper part, often concentrated to vertical streaks. Paraphyses abundant, 1.2–1.5–1.7 μm wide
in mid-hymenium (s=0.1, N=30), unbranched or sparingly branched in upper part, infrequently anastomosed; apices
± clavate, 1.6–3.3–4.7 μm wide (s=0.9, N=30), with or without surrounding gelatinous cap with green pigment. Asci
clavate, 8-spored, Bacidia-type sensu Hafellner (1984). Ascospores colourless, without perispore or ornamentation,
straight or slightly curved, short-bacilliform, narrowly ellipsoid, or fusiform, sometimes slightly tapering towards one
end, with blunt or slightly pointed ends, 11–14–19 μm long (s=2, N=30), 2.6–3.3–3.9 μm wide (s=0.3, N=30), with 3
septa at maturity.
Pycnidia often present, semi-immersed in thallus except for a blackish ostiolar region, 30–80 μm diam., unilocular,
with a red-brown (K+ purplish, N+ orange-red) pigment (but no green pigment) in upper part; inside of cavity lined
with conidiophores terminated by cylindrical or narrowly ampulliform conidiogenous cells, 5–8 × 1.2–1.6 μm. Conidia
acrogenously formed, filiform, more or less curved, non-septate, 8–16 × c. 1.0 μm.
Chemistry:—No lichen substances detected.
Habitat and distribution:—Apparently on calcareous rock, sometimes in anthropogenic settings (e.g., cement
constructions). Currently known only from low or moderate elevations in northern Italy, but is likely to be more
widespread in the Mediterranean region.
Notes:—Very similar to B. granosa, differing from this in the warted thallus surface never becoming
microsquamulose, the consistent lack of a dark prothallus, and the lack of green pigment in the pycnidial wall. Both
species differ from B. coprodes in the hypothecium being distinctly paler than the surrounding proper exciple (Fig.
4E–F).
Additional specimens examined:—ITALY. Friuli-Venezia Giulia: Udine, “Alto Friuli presso Bueris, 250 m, su
vecchio muretto d’arenaria,” 10 November 1983, P. L. Nimis (TSB 3769). Liguria: Genova, “Quarto, ad maceriam,
herba tectam,” 20 November 1936, C. Sbarbaro (UPS L-160570). Genova, “Val Bisagno, S. Pantaleo”, June 1946, C.
Sbarbaro (UPS L-634654).
EKMAN
72 • Phytotaxa 191 (1) © 2014 Magnolia Press
FIGURE 2. Bacidia notarisiana, thalli with apothecia. A. Italy (UPS L-634654). B. Italy (TSB 3769).—Scales = 0.5 mm.
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 73
Bacidia granosa (Tuck.) Zahlbruckner (1926b: 203) (Fig. 3A, 4F)
Lecidea granosa Tuckerman (1862: 420). Toninia granosa (Tuck.) Bouly de Lesdain (1921: 68). Bilimbia granosa
(Tuck.) Fink (1935: 225). Type: UNITED STATES OF AMERICA. South Carolina: Berkeley Co., “Santee Canal, on
brick walls”, H. W. Ravenel 191 (lectotype, selected by Ekman 1996: FH-TUCK 00060062!).
Lecidea trachonopsis Nyl. in Nylander & Crombie (1883: 64). - Bacidia trachonopsis (Nyl.) Zahlbruckner (1926a:
157). Type: CHINA. Shanghai: “prope Shanghai”, 1862, A. C. Maingay (H-NYL 18967!)
Bilimbia pammellii Fink in Hedrick (1934: 156). Bacidia pammellii (Fink) Zahlbruckner (1939: 365). Type:
UNITED STATES OF AMERICA. Iowa: Boone Co., “The Ledges, sandstone”, 27 July 1903, B. Fink (lectotype,
selected by Ekman 1996: MU L1397; isolectotype: MICH 68325!).
Nomenclature:—Fink should be cited as the author of Bilimbia pammellii, as both the name and validating
description were ascribed to him (ICN Art. 46.2). The ascription of the validating description to Fink was made by
Hedrick (1933) with the statement “the descriptions were prepared by Dr. Fink”. This is considered here as ‘internal
evidence’ in the sense of Art 46.7, as the two papers (Hedrick 1933, 1934) have the same title and author and are parts
of the same publication in the sense of Art 35.5.
Thallus crustose, thin to rather thick, pale green to grey-green, finely areolate; areoles ± effigurate with warted surface
or becoming ± microsquamulose with age. Prothallus lacking or sometimes present, forming a thin, dark line along
edge of thallus.
Apothecia lecideine, 0.2–0.3–0.6 mm diam. (s=0.1, N=75), rounded to irregularly lobate, at first flat, becoming
more or less convex with age, without pruina. Disc black. Margin concolorous with disc, distinct, slightly raised above
disc in young apothecia, soon level with disc, becoming excluded in convex apothecia.
Proper exciple 37–56 μm thick, without crystals; composed of radiating, mainly dichotomously branched hyphae
with thick gelatinous walls and cell lumina not or only slightly constricted at septa; cell lumina 7–12 × 1.5–3 μm in
inner part of exciple, gradually wider towards edge, terminal cells up to 5 μm wide; dark red-brown or blackish green,
with a mixture of a red-brown (K+ purplish, N+ orange-red) and a sordid green pigment (K+ intensifying green, N+
purple with the deposition of blue crystals), rim sometimes paler than inner exciple. Hypothecium pale orange-brown
to medium red- or purple-brown throughout (with a red-brown, K+ purplish and N+ orange-red pigment and, at least
in paler versions, an orange-brown, K+ intensifying, N- pigment), distinctly paler than and contrasting with exciple.
Hymenium 44–54–67 μm tall (s=6, N=32), colourless below, upper part with unevenly distributed green pigment,
which is often concentrated to vertical streaks. Paraphyses abundant, 1.2–1.5–2.0 μm wide in mid-hymenium (s=0.2,
N=55), unbranched or sparingly branched in upper part, infrequently anastomosed; apices ± clavate, 2.0–3.2–5.4 μm
wide (s=0.9, N=70), with or without surrounding gelatinous cap with green pigment. Asci clavate, 8-spored, Bacidia-
type sensu Hafellner (1984). Ascospores colourless, without perispore or ornamentation, straight or slightly curved,
short-bacilliform, narrowly ellipsoid, or fusiform, sometimes slightly tapering towards one end, with blunt or slightly
pointed ends, 11–15–28 μm long (s=3, N=72), 2.6–3.3–4.2 μm wide (s=0.3, N=72), with 3 septa at maturity.
Pycnidia often present, semi-immersed in thallus except for a blackish ostiolar region, 30–80 μm diam., unilocular,
with a mixture of a red-brown (K+ purplish, N+ orange-red) and a sordid green (K+ intensifying, N+ purple) pigment in
upper part; inside of cavity lined with conidiophores terminated by cylindrical or narrowly ampulliform conidiogenous
cells, 5–8 × 1.0–2.0 μm. Conidia acrogenously formed, filiform, more or less curved, non-septate, 7–20 × 0.5–1.0 μm.
Chemistry:—No lichen substances detected.
Habitat and distribution:—On sandstone or limestone rocks, sometimes on man-made substrates such as brick,
probably mostly or always in shade, at low elevations. Currently known from the continental eastern United States,
Cuba, Bermuda, and coastal eastern China.
Notes:—Very similar to B. notarisiana (see that species).
Additional specimens examined:—UNITED STATES OF AMERICA. Iowa: Clayton Co., “Bixby, north of
Edgewood”, 8 September 1961, H. A. Imshaug 28034A (MSC 0097748). Fayette Co., 3 December 1895, B. Fink in
Cummings, Williams & Seymour: Decades of North American Lichens 245 and Cummings, Williams & Seymour:
Lichenes Boreali-Americani 186 (LD, MSC 0143353, MSC 0132953); 1895, B. Fink (UPS L-649436, L-649437).
Fayette Co., “Echo Valley State Park, east of West Union”, 30 October 1965, C. Wetmore 13629 (MSC 0097750).
Jackson Co., “Maquoketa Caves Park, northwest of Maquoketa”, 9 September 1961, H. A. Imshaug 28087 (MSC
0097751). Jones Co., “Wapsipinicon, south of Anamosa”, 9 September 1961, H. A. Imshaug 28116 (MSC 0097747).
Louisiana: Orleans par., “graveyard in New Orleans”, 1851, J. Hale (FH-TUCK 00060061). Michigan: Alpena Co.,
“limestone sinkholes c. 3 mi N of Long Rapids”, 3 August 1974, R. C. Harris 9347 (MSC 0097752). Missouri: Taney
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74 • Phytotaxa 191 (1) © 2014 Magnolia Press
FIGURE 3. Bacidia granosa and B. inornata, thalli with apothecia. A. B. granosa, U.S.A. (MSC 0097751). B. B. inornata, Sweden (UPS
L-649188). C. B. inornata, Sweden (UPS L-637820).—Scales = 0.5 mm.
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 75
Co., “Mark Twain National Forest, Hercules Glades, 1 mile north of Persimmon Hollow”, 30 August 1991, C. M.
Wetmore 68454 (LD). South Carolina: undated, H. W. Ravenel (FH-TUCK 00060063). Charleston Co., “Charleston,
brick wall”, 1853, H. W. Ravenel 385 (FH-TUCK 00060064). BERMUDA. “The Flatts, on limestone”, April 1916, A.
B. Hervey (MSC 0097745). CUBA. Wright: Lichenes Cubae 236 (UPS L-662824).
Bacidia inornata (Nyl.) Blomb. & Forssell in Nordstedt et al. (1880: 81) (Fig. 3B–C, 4D)
Lecidea inornata Nylander (1874: 11). Type: FINLAND. Tavastia australis: “Padasjoki, Nyystölä, kiven reunalla”,
1872, E. Lang a.k.a. E. A. Vainio (lectotype, designated here: TUR-V 20450!; isolectotype: H-NYL 17944).
Nomenclature:—Nylander clearly saw TUR-V 20450, as he mentions in the diagnosis that the specimen was
growing “cum Verrucaria gibbosula”, a note that only appears on the material in TUR-V. The small piece in H-NYL
(17944) probably represents a part of Vainio’s original collection that Nylander kept for his own herbarium. As the
TUR-V specimen is more abundant, it is selected here as the lectotype.
Thallus crustose thin, pale green to grey-green, almost granular or finely areolate, with ± smooth, warted, or scurfy
surface. Prothallus thin, white, arachnoid, visible along edge of thallus and between discrete areoles.
Apothecia lecideine, 0.2–0.3–0.7 mm diam. (s=0.1, N=30), rounded or irregularly lobed, flat, somewhat convex
with age, without pruina, often becoming tuberculate by regeneration of several young apothecia from aging apothecia.
Disc yellow-brown to red-brown to black, sometimes piebald. Margin darker than or concolorous with disc, matt,
distinct, slightly raised above disc in young apothecia, soon level with disc, persistent.
Proper exciple 44–72 μm thick, without crystals; composed of radiating, mainly dichotomously branched hyphae
with thick gelatinous walls and cell lumina not or only slightly constricted at septa; cell lumina 7–12 × 2–2.5 μm in
inner part of exciple, gradually wider towards edge, terminal cells up to 8 μm wide; blackish purple-brown or blackish
green, with a mixture of a red-brown (K+ purplish, N+ orange-red) and,in the upper lateral parts, a sordid green
pigment (K+ intensifying green, N+ purple with the deposition of blue crystals), rim concolourous with inner exciple.
Hypothecium in upper part dark red-brown, concolourous with proper exciple or slightly paler, in lower part medium
to dark red-brown, concolourous with or paler than upper part of hypothecium (all parts with a red-brown, K+ purplish
and N+ orange-red pigment). Hymenium 32–39–46 μm tall (s=4, N=15), colourless or dilute red-brown, upper part
also with unevenly distributed spots of red-brown and sordid green pigment. Paraphyses abundant, 1.2–1.4–1.6 μm
wide in mid-hymenium (s=0.2, N=15), unbranched or sparingly branched in upper part, not anastomosed; apices ±
clavate, 2.0–2.9–3.9 μm wide (s=0.6, N=30), with or without surrounding gelatinous cap with red-brown or green
pigment. Asci clavate, 8-spored, Bacidia-type sensu Hafellner (1984). Ascospores colourless, without perispore or
ornamentation, straight, bacilliform or short-acicular with blunt ends, 11–15–19 μm long (s=2, N=30), 1.9–2.1–2.3 μm
wide (s=0.1, N=30), with (0–)3(–5) septa at maturity, septa apparently forming late during ascospore development.
Pycnidia often present, sessile to stipitate, 60–90 μm diam., unilocular; wall with a red-brown (K+ purplish, N+
orange-red) pigment; inside of cavity lined with conidiophores terminated by cylindrical or narrowly ampulliform
conidiogenous cells, 6–8 × 1.0–1.2 μm. Conidia acrogenously formed, bacilliform to narrowly ellipsoid, straight, non-
septate, 4–7 × 1.2–1.5 μm.
Chemistry:—No lichen substances detected.
Habitat and distribution:—B. inornata inhabits acid, preferably soft and easily weathering rocks and boulders
in humid and shady situations, often near water. Currently known from a few places in Sweden, Finland, Wales, and
Scotland.
Notes:—B. inornata is similar to B. coprodes in the pigmentation of the proper exciple and hypothecium as well as
conidium shape. B. inornata can be distinguished, however, on account of the tuberculate apothecia (Fig. 3C), thinner
hymenium, more red-brown and less green pigment in the epihymenium (Fig. 4D), narrower ascospores, and (when
present) sessile to stipitate pycnidia. Ascospores are poorly developed in the type material, which is probably why
Nylander (1874) incorrectly described the ascospores as acicular and 32–36 × 1–1.5 μm. In the otherwise excellent
description of this species by Vainio (1922: 166), ascospore shape and size were taken unaltered from Nylander’s
original description (“secund. Nyl. l. c.”), possibly because Vainio also had problems finding mature ascospores.
Symptomatically, the hand-written notes by Vainio on the envelope of the TUR-V specimen contain details about
almost every aspect of the morphology of the species with the exception of ascospores. Apart from the descriptions
by Nylander and Vainio, the only other description of this poorly known species was provided by Foucard (1990: 83).
First reported from Sweden (Västergötland) by Magnusson (1955), but that specimen is Micarea marginata (UPS L-
200622, determined by B. J. Coppins).
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76 • Phytotaxa 191 (1) © 2014 Magnolia Press
FIGURE 4. Sections of apothecia. A. Bacidia coprodes, Sweden (UPS L-647836). B. B. coprodes, Germany (ZT Myc-3656, lectotype
of Lecidea chytrina var. hormospora). C. B. coprodes, Norway (UPS L-138582, holotype of Biatorina seposita). D. B. inornata, U.K. (E
00425709). E. B. notarisiana, Italy (UPS L-634654). F. B. granosa, U.S.A. (UPS L-649436).—Scales = 100 μm.
RAMALINACEAE, LECANOROMYCETES, ASCOMYCOTA Phytotaxa 191 (1) © 2014 Magnolia Press • 77
Additional specimens examined:—SWEDEN. Härjedalen: Tännäs par., “1.2 km N of Ramundbergsvallen, the W side
of the stream Sveån”, elev. 825 m, 62°43’N 12°22’E, 3 June 2007, M. Svensson 1030 (UPS L-649188). Jämtland: Åre
par., “Handöl, Handöl rapids, E river bank S of the suspension bridge and N of the small hill with boulders c. 400
m SSW of the suspension bridge”, elev. 570–620 m, 63.23°N 12.45°E, 31 July 1993, A. Nordin, B. Owe-Larsson &
G. Thor 136 (UPS L-637820). Ångermanland: Nordingrå par., “Omne, p. 255, NO-sluttning, vittrad sten på hylla i
skugga”, 15 July 1958, S. Sundell 1474 (UPS L-200623). UNITED KINGDOM. Camarthen (VC 44): “13 km N of
Llandovery: Allt Rhyd-y-gross NNR, wood SE of Cribyn Du, on sheltered underhang on rock face by stream”, 24 July
1981, B. J. Coppins 8597 & R. G. Woods (E 00425712). Mid-Perth (VC 88): “Aberfeldy, Moness Wood, on shaded
rocks under trees, by river”, 6 April 1986, B. J. Coppins 11270 (E 00425710). “Crieff, Drummond Park, Drummond
Wood, shaded boulder near stream”, 9 August 1978, B. J. Coppins 3617 (E 00425711). “Allt na Ceardaich, Killin, base
of shaded acid-stone wall, 175 m”, 25 January 1991, A. M. Fryday 2024 (E 00425709).
Key
The key below includes saxicolous, lichenized members of the Ramalinaceae with brown pigment in the proper exciple
and hypothecium, a green (K+ intensifying, N+ purple) pigment in the epihymenium, and 3-septate, bacilliform,
ellipsoid, or fusiform ascospores shorter than 30 μm. All taxa keyed out below have an ascus apex with a more or less
conical ocular chamber and likewise conical axial body (more or less ‘Bacidia-type’ in the sense of Hafellner 1984).
The part of the key dealing with species of Toninia has been adapted from Timdal (1991). Note that Hitch et al. (2009),
unlike Timdal (1991), treat T. fusispora as a species separate from T. aromatica, mainly on account of ascospore
shape.
In addition to the species keyed out below, attention should be paid to other similar species with more or less
brown proper exciple and hypothecium and green epihymenium, and septate ascospores, viz. Bacidia bagliettoana (A.
Massal. & De Not.) Jatta, Bacidia illudens (Nyl.) H. Olivier, Toninia coelestina (Anzi) Vězda, and T. cretica Timdal.
All of these species inhabit soil, detritus, or moribund bryophytes, and never occur directly on rock. Ascospores mostly
tend to be longer than in the species treated here, often longer than 25 μm. Furthermore, 5–7-septate ascospores are
frequent in B. bagliettoana, B. illudens, and T. coelestina. Bilimbia sabuletorum (Schreb.) Arnold and B. microcarpa
(Th. Fr.) Th. Fr. are also frequently misidentified as members of the Bacidia coprodes group. Bilimbia, however, is
characterized by stout paraphyses and excipular hyphae and distinctly thick-walled asci. B. sabuletorum and B. lobulata,
the most widespread species of that genus, are also characterized by a finely warted epispore. Finally, members of
the B. coprodes group may be confused with an undescribed species of Fellhaneropsis currently known from shaded
siliceous rocks in a few sites in central Sweden and one in central Germany. This species also has black apothecia with
a well developed and persistent proper margin, 3-septate ascospores and a dark brown hypothecium. Unlike members
of the B. coprodes group, however, this species has abundant pycnidia with very long and filiform conidia, a dark-
amyloid tube structure in the axial body, somewhat constricted ascospore septa, and a micareoid photobiont.
1. Thallus thin, continuous and finely areolate, of discontiguous areoles with warted surface, or granular (granules then up to 0.25
mm wide) ................................................................ ...........................................................................................................................2
- Thallus composed of discrete or contiguous, up to 4 mm wide and flat to moderately convex, pale grey to dark brown squamules
with pruinose or epruinose surface ............................ ........................................................................................................................6
2. Ascospores broadly ellipsoid, at least 5 μm wide ......................................... ...................................................................T. mesoidea
- Ascospores less than 4.5 μm wide (members of the Bacidia coprodes group) ................. ................................................................3
3. Hymenium up to c. 45 μm, colourless to dilute red-brown; epihymenium with mixed red-brown and green pigment; ascospores
mostly c. 2 μm wide; pycnidia sessile or stalked; conidia bacilliform to narrowly ellipsoid, straight ............................. B. inornata
- Hymenium mostly taller than 45 μm, colourless and with green pigment in epihymenium and sometimes in vertical streaks; as-
cospores ≥ 2.5 μm wide; pycnidia immersed; conidia filiform, bacilliform, narrowly ellipsoid, or tear-shaped, straight or curved
........................ ....................................................................................................................................................................................4
4. Hypothecium at least in upper part dark red-brown to greenish black, concolorous with or darker than inner exciple; conidia either
filiform and curved or short-bacilliform to narrowly ellipsoid to tear-shaped, straight ...................................................B. coprodes
- Hypothecium evenly pale orange-brown to medium red-brown, paler than and distinctly contrasting with red-brown to blackish
green inner exciple; conidia filiform, curved......... ............................................................................................................................5
5. Areoles somewhat effigurate, with age forming minute squamules; prothallus, when present, forming a black line; pycnidial wall
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78 • Phytotaxa 191 (1) © 2014 Magnolia Press
with mixture of green and red-brown pigment .................................................................................................................. B. granosa
- Areoles warted, never becoming effigurate or microsquamulose; prothallus absent; pycnidial wall with red-brown pigment only
.................................................................. .....................................................................................................................B. notarisiana
6. Squamules with irregular pale flecks; upper part of hymenium with green pigment only ..............T. aromatica (incl. T. fusispora)
- Squamules without irregular pale flecks; upper part of hymenium dominated by red-brown pigment .. ................ T. verrucarioides
Additional names
The following observations were made on types of three names that I came across during the course of this revision.
Bilimbia clavigera Zahlbruckner (1903: 180)
Bacidia clavigera (Zahlbr.) Zahlbruckner (1926a: 107). Type: CROATIA. Split-Dalmatia: Hvar, “Insel Lesina, am
Wege von Lesina nach Cittavecchia, an Kalkfelsen”, November 1902, J. Lütkemüller (W!).
Zahlbruckner (1903) considered his new species to be closely related to Bacidia coprodes. The type material in W,
from the island Hvar off the Dalmatian coast in Croatia, is probably Toninia aromatica with a poorly developed thallus
consisting of scattered small whitish squamules.
Bilimbia trachona var. intercedens Arnold (1873: 505)
Bilimbia intercedens (Arnold) Arnold (1877: 570). Bacidia trachona var. intercedens (Arnold) Zahlbruckner (1926b:
209). Type: AUSTRIA. Tirol: “Larix-Rinde, Waldrast in Tirol”, August 1872, F. Arnold (UPS L-664328!).
Original material could not be located in M. The type specimen in UPS was probably sent from Arnold to T.
M. Fries. This specimen was labelled “Bilimbia --/n. sp” and annotated “a me adhuc denominanda” (= “should still
be named by me”) by Arnold, and surely represents the taxon later described as Bilimbia trachona var. intercedens
Arnold. This taxon was the only taxon in Bilimbia described as new from the area in question, and the locality data
fully agree with the material discussed in detail by Arnold (1873: 511–512). The type material of Bilimbia trachona
var. intercedens in UPS belongs to Bacidia verecundula (Th. Fr.) H. Magn., which was originally described as Bilimbia
verecundula Th. Fries (1874: 387). Note that Arnold (1877), when combining his taxon to species level, put forward
the idea of a close relationship with Bacidia verecundula, possibly after having communicated with Fries on the matter.
Also note that Patellaria intercedens Müll. Arg., on which Bacidia intercedens (Müll. Arg.) Zahlbr. is based, is a
different taxon.
Bilimbia subcoprodes de Lesd. in Crozals (1924: 107)
Bacidia subcoprodes (de Lesd.) Zahlbruckner (1932: 409). Type: FRANCE. Provence-Alpes-Côte d’Azur: Var,
“Giens”, February 1924, A. de Crozals (US 01138555!).
The type material in US is Catillaria aphana (Nyl.) Coppins. Most ascospores are 1-septate, although some are
non-septate and some 3-septate. This variation in ascospore septation is often observed in that species.
Acknowledgements
I am grateful to the curators listed in the Material and Methods for providing essential loans of herbarium material, to
Brian Coppins for directing my attention to British specimens that proved to belong to B. coprodes and B. inornata, and
to Francesco Di Carlo for hospitality during a visit to the Massalongo herbarium in Verona. The Swedish Taxonomy
Initiative is thanked for financial support (grant no. 146/07 1.4).
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