The genus Mycoblastus in the cool temperate Southern
Hemisphere, with special reference to Tasmania
Abstract: The genus Mycoblastus in cool temperate latitudes of the Southern Hemisphere is reviewed.
Eight species are treated in detail: M. bryophilus Imshaug ex Kantvilas sp. nov., from Campbell Island
and Tasmania; M. campbellianus (Nyl.) Zahlbr., M. coniophorus (Elix & A.W. Archer) Kantvilas & Elix
comb. nov. and M. dissimulans (Nyl.) Zahlbr., all widespread across the austral region; M. disporus
(C. Knight) Kantvilas comb. nov., from New Zealand and Tasmania; M. kalioruber Kantvilas sp. nov,
from Tasmania; M. sanguinarioides Kantvilas sp. nov., from Tasmania and south-eastern Australia;
and M. leprarioides Kantvilas & Elix sp. nov., from south-eastern Australia (Victoria). Notes are
provided on many other species of Mycoblastus, including those recognised for the Northern
Hemisphere, and those originally described from austral regions but now excluded from the genus.
Major characters of the genus are discussed, including thallus morphology and chemistry, apothecial
pigments and ascus structure. It is suggested that the genus is heterogeneous and that some of its
closest afﬁnities may lie with the family Megalariaceae and the genus Japewia.
Key words: ascus, austral, Japewia, lichens, Megalaria,Megalariaceae,Mycoblastaceae
The genus Mycoblastus was introduced by
Norman (1853), based on the widespread
species, Mycoblastus sanguinarius. Despite
being visually striking, the genus has received
scant attention from taxonomists. For the
temperate Northern Hemisphere, useful
accounts have been published by Schauer
(1964), James (1971), Tønsberg (1992) and
Watson & James (1992), the last reference
serving as a sound treatment of species
that occur in Europe and North America.
Additional species have been described from
Asia, including the Himalayas (M. indicum;
Awasthi & Agarwal 1968), Java (M. endo-
xanthus and M. grisomarginatus; Groenhart
1950, 1954), the Philippines (M. dendropho-
rus; Vainio 1921) and Japan (M. japonicus;
Müller Argoviensis 1891), whereas Aptroot
et al. (1997) recorded an unidentiﬁed species
from New Guinea. Most of the earlier
authors initially named their species in
Lecidea, and the transfer to Mycoblastus of
these as well as Southern Hemisphere taxa
was undertaken by Zahlbruckner (1926).
The ﬁrst species of Mycoblastus recognized
from austral regions was from Chile,
described by Nylander (1855) as Lecidea
dissimulans. Soon after, M. campbellianus
was described from Campbell Island (also
as a Lecidea) by Nylander (1876). Other
taxa soon followed, mainly based on New
Zealand collections, and the transfer of some
of these to Mycoblastus was effected by
Müller Argoviensis (1894). In more recent
times, Zahlbruckner (1941) and Dodge
(1948, 1965, 1970) added taxa, although the
latter author’s have since proved to have
been misplaced generically. However, apart
from the accounts by Galloway (1985, 2007)
and the unpublished thesis of Kantvilas
(1985), there has been no attempt to under-
take a revision of austral species, even though
herbarium collections clearly indicate that
the genus is a signiﬁcant component of local
Mycoblastus is characterized by a crustose
thallus containing a green coccoid photo-
biont, large, conspicuous, typically black or
G. Kantvilas: Tasmanian Herbarium, Private Bag 4,
Hobart, Tasmania, Australia 7001. Email:
The Lichenologist 41(2): 151–178 (2009) © 2009 British Lichen Society
doi:10.1017/S0024282909008238 Printed in the United Kingdom
dark-coloured, immarginate apothecia, fre-
quently rich in colourful pigments, richly
branched, anastomosing paraphyses, and
lecanoralean asci, typically containing either
one or two relatively large, hyaline, usually
simple ascospores. The genus was accommo-
dated within its own family, Mycoblastaceae,
by Hafellner (1984) and is retained there,
together with Tephromela, by Lumbsch &
Species of Mycoblastus occur mostly on
organic substrata, such as bark, humus or
peaty turf, in cool moist environments. On
the basis of the present study, the genus
consists of approximately 20 species, with
roughly equal representation in both hemi-
spheres. Whereas many species are wide-
spread in their particular hemisphere, no
species are shared between northern and
southern temperate zones.
Material and Methods
The study is based chieﬂy upon extensive herbarium
collections held in MSC, assembled by Dr H.A.
Imshaug and his co-workers during several expeditions
to various austral regions in the 1960s and 1970s, on the
author’s ﬁeld experience and collections from Tasmania
and south-eastern Australia, held in HO, and on selected
reference and type material from other herbaria (as cited
in the text).
Observations of apothecial anatomy were undertaken
using light microscopy on hand-cut sections, mounted
in water, 10% KOH (K), 50%HNO
(N), Lugol’s Iodine
(IKI) and lactophenol Cotton Blue. Initial measure-
ments of apothecial tissues were made in water, whereas
asci, ascospores, paraphyses and hyphae were measured
in K. With the exception of M. disporus,M. bryophilus
and M. leprarioides, for which fertile material was limited
(or absent), ascospore measurements are based on at
least 100 observations, and are presented in the format:
smallest measurement–mean–largest measurement;
single outlying values are given in parentheses.
Chemical analyses using thin-layer chromatography
followed standard methods (Orange et al. 2001), mainly
using solvents A and C, and comparison with a range of
reliable reference specimens. High-performance liquid
chromatography (Feige et al. 1993) of type material
and selected specimens was undertaken by J. A. Elix
in Canberra. Nomenclature of ascus types follows
In order to simplify the species descriptions, the
reactions of the main pigments in water, K and N are not
repeated throughout. The two standard pigments,
‘cinereorufa-green’ of Meyer & Printzen (2000) and
‘fucatus-violet’ are referred to as such in the text, and
their characteristics are summarized in the general
Authorities for lichen taxa cited in text are included
only at their ﬁrst mention.
Comparative reference material examined. Megalaria
grossa (Pers. ex Nyl.) Hafellner: Australia: Tasmania:
Little Fisher River, 41°45'S 146°20'E, 920 m, 1987, G.
Kantvilas 56/87 (BM, HO).
Megalaria laureri (Th. Fr.) Hafellner: USA:
Michigan: W of Kingston Lake, 1970, R. C. Harris 6055
Megalaria melaloma (C. Knight) Kantvilas:
Australia: New South Wales: Braidwood district, SE
of Rossi, 1967, W. A. Weber & D. McVean L-49319
Tasmidella variabilis Kantvilas, Hafellner & Elix
var. variabilis:Australia: Victoria: Baw Baw NP, Mt
Erica car park, 37°53'S 146°21'E, 1050 m, 2008, G.
Kantvilas 187/08 (HO, MEL).
Tephromela atra (Huds.) Hafellner: Autralia:
Tasmania: Edwards Road, 1980, G. Kantvilas 324/80
Japewia tornoensis (Nyl.) Tønsberg: USA.:
Washington: Falls Creek near Carson, 1994, G.
Kantvilas, J. Davis & D. Shaw s.n. (HO).
Japewiella tavaresiana (H. Magn.) Printzen: Ireland:
Lough Corrib, 53°30'N 9°22'W, 3 m, 1998, H. Hertel
39505 (H. Hertel: Lecideaceae Exsicc. 303) (HO).
General features of the genus
Species of Mycoblastus have a crustose,
potentially variable thallus, and the same
taxon may exhibit forms ranging from thin
and effuse to thick, warty or papillate. In
many cases, thallus morphology can be inter-
preted in terms of substratum and habitat
characteristics. At least one taxon has an
entirely granular-sorediate thallus resem-
bling a Lepraria. A prothallus is usually evi-
dent, and when the thallus is particularly thin
or dispersed, this may lend a purplish hue to
the whole lichen. The presence or absence
of soredia is usually important and used in
this account as a species-deﬁning charac-
ter. However, at present no obvious ‘species-
pairs’ in the sense of Poelt (1970) are
known, with the possible exception of the
Northern Hemisphere’sM. afﬁnis (fertile,
esorediate) and M. alpinus (usually sterile,
152 THE LICHENOLOGIST Vol. 41
Thallus chemistry is an important and use-
ful taxonomic character. The most common
compound occurring in Southern Hemi-
sphere taxa is the depside, perlatolic acid. In
the Northern Hemisphere, this compound is
known only from M. caesius, but the some-
what similar compound, planaic acid, is
more widespread. Atranorin, fumarproto-
cetraric acid and compounds allied to the
latter (physodalic and virensic acids) are also
known from the genus. Fatty acids have been
detected in most species but hitherto, despite
concerted efforts, no consistent pattern in
their occurrence has been found. They
often occur in only trace concentrations,
pose technical difﬁculties as regards their
detection and identiﬁcation, and are hence
considered of limited taxonomic value.
Apothecial pigments provide a major
character in the description of species of
Mycoblastus. Two main pigments were
observed in the species studied. First, there is
a blue-green pigment that approximates to
the ‘cinereorufa-green’ pigment of Meyer &
Printzen (2000). This pigment varies from
vivid to dull blue-green or olive-green. The
intensity and clarity of the pigment may
depend on habitat, but is also affected by
whether the apothecial tissues are inspersed
with granules, crystals or oil droplets. The
pigment is ± unchanged in K or, more
frequently, turns an olive-green to grey-
green. In N, the pigment turns crimson.
Cinereorufa-green pigment is recorded by
Meyer & Printzen (2000) in M. sanguinarius
and appears to be widespread in the genus.
The second pigment recorded is an intensely
violet pigment termed here ‘fucatus-violet’.
This pigment was described by James (1971)
from the European species, M. fucatus
(Stirt.) Zahlbr. It gives a K+ vivid turquoise-
green reaction; in N it slowly turns orange
and gradually fades to colourless.
These two pigments may occur separately
or together. When overlying each other or
intermixed, the resulting colour is often a
deep indigo-blue. In doubtful cases, conﬁr-
mation of which pigment is present can be
achieved by using the two main reagents:
using K, even minute quantities of fucatus-
violet will produce a vivid turquoise-green;
using N, even minute traces of cinereorufa-
green will yield a crimson reaction.
The disposition and intensity of the pig-
ments can vary greatly, but this is not consid-
ered an important taxonomic character.
Pigments may be conﬁned to an epihymenial
layer, or penetrate between the asci into the
hymenium, or be present at the edge of or
within the excipulum. Sometimes they may
also extend into a subhypothecial band. Such
variation can sometimes be interpreted as a
response to microhabitat (with increasing
exposure generally leading to heavier pig-
mentation) or to age of the apothecia (older
apothecia are typically darker).
Mycoblastus sanguinarius and its allies also
contain a deep red subapothecial pigment
that some authors (e.g. Watson & James
1992) call rhodocladonic acid. The relatively
rare yellow-orange pigment, hybocarpone,
occurs beneath the apothecia of M. kalioru-
ber; it yields an intense K+ red reaction.
Other unknown pigments (usually reddish
or yellowish) occur in some species, and
secalonic acid appears to be responsible
for some of the yellow, K+ orange-yellow
coloration in M. coniophorus.
Although the presence of crystals and oil
droplets in the apothecial tissues can be very
striking, it is applied only to a limited extent
as a taxonomic character in this work. The
degree of inspersion may also be linked to
habitat and age.
The photobiont of Mycoblastus is a uni-
cellular green alga with essentially globose to
irregularly subglobose cells, 5–16 µm wide.
The individual cells separate readily in
squash preparations in KOH. The photo-
biont is not Trebouxia (as seen, for example,
in Lecanora); this has substantially larger
cells. On the basis of purely qualitative, com-
parative observations of sections of lichen
thalli, the photobiont is similar to that seen in
Pertusaria or Megalaria. The photobiont of
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 153
the latter genus is thought to be Dictyochlo-
ropsis (Tschermak-Woess 1984).
The ascus of Mycoblastus sanguinarius was
depicted and described by Hafellner (1984)
who noted one-spored asci with a thick amy-
loid tholus, with a distinct ocular chamber
and masse axiale in the form of a pale border;
the ascus also had a prominent amyloid cap
composed of compacted gel. This type of
ascus, termed the ‘Mycoblastus-type’ after
Hafellner (1984), is also found in the other
one-spored taxa, including M. sanguinari-
oides (described below); it is depicted in
Fig. 1A where the similarities to Hafellner’s
drawing are clearly evident. The amyloid
reaction is restricted to the asci and sur-
rounding gel, and does not occur through the
whole hymenium. So intense is the amyloid
reaction that ascus structure is best observed
using very dilute Lugol’s reagent after
pre-treatment with KOH and washing with
A different type of ascus is found in the
other austral species studied. In these
species, the ascus is at least two-spored. The
tholus is very well-developed and intensely
amyloid, but is pierced totally (or almost so)
by a barrel-shaped to conical, weakly amy-
loid masse axiale; the tholus tissue adjacent
to the masse axiale usually stains slightly
more intensely. There is also a blunt to beak-
like ocular chamber, although this may be
absent in older asci where the apex of the
F. 1. Mycoblastus species, asci with the amyloid parts stippled. A, M. sanguinarioides (Kantvilas s.n., 1996); B,
M. dissimulans (Kantvilas 1086/81). Scale = 50 µm.
154 THE LICHENOLOGIST Vol. 41
ascoplasm is ± neatly rounded. The ascus
is surrounded by a thin, external, ‘fuzzy’
amyloid layer, but there is no thick, heavily
amyloid cap (Fig. 1B). This type of ascus
more closely approximates a Biatora-type or
Lecidella-type, and is not unlike that seen in
certain species of Megalaria (for example, see
Kantvilas 2008). As in M. sanguinarius, the
amyloid reaction is restricted to the asci
themselves, and is best observed using
extremely dilute Lugol’s reagent.
The structure of the paraphyses differs
between the species studied. Paraphyses are
invariably highly branched, but their thick-
ness varies, as does the degree to which they
separate in K. In M. dissimulans, for example,
they remain densely conglutinated, whereas
in M. campbellianus they are usually lax. The
paraphyses may also be rather robust and
become ± moniliform in the uppermost part.
The apices of all species generally remain
conglutinated, and a gelatinous cap is fre-
quently developed that encases the apical
cell. Pigment tends to occur around these
caps, so that in section, they give the impres-
sion of a cellular epithecium. It is still unclear
to what extent the variation in paraphyses is
induced by age or habitat, and this character
is not applied rigorously in the delimitation
The size of ascospores and their number in
the ascus is critical for species identiﬁcation.
Two states are found in Mycoblastus: broadly
ellipsoid to cylindrical ascospores that occur
almost always singly in the ascus (as found
in M. sanguinarius and M. sanguinarioides,
hereafter referred to as the ‘M. sanguinarius
group’; Fig. 2A); and ovate to broadly ellip-
soid ascospores that are usually two per ascus
(as found in the other species studied; e.g.
M. dissimulans; Fig. 2B). In the latter type, 3-
or even 4-spored asci are known, but these
additional spores are usually aborted before
The ascospores have a double wall that is
clearly discernible in light microscopy: there
is a thin inner wall and a thick, gelatinous
outer wall. Measurements cited in this paper
include both layers. Occasionally the outer
wall swells noticeably and ruptures, releasing
an ascospore with only the thin, inner wall.
Mycoblastus is generally regarded as
simple-spored (Hafellner 1984) and this is
certainly true for the M. sanguinarius group.
However, apparently septate ascospores have
been consistently observed in other species.
In many cases, these septate spores are
clearly old and deformed, and the septum is
frequently thin, crooked and more akin to a
plasma bridge as seen in many normally
simple-spored lichens. However, well-
formed ascospores with a distinct true
septum have been observed, albeit very occa-
sionally, in all members of the M. dissimulans
group (Fig. 2B).
Despite their superﬁcial similarities, there
appear to be two distinct groups evident
in Mycoblastus, supported by differences
in ascus structure, ascospores and thallus
chemistry. This infra-generic division
requires further exploration, a task that is in
progress but beyond the scope of the present
paper. Some species (M. afﬁnis and M. san-
guinarius) have been included in several
studies dealing with systematic relationships
based on DNA sequence data, albeit in a
cursory way. Strong support for the relation-
ship between Mycoblastus and Tephromela,
as well as with Calvitimela, was found by
Arup et al. (2007), Ekman et al. (2008) and
Mia˛dlikowska et al. (2006), although all
these authors concede that further studies are
needed. A comparative morphological and
anatomical analysis of the systematic place-
ment of Mycoblastus will be explored in a later
contribution. However, on the basis of ascus
structure, the presence of pigments, granules
and oil droplets, and the generally large, sim-
ple or one-septate ascospores, a relationship
with Megalaria and the family Megalariaceae
may be worthy of serious consideration. In
this family, the genus Tasmidella has occa-
sionally septate, double-walled ascospores
(Kantvilas et al. 1999). Even more similar,
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 155
and probably related, is the genus Japewia
(Tønsberg 1990; Printzen 1999; Spribille &
Printzen 2007). Japewia shares many charac-
ters with the M. dissimulans group, including
aLecidella-type ascus, reduced exciple,
branched and anastomosing paraphyses and
double-walled ascospores. It differs mainly
by lacking an inspersed hymenium and dark,
K+, N+ pigments. Salient features of these
genera and other possibly related taxa are
compared in Table 1.
Key to austral species of Mycoblastus
1 Thallusnotsorediate ................................2
F. 2. Mycoblastus species, ascospores. A, M. sanguinarioides (Kantvilas s.n., 1996); B, M. dissimulans (Kantvilas
1086/81). Scale = 50 µm.
156 THE LICHENOLOGIST Vol. 41
2(1) Asci one-spored, with the ascospore ellipsoid to oblong, usually >60 µm long;
apothecia frequently with small patches of reddish pigment beneath; thallus
containing atranorin . . . . . . . . . . . . . . . . . . . . M. sanguinarioides
Asci at least two-spored, with ascospores ellipsoid to ovate, usually <60 µm long; red
pigments not present beneath apothecia; thallus lacking atranorin but containing
3(2) Thallus P+ orange-red (containing physodalic acid) . . . . . . . . . M. disporus
Thallus P−(physodalic acid lacking) . . . . . . . . . . . . . . . . . . . . . . . . 4
4(3) Upper part of hymenium containing cinereorufa-green and/or fucatus-violet pig-
ments; hypothecium colourless to pale yellow, usually K+ yellowish or yellow-
orange; containing perlatolic acid ± fatty acids . . . . . . . . M. dissimulans
Upper part of hymenium with cinereorufa-green pigment only; hypothecium vivid
yellow, K+ intense blood red; containing hybocarpone in addition to perlatolic
andfattyacids........................... M. kalioruber
5(1) Soredia usually whitish or at least paler than the thallus, occurring in discrete soralia,
becoming diffuse and conﬂuent only in later stages of development . . . . . . 6
Soredia concolorous with the thallus, not in discrete soralia, arising from cracks and
soon becoming scattered and conﬂuent, sometimes forming a granular crust . 7
6(5) Soralia P+ orange-red (containing virensic acid in addition to perlatolic acid);
soredia farinose; typically epiphytic or on wood . . . . . . M. campbellianus
Soralia P−(containing perlatolic acid, ± fatty acids); soredia granular; typically on
7(5) Thallus typically fertile; soredia scattered at ﬁrst, later becoming conﬂuent, P−
(containing perlatolic acid, ± fatty acids); very common and widespread . . . .
Fertile material unknown; soredia conﬂuent from the outset and forming a thick,
granular crust, P+ red (containing protocetraric acid in addition to perlatolic acid)
.................................. M. leprarioides
Mycoblastus bryophilus Imshaug ex
Kantvilas sp. nov.
Mycoblasto dissimulanti (Nyl.) Zahlbr. similis sed thallo
granulari, areolato vel noduliformi, sorediis grossis
granularibusque in soraliis discretis, 0·5–2·5 mm latis
Type: New Zealand, Campbell Island, rock outcrops
on north side of west end of Lyall ridge at 500–600 ft
[150–180 m] elevation, 18 January 1970, R. C. Harris
5539 (MSC—holotypus; HO—isotypus).
(Fig. 4A & B)
Thallus whitish cream, rarely greenish or
brownish grey, matt, composed of scattered,
crowded or contiguous granules, areoles or
nodules mostly 0·2–1·5 mm wide, smooth or
abraded, becoming sorediate; soralia con-
colorous with the thallus or paler, rather
erumpent, usually ± discrete, c. 0·5–2·5 mm
wide; soredia very coarsely granular, some-
times fusing in irregular patches to 10 mm
wide but not forming a continuous sorediate
crust; prothallus effuse, often rather glossy
and ± gelatinous, blue-grey to brownish; cor-
tex absent; medulla white; photobiont cells ±
globose, 5–12 µm diam.
Apothecia very rare and usually not devel-
oped, 0·3–1·5 mm diam., dull to glossy black
to brown-black, superﬁcial, markedly convex
to subglobose, basally constricted, immar-
ginate. Exciple in section, 20–90 µm thick,
hyaline to pale brownish, usually streaked
with varying amounts of cinereorufa-green
and fucatus-violet pigments, composed of
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 157
T 1. Comparison of salient features of Mycoblastus and superﬁcially similar genera (based on observation of selected specimens and literature).
atra-red red-brown, K±
paraphyses branched and
mainly simple simple or
mainly simple branched and
158 THE LICHENOLOGIST Vol. 41
Table 1. continued.
ascospores ellipsoid to ob-
100 µm long
to 35 µm
to 45 µm
walled, to 20
walled, to 16
walled, to 20
walled, to 20
conidia bacilliform bacilliform ellipsoid to ob-
ampulliform bacilliform to
thread-like unknown bacilliform
chemistry atranorin, ±
± fatty acids
nil mostly nil, ±
Ekman & Tønsberg (1996);
Kantvilas et al. (1999);
Spribille & Printzen (2007);
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 159
radiating, heavily conglutinated hyphae,
1–2·5 µm thick. Hypothecium 140–250 µm
thick, hyaline to pale yellow-brown, dilute
yellowish in K, inspersed with oil droplets
2–12 µm wide, subtended by a hyaline to pale
yellowish subhypothecium, intensifying yel-
low in K, and a band of cinereorufa-green
pigment. Hymenium to c. 150 µm thick, hya-
line to pale yellowish, likewise inspersed with
oil droplets, pigmented in the upper part with
a thick layer of cinereorufa-green pigment
that overlies additional fucatus-violet pig-
ment. Asci 2-spored, approximating the
Biatora- or Lecidella-types, 130–150 ×40–50
µm; very few mature asci observed. Paraphy-
ses numerous, highly branched, separating
easily in K or remaining conglutinated, 1·5–4
µm thick; apices mostly 3–6 µm thick, occa-
sionally with the ultimate cells encased in
pigment. Ascospores ovate to broadly ellip-
soid, simple, 50–58·7–66 ×26–34·5–40 µm
(very few observed); wall (1–)3–4 µm thick.
Pycnidia not found.
Chemistry. Perlatolic acid, rangiformic acid
(± trace), norrangiformic acid (± trace) and
norcaperatic acid (± trace); thallus K−,
KC−, C−, P−, UV± whitish.
Etymology. The speciﬁc name refers to the
habitat of the new species, which commonly
overgrows terricolous bryophytes (and other
small plants). The name was ﬁrst applied as
an unpublished herbarium name to material
collected and distributed by Henry Imshaug
Remarks. Mycoblastus bryophilus is closely
related to M. dissimulans (see below) in so far
as both species contain perlatolic acid as
the major compound, and have apothecia
usually with both cinereorufa-green and
fucatus-violet pigments. They differ by the
former being sorediate. The soredia in this
taxon differ from those of the other sorediate
species found in the study area in that they
occur in delimited, albeit irregular soralia
and are extremely coarse. In M. coniophorus
and M. leprarioides, the soredia are diffuse
and occur ± across the entire thallus, where-
as in M. campbellianus they are farinose.
Although the ascospores of M. bryophilus
appear to be marginally larger than those of
M. dissimulans, this may be an artefact of a
signiﬁcantly smaller sample size for the
In cold, treeless environments, both M.
bryophilus and M. dissimulans occur together
and intermixed; indeed much of the MSC
material distributed under the former name
is non-sorediate and, in this study, is ascribed
to M. dissimulans. Specimens need to be
checked carefully, bearing in mind that the
thallus of M. dissimulans from terricolous
habitats may be likewise granular-nodulose
and quite abraded but does not form soredia,
whereas M. bryophilus forms discrete soralia
that sometimes become very prominent and
erumpent. Such specimens may resemble
sterile thalli of Dibaeis arcuata (Stirt.) Kalb &
Gierl that may grow in similar habitats.
Only two specimens of this taxon had
apothecia, and then only very few and with
very few, well-developed ascospores; conse-
quently the description and measurement of
fertile characters are scant and provisional.
Distribution and ecology. Mycoblastus bryo-
philus is known from Campbell Island, where
the abundance of collections suggests it is
common, and from Tasmania. It occurs
mainly over bryophyte turfs, peaty soil, litter
or the rotting bases of plant tussocks in cold,
windswept, treeless habitats. However, of the
ﬁve Tasmanian collections studied, one is
from the papery bark of a Leptsopermum in
highland woodland, another is from the dead
fronds of the tiny epiphytic fern, Apteropteris
applanata, which grows on Athrotaxis in
alpine woodland, and a third is from rock.
Specimens examined. New Zealand: Campbell Island:
St Col Peak, 1969, H. A. Imshaug 45990 (MSC); sum-
mit of Mt Dumas, 1970, H. A. Imshaug 47005 (MSC);
Lyall ridge, 1969, R. C. Harris 4399 (HO, MSC); Paris-
Villarceau ridge, 1970, H. A. Imshaug 46616 (HO,
MSC); NE of Mt Sorenson, 1970, R. C. Harris 5120
(MSC).—Tasmania: c. 2 km SE of Lake Emmett,
42°38'S 146°33'E, 980 m, 1990, G. Kantvilas 151/90
(HO); summit of Black Bluff, 41°27'S 145°57'E, 1335
m, 2000, G. Kantvilas 132/00 (HO); Mt Wedge Track,
42°51'S 146°17'E, 1000 m, 2002, G. Kantvilas 622/02
(HO); King William Saddle, 42°13'S 146°06'E, 840 m,
1984, G. Kantvilas 177/84 & P. W. James (BM, HO).
160 THE LICHENOLOGIST Vol. 41
Flinders Island: head of Leventhorpe Gully, 40°04'S
148°05'E, 330 m, 2007, G. Kantvilas 68/07 (HO).
Mycoblastus campbellianus (Nyl.)
Cat. lich. univ. 4: 3 (1926) —Lecidea campbelliana Nyl.,
C. r. hebd. Séanc. Acad. Sci. Paris 83: 90 (1876); type:
New Zealand, Insula Campbell, 1874, Filhol (H-NYL
Mycoblastus minor Zahlbr., Denkschr. Akad. Wiss.
Wien math.-naturwiss. Kl. 104: 294 (1941); type: New
Zealand, East Waiotapu Valley, Rotorua, June 1936, on
manuka twigs, K.W. Allison 28 (CHR 378615!—
lectotype, ﬁde Galloway 1985).
(Fig. 3A & B)
Thallus whitish, cream, grey-brown or dull
greyish, sometimes patchily blue-grey due to
the underlying prothallus, matt, ± continu-
ous, smooth or a little uneven, generally
deeply cracked, 40–180 µm thick, sorediate;
soralia white, 0·2–0·5(−1·2) mm wide,
initially discrete, speck-like or sometimes
tuberculate, soon becoming diffuse or con-
ﬂuent and spreading unevenly across the
thallus; soredia farinose to granular; cortex
absent; medulla white; photobiont cells ±
subglobose, 5–11 µm diam.
Apothecia 0·3–0·5(−0·8) mm diam., scat-
tered, ± superﬁcial, basally constricted,
strongly convex and top-shaped when well
developed, occasionally sunken in the thallus
surface, abraded or excavate, glossy black to
blue-black, rarely dark reddish brown or pie-
bald, immarginate. Exciple in section often
excluded, in young apothecia 20–60 µm
thick, with bright cinereorufa-green pigment
at the outer edge, composed of radiating,
very heavily conglutinated hyphae, 2–4 µm
thick, at the outer edge expanding to 4–5 µm
thick. Hypothecium 85–200 µm thick, colour-
less to pale yellowish, intensifying in K and
N, sometimes yellow-orange, inspersed with
minute crystals that mostly dissolve in K,
occasionally with a subhypothecial band of
cinereorufa-green pigment. Hymenium 80–
140 µm thick, inspersed with oil droplets
and minute granules that mostly dissolve in
K, entirely or in the upper part bright
cinereorufa-green pigmented. Asci 2(−4)-
spored, approximating the Biatora- or
Lecidella-types, 70–98 ×24–47 µm. Para-
physes numerous, highly branched, mostly
separating readily in K, 2–3·5 µm thick, ±
moniliform in the upper part; apices typically
enlarged, 3–5 µm thick, in extreme cases
the ultimate cells developing roundish,
F. 3. Mycoblastus campbellianus (Kantvilas 369/07). A, habit; B, ascospores. Scales: A = 1 mm; B = 40 µm.
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 161
blue-green pigmented, gelatinous caps, 8–13
×7–12 µm that form a conglutinated, ‘cellu-
lar’ epithecial layer 15–40 µm thick. Asco-
spores ovate to broadly ellipsoid, 27–38·7–
51(−56) ×17–22·4–29(–32) µm, 0(–1)-
septate; wall (1·5–)2·5–5 µm thick.
Pycnidia not found.
Chemistry. Perlatolic acid, hyperperlatolic
acid, virensic acid and protocetraric acid (±);
very rarely with additional caperatic or nor-
caperatic acids; thallus and soralia K−, C−,
KC−, P+ orange-red, UV+ white. Perlatolic
acid is invariably the major compound
present whereas the concentration of the
other compounds varies. Virensic acid is
sometimes barely detectable on a TLC
plate, but the soralia nevertheless give a P+
Remarks. Mycoblastus campbellianus is a
very distinctive species in the austral ﬂora,
characterized by a thallus with farinose sore-
dia that occur in discrete, roundish, white
soralia, by the presence of virensic acid in
addition to perlatolic acid, and by the rela-
tively small apothecia and ascospores. The
presence of the blue-green, cinereorufa-
green pigment in the apothecia allies this
species to M. bryophilus, M. coniophorus, M.
dissimulans and the Northern Hemisphere’s
M. afﬁnis, M. caesius and M. sanguinarius, but
it differs from all of these species by its chemi-
cal composition, readily conﬁrmed by the P+
orange-red reaction of the soralia. It differs
further from M. dissimulans (with which it
often grows) by a range of characters: the
thallus of M. dissimulans is much thicker
and often areolate, papillate or verrucose,
whereas that of M. campbellianus is thin and
generally ± smooth; M. dissimulans may be
coarsely abraded but not truly sorediate. Of
the species listed above, only M. bryophilus,
M. caesius and M. coniophorus are truly
sorediate but in those species the soredia are
very coarse, and form at cracks in the thallus,
at the edges of slightly upturned areoles or on
points of abrasion, rather than in discrete
The apothecia and ascospores of M. camp-
bellianus are generally a little smaller than
those of the other species studied. The
hymenium is also not as densely inspersed,
and then mainly with crystals rather than oil
droplets; probably for this reason, the blue-
green pigment appears much clearer and
brighter. The crystals ﬂuoresce in polarized
light and dissolve in KOH. The paraphyes of
M. campbellianus are also distinctive, separat-
ing readily in K and being generally more
robust and moniliform than those of the
other species mentioned above.
Two unusual specimens of this species
have been collected in Tasmania. One
(Kantvilas s.n.) lacks virensic acid but
accords with M. campbellianus in every other
macro- and micro-character. The other
(Kantvilas 22/00) is from rock and contains
additional atra-red (Meyer & Printzen 2000)
pigment in the subhypothecium. A further,
as-yet unidentiﬁed, superﬁcially similar
species occurs in the New Guinea highlands
(collected by A. Aptroot); it differs by con-
taining perlatolic acid and atranorin, and has
fucatus-violet pigment in the apothecia.
The thallus of M. campbellianus varies from
being thick and prominent to very thin,
but the white ﬂeck-like soralia are always
conspicuous. Sterile specimens are common
and may resemble three other sorediate,
frequently sterile lichens that occur in simi-
lar habitats: Austroblastenia pauciseptata
(Shirley) Sipman (containing pannarin; P+
orange), Pertusaria novaezelandiae Szatala
(containing hypothamnolic acid; K+ violet)
and an unidentiﬁed Trapelia (containing
gyrophoric acid; C+ red).
Although the detailed description of Myco-
blastus minor given by Zahlbruckner (1941)
concurs with all the salient features of M.
campbellianus, the synonymy of these names,
proposed by Galloway (1985) and accepted
here, could not be conﬁrmed by examination
of the type of the former. The specimen
studied (in CHR) includes a wide range of
crustose lichens, dominated by a Tephromela,
but no species of Mycoblastus was evident.
Distribution and ecology. Mycoblastus camp-
bellianus is a species of cool to cold climates,
and has been recorded from the North and
South Islands of New Zealand (Galloway
162 THE LICHENOLOGIST Vol. 41
2007), Tasmania (Kantvilas et al. 2008),
Campbell Island, Macquarie Island (Dodge
1948), the highlands of south-eastern
Australia, Staten Island, Tierra del Fuego
and southern Chile. It typically occurs as an
epiphyte, although one specimen seen is
from peaty soil and another is from rock.
In Tasmania, it occurs in a wide range of
vegetation types, including cool temperate
rainforest, wet eucalypt forest, wet scrub,
subalpine heathland and alpine communi-
ties. It typically colonizes a wide range of
bark types as well as decorticated wood, and
is part of a diverse community that includes
Coccotrema cucurbitula (Mont.) Müll. Arg.,
Fuscidea australis Kantvilas, Loxospora soleno-
spora (Müll. Arg.) Kantvilas, Melanelia sub-
glabra (Räsänen) Essl., Menegazzia globulif-
era R. Sant., M. pertransita (Stirt.) R. Sant.,
Pannaria microphyllizans (Nyl.) P.M. Jørg.,
Parmelia tenuirima Hook.f. & Taylor, P.
salcrambidiocarpa Hale, Tasmidella variabilis
Kantvilas et al. and Thelotrema lepadinum
(Ach.) Ach. Also often present together with
M. campbellianus are M. coniophorus, M. dis-
simulans and M. kalioruber.
Selected specimens examined. Argentina: Isla de los
Estados: Puerto Basil Hall, 54°45'S 64°10'W, 1971,
H. A. Imshaug 51405 & K. Ohlsson (MSC); Puerto Cook,
54°46'S 64°02'W, 10 m, 1971, H. A. Imshaug 51636 &
K. Ohlsson (MSC); Puerto San Juan, 54°45'S 63°54'W,
1971, H. A. Imshaug 51825 & K. Ohlsson (MSC); Puerto
Parry, 54°47'S 64°22'W, 1971, H. A. Imshaug 53885 &
K. Ohlsson (MSC); Bahia Capitan Canepa, 54°49'S
64°28'W, 45 m, 1971, H. A. Imshaug 53084 & K.
Ohlsson (MSC); Bahia Crossley, 54°48'S 64°40'W,
1971, H. A. Imshaug 50844 & K. Ohlsson (MSC).—
Tierra del Fuego: Bahia Valentin, 54°53'S 65°29'W,
10–15 m, 1971, H. A. Imshaug 50488a & K. Ohlsson
(MSC).—Chile: Prov. Magallanes: B. Fortescue,
53°42'S 72°01'W, 1969, H. A. Imshaug 44924 & K.
Ohlsson (MSC).—New Zealand: Campbell Island: S of
Tucker Cove Station, 1969, R. C. Harris 4878 (MSC);
N of Beeman Station, 1970, R. C. Harris 5771 (MSC);
Garden Cove toward Filhol Peak, 1970, R. C. Harris
5197 (MSC); N side of Perseverance Harbour, 1969,
R. C. Harris 4833 (MSC); E of Mt Sorenson, 1970,
R. C. Harris 5164 (MSC); S slope of Mt Honey, 1970,
H. A. Imshaug 4783 (MSC); base of Lyall Ridge, 1969,
H. A. Imshaug 46075 (MSC).—Tasmania: Little
Fisher River, 41°45'S 146°20'E, 880 m, 1984, G.
Kantvilas 437/84 & P. James (BM, HO); Lake Dobson,
42°41'S 146°35'E, 1030 m, 1981, G. Kantvilas 652/81
&P. James (BM, HO); track to Clear Hill, 42°41'S
146°16'E, 1000 m, 2000, G. Kantvilas 27/00 (HO); Mt
Maria, 42°37'S 148°07'E, 705 m, 2000, G. Kantvilas
22/00 (HO); Lake Adelaide, 41°50'S 146°15'E, 1060
m, 2002, G. Kantvilas 68/02 (HO); near Little
Florentine River, 42°44'S 146°25'E, 430 m, 2000, G.
Kantvilas 70/00 (HO); Boyd Lookout, 42°49'S
146°21'E, 2006, G. Kantvilas 282/06 (HO); Little
Fisher River, 41°45'S 146°20'E, 880 m, 1982, G.
Kantvilas s.n. (HO); Long Tarns, 41°47'S 146°21'E,
1270 m, 2006, G. Kantvilas 405/06 (HO); Blue Peaks,
41°43'S 146°23'E, 1290 m, 2006, G. Kantvilas 416/06
(HO).—Australia: Victoria: Errinundra Saddle, Rain-
forest Walk, 37°19'03$S 148°50'19$E, 910 m, 2008,
G. Kantvilas 177/08 & J. Elix (HO, MEL); Errinundra
Plateau, Bonang River Picnic Area, 37°17'02$S
148°48'25$E, 810 m, 2008, G. Kantvilas 131/08 & J.
Mycoblastus coniophorus (Elix & A. W.
Archer) Kantvilas & Elix comb. nov.
Pertusaria coniophora Elix & A. W. Archer, in J. A. Elix,
A. Aptroot & A. W. Archer, Mycotaxon 64: 20 (1997);
type: Australia, Victoria, Cape Conran, 18 km E of
Marlo. 37°49'S 148°44'E, on dead wood, 21 November
1978, J. A. Elix 5297 (CANB!—holotype).
Pertusaria coniophorella Elix & A. W. Archer, Austral.
Lichenol. 60: 21 (2007); type: Australia, New South
Wales, Cotten-Bimbang National Park, junction of
Oxley Highway and Tobins Road, c. 70 km E of Walcha,
31°22'22$S 152°03'37$E, 1040 m, on dead wood in
wet Eucalyptus forest with tree fern understorey, 28 April
2005, J. A. Elix 36398 (CANB!—holotype).
Thallus whitish cream to pale yellowish
white, occasionally tinged a little greyish
green, 40–350 µm thick, continuous, smooth
to deeply cracked to rather scurfy-areolate,
sometimes rather verruculose-lumpy, soredi-
ate; soredia concolorous with the thallus,
very coarsely granular, arising from cracks in
the thallus or at the edges of areoles, at length
spreading across the thallus and sometimes
forming a granular-sorediate crust to c. 1 mm
thick; cortex absent; medulla white, but
sometimes locally yellowish beneath the
apothecia; photobiont cells subglobose, 7–12
Apothecia 0·5–1·5(−2) mm diam., scat-
tered or rarely conﬂuent, superﬁcial, strongly
convex to subglobose, basally constricted,
dull or glossy blue-black to black, occasion-
ally devoid of pigment and then pale reddish
brown or piebald, immarginate. Exciple in
section usually well developed, persistent,
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 163
F.4.Mycoblastus species, habit. A, M. bryophilus (Kantvilas 622/02); B, M. bryophilus (isotype); C, M. coniophorus
(Kantvilas 97/85); D, M. disporus (holotype); E, M. dissimulans (Kantvilas 1086/81); F, M. kalioruber (Kantvilas
154/90); G, M. leprarioides (holotype); H, M. sanguinarioides (Kantvilas s.n., 1996). Scale = 2 mm.
164 THE LICHENOLOGIST Vol. 41
35–75 µm thick, hyaline to pale yellow-
brown, colour intensifying in N, K+ yellow,
composed of very densely conglutinated,
branched and anastomosing, radiating
hyphae 1–3 µm thick. Hypothecium 140–200
µm thick, hyaline to pale yellow brown, ±
unchanged or intensifying yellow in K and N,
densely inspersed with oil droplets 3–20 µm
wide and minute granules mostly insoluble in
K, subtended by a pale yellow-brown sub-
hypothecium, intensifying yellowish in N,
K+ intense yellow to orange-yellow, occa-
sionally with a band of cinereorufa-green pig-
ment beneath. Hymenium 110–170 µm thick,
hyaline to pale yellow-brown, intensifying
yellowish in K, inspersed with granules and
oil droplets as in the hypothecium, in the
upper part densely cinereorufa-green pig-
mented, sometimes with the pigment diffus-
ing into the whole hymenium. Asci 2-spored,
approximating the Biatora- or Lecidella-
types, 90–125 ×30–44 µm. Paraphyses highly
branched, persistently heavily conglutinated
and not separating in K, ± evenly 1·5–3 µm
thick, occasionally somewhat moniliform
towards the apices; apices not or slightly
enlarged to 3–4 µm, sometimes with gelati-
nous caps 8–15 µm wide. Ascospores broadly
ellipsoid to ovate, (27–)29–43·1–60 ×17–
24·2–34(–38) µm, simple or rarely spuriously
1-septate; wall 2–5 µm thick.
Pycnidia not found.
Chemistry: Perlatolic acid, frequently
accompanied by either, or both, caperatic
acid and norcaperatic acid, often only in
trace concentrations; secalonic acid some-
times detectable by TLC (possibly conﬁned
to apothecia); thallus (particularly sorediate
areas) K−, C−, KC−, P± weak orange, UV+
Remarks. This species is characterized by
the sorediate thallus containing perlatolic
acid, the two-spored asci and by the presence
of cinereorufa-green pigment in the apoth-
ecia. Only three other sorediate species are
known in the study area: M. campbellianus,
which has farinose soredia in discrete soralia
and contains additional virensic and/or
protocetraric acids; M. bryophilus, which is
chieﬂy terricolous, often contains additional
fucatus-violet pigment, and has coarse sore-
dia in discrete, erumpent soralia; and M.
leprarioides, which forms an entirely granular-
sorediate crust, and contains additional
The most similar species to M. coniophorus
is the Northern Hemisphere taxon, M.
caesius. Specimens from the Paciﬁc North-
West of North America (collected and deter-
mined by T. Tønsberg) have a similar
thallus, likewise containing perlatolic acid,
with scattered coarse soredia and similar-
sized ascospores, 42–50 ×20–29 µm. How-
ever, M. caesius differs by several subtle
characters. Its thallus is thinner, often rather
dispersed and discontinuous, and conspicu-
ously delimited and underlain by a dark
prothallus, giving it a purplish tinge. The
apothecia of M. caesius sometimes contain
additional ‘fucatus-violet’ pigment together
with or just beneath the cinereorufa-green
pigment, yielding an indigo-blue tinge;
fucatus-violet pigment has never been ob-
served in M. coniophorus. Furthermore, the
K+ yellow reaction in the exciple and sub-
hypothecium of M. caesius is absent or very
weak, the hymenium is less inspersed, and
the paraphyses are generally more robust and
more lax in K. Whereas some particular
specimens of M. coniophorus could well be
accommodated within a broad concept of M.
caesius, after an examination of a very large
range of material, I have preferred to keep the
taxa separate. Certainly no material of M.
caesius seen ever approaches the typical form
of M. coniophorus, with its relatively thick,
coarsely granular sorediate, cream-coloured
The species was initially described as a
Pertusaria, based on a sterile specimen. It was
characterized by its chemical composition,
comprising perlatolic, conﬂuentic and 2-O-
methylconﬂuentic acids, and by the presence
of isidia (Elix et al. 1997). The interpretation
of the latter feature is unusual, because
the type specimen has a clearly abraded,
verruculose-sorediate thallus. The cited
chemical composition was incorrect and
the type specimen contains perlatolic acid
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 165
Distribution and ecology. This species is
currently known from southern Chile, Juan
Fernandez, Auckland Island, Macquarie
Island, Tasmania and south-eastern
Australia, where it is usually found as an
epiphyte in scrub and forest, or more rarely,
on soil and turf in treeless vegetation. With
some hesitation, it is also recorded here from
Prince Edward Island, based on material pre-
viously identiﬁed as M. caesius by Øvstedal &
Gremmen (2007); this small, sterile speci-
men is highly abraded, and is not greatly
dissimilar from some forms of M. bryophilus.
On the basis of herbarium material available,
M. coniophorus is particularly common in
Tasmania, but is much rarer elsewhere. It
seems surprising that no specimens from the
main islands of New Zealand could be
located, nor is any species that matches M.
coniophorus referred to by Galloway (1985,
In Tasmania, M. coniophorus is extremely
common and widespread, although mainly in
the lowlands and essentially outside of
the south-west of the island, a region with
many distinctive and/or endemic elements
(Kantvilas 1995). It is found mostly in cool
temperate rainforest, wet sclerophyll forest
and wet scrub. Here it occurs on a wide
variety of trees and shrubs, including species
with rough, ﬂaky bark such as Nothofagus
cunninghamii, species with smooth bark such
as Nematolepis squamea, and species with
papery bark such as Melaleuca species. It is
typically part of a rich assemblage of chieﬂy
crustose lichens, dominated by Thelotrema
lepadinum and Coccotrema cucurbitula. It is
also occasionally found in dry sclerophyll
forest where it favours locally moister,
more shaded sites such as fallen logs. Two
Tasmanian specimens are from rock: one in
buttongrass moorland, the other in eucalypt
Selected specimens examined. Chile: Prov. Magallanes:
SE point of Isla Williams, 48°46'S 74°24'W, 1969, H.
A. Imshaug 43419 & K. Ohlsson (MSC). Prov. Chiloe:
Puerto Ballena, 44°08'S 73°29'E, 1969, H. A. Imshaug
43134 & K. Ohlsson (MSC).—Juan Fernandez: Mas a
Tierra: Valle Anson, slope of Cerro Damajuana, 210 m,
1965, H. A. Imshaug 38256B (MSC).—New Zealand:
Auckland Island: Sealers Creek Cove, mouth of Laurie
Harbour, 1973, H. A. Imshaug 57862 (MSC).—
Macquarie Island: between Bauer Bay and Mawson
Point, 54°33'35$S 158°51'40$E, 6 m, 1986, R. D.
Seppelt 15918 (HO).—Tasmania: Mt Cameron,
40°59'S 147°56'E, 350 m, 1997, G. Kantvilas 124/97 &
J. A. Elix (HO); Ben Ridge Road, 830 m, 1981, G.
Kantvilas 1100/81 (HO); Granville Harbour, 41°49'S
145°02'E, 20 m, 1984, G. Kantvilas 231/84 & P. James
(BM, HO); summit ridge of Mt Scott, 41°18'S
147°31'E, 965 m, 2002, G. Kantvilas 223/02 (HO); Cox
Bight, West Beach, 43°29'S 146°13'E, seal-level, 1985,
G. Kantvilas 97/85 (BG, BM, HO, MSC); W of Tahune
Bridge, 43°06'S 146°40'E, 90 m, 2003, G. Kantvilas
100/03 (HO); c. 3 km S of Teepookana, 42°13'S
145°26'E, 220 m, 1990, G. Kantvilas 651/90 (HO);
Bruny Island, Coal Point, 43°20'S 147°19'E, 2 m,
2005, G. Kantvilas 140/05 (HO); Maria Island,
Montgomery Creek, 42°40'S 148°07'E, 5 m, 2003,
K. Felton s.n. (HO); slopes of Mt Albert, 42°21'S
147°52'E, 850 m, 2008, G. Kantvilas 251/08 (HO).
Flinders Island: Mt Killiecrankie, 39°49'S 147°52'E,
310 m, 2006, G. Kantvilas 37/06 (HO).—Australia:
Victoria: Errinundra Saddle, Rainforest Walk,
37°19'03$S 148°50'19$E, 910 m, 2008, G. Kantvilas
174/08 & J. Elix (HO, MEL); Errinundra Plateau,
Bonang River Picnic Area, 37°17'02$S 148°48'25$E,
810 m, 2008, G. Kantvilas 132/08 & J. Elix (HO). New
South Wales: Mt Hyland Nature Reserve, 20 km N of
Hernani, 30°10'44$S 152°25'19$E, 1340 m, 2005,
J. A. Elix 36633 (CANB, HO); Monga State Forest,
near Mongarlowe River, 35°35'S 149°55'E, 1986, G.
Kantvilas s.n. (HO).—Prince Edward Island: S of
Golden Gate, 2003, N.J. Gremmen G03-204 (BG).
Mycoblastus disporus (C. Knight)
Kantvilas comb. nov.
Megalospora dispora C. Knight, Trans. N. Z. Inst. 12: 378
(1880); type: New Zealand, Charles Knight 305
(WELT–hb. Knight Vol. 58A, p. 4!—holotype).
Thallus whitish, matt, rather waxy, ± con-
tinuous, smooth or more commonly un-
evenly lumpy and verruculose, cracked, to
300 µm thick, not sorediate; cortex absent;
medulla white; photobiont cells ± globose,
8–16 µm diam.
Apothecia 0·5–1·5 mm diam., scattered,
superﬁcial, basally constricted, strongly
convex to hemispherical, glossy black to
blue-black or brown-black, immarginate.
Exciple in section soon ± excluded, in young
apothecia 30–60 µm thick, hyaline to pale
brownish, or with cinereorufa-green and
fucatus-violet pigments throughout or at the
outer edge, composed of radiating, branched
and anastomosing, conglutinated hyphae,
166 THE LICHENOLOGIST Vol. 41
2–3 µm thick, expanding to 3–4 µm thick at
the outer edge. Hypothecium 100–160 µm
thick, colourless, unchanged in K and N,
usually inspersed with oil droplets 2–10 µm
diam., subtended by a colourless to pale
straw-coloured subhypothecial band, like-
wise inspersed, unchanged or intensifying
yellowish in K. Hymenium 150–180 µm thick,
inspersed with oil droplets, in the upper part
with a thick band of cinereorufa-green pig-
ment, with or without additional fucatus-
violet pigment, with the pigments diffusing
down between the asci. Asci elongate-clavate,
(1–)2(–4)-spored, approximating the Biatora-
or Lecidella-types, 100–150 ×36–50 µm.
Paraphyses numerous, highly branched,
mostly separating readily in K, ± evenly
2–3 µm thick; apices sometimes pigmented
and usually enlarged to 3–5(–6) µm wide.
Ascospores ovate to broadly ellipsoid, 40–
61·5–66(–70) ×18–34·3–38 µm, simple or
spuriously 1-septate; wall 3–4(–6) µm thick.
Pycnidia not found.
Chemistry: Perlatolic acid, physodalic acid,
with traces of protocetraric acid; thallus
K−, KC−, C−, P+ orange-red, UV+ faint
Remarks. The esorediate thallus contain-
ing perlatolic acid, the two-spored asci and
the presence of cinereorufa-green and
fucatus-violet pigments ally this species to
M. dissimulans. It differs solely in its chemi-
cal composition, namely the presence of
additional physodalic and protocetraric acids
and the absence of any fatty acids. Only three
specimens of this taxon were available for
study. The type (from New Zealand) differs
from the two Tasmanian specimens by hav-
ing somewhat smaller ascospores (40–48 ×
18–24 µm); it contains only cinereorufa-
green pigment. The two Tasmanian speci-
mens differ from each other in that one
(Kantvilas 66/99) has high concentrations of
both pigments, whereas the other has only
cinereorufa green. However, so unusual is
the chemistry of the three specimens, notably
in the presence of physodalic acid, unre-
corded in any other species of the genus, that
I have elected to treat all three specimens
as conspeciﬁc pending further material
Distribution and ecology. Mycoblastus dis-
porus has been recorded from Tasmania,
where it is uncommon, and from New
Zealand. In Tasmania, it has been collected
at only two localities: from the bark of the
low shrub, Richea sprengelioides, in alpine
heathland, and from trunks of Athrotaxis
cupressoides in open montane rainforest.
Specimens examined.Tasmania: unnamed lake, 1·5
km W of Chalice Lake, 41°53'S 146°08'E, 1180 m,
1999, G. Kantvilas 66/99 (HO); Mt Wedge, 42°51'S
146°18'E, 1140 m, 1981, G. Kantvilas 880/81 (HO).
Mycoblastus dissimulans (Nyl.) Zahlbr.
Cat. lich. univ. 4: 3 (1926) —Lecidea dissimulans Nyl.,
Ann. Sci. Nat., sér. 4 (Bot.) 3: 167 (1855); type: Chile,
ad cortices arborum, C. Gay (H-NYL 10905!—
lectotype, here designated).
Lecidea concordans Nyl., Lich. Novae Zealandiae: 108
(1888); type: New Zealand, Greymouth, 1886, Richard
Helms 236 (H-NYL 10903! —holotype).
?Mycoblastus hypomelinus (Stirt.) Müll. Arg., Bull.
Herb. Boissier 2, App. 1: 57 (1894) —Lecidea hypomelina
Stirton, Proc. Phil. Soc. Glasgow 10: 305 (1877); type:
New Zealand, Wellington, J. Buchanan (GLAM—
Arthonia diaphora Stirt., Proc. Phil. Soc. Glasgow 10:
301 (1877); type: New Zealand, near Wellington,
August 1876, J. Buchanan (GLAM—lectotype, ﬁde
Galloway 1985; BM! —isolectotype).
(Figs 1B, 2B, 4E)
Thallus whitish, cream or dull olive-grey, in
part blue-grey due to the underlying prothal-
lus, or sometimes very thin and ± inapparent,
25–250(−500) µm thick, matt to rather waxy
and glossy, ± continuous and smooth to
rather deeply cracked and areolate, at times
rather warty-papillate, sometimes becoming
abraded and scurfy but never sorediate; cor-
tex absent; medulla white; photobiont cells
globose, 5–10 µm diam.
Apothecia 0·3–1·5(−1·8) mm diam., scat-
tered or crowded and conﬂuent, superﬁcial,
basally contricted, strongly convex to sub-
globose, black to blue-black, rarely pale to
reddish brown or piebald, smooth and
glosssy or uneven and scabrid, sometimes
excavate, occasionally (when very young)
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 167
with a thin proper margin but soon becom-
ing immarginate. Exciple in section mostly
excluded, in young apothecia 20–80 µm
thick, hyaline to pale yellowish, intensifying
yellowish in K and N, mostly also with
patches of fucatus-violet and/or cinereorufa-
green pigments, composed of radiating,
branched and anastomosing, heavily conglu-
tinated hyphae 1–3 µm thick. Hypothecium
90–200(–350) µm thick, hyaline to pale yel-
lowish brown, intensifying yellowish in K
and N, inspersed with crystals, granules and
oil droplets 3–10 µm wide that partly dissolve
in K or N, subtended by a yellowish to orange
subhypothecium, usually K+ yellow to
yellow-orange, N+ deep yellow, typically
also with a cinereorufa-green band. Hy-
menium 100–150(–180) µm thick, likewise
densely inspersed with crystals, granules and
oil droplets to 20 µm diam., hyaline to pale
yellowish brown, in the upper part usually
continuously or patchily pigmented with
cinereorufa-green and/or fucatus-violet, with
the pigments sometimes diffusing down
between the asci. Asci 2-spored, approxi-
mating the Biatora- or Lecidella-types, 86–
120(–147) ×(29–)34–50 µm. Paraphyses
numerous, highly branched, separating
rather well in K, relatively stout, unevenly
1–4 µm thick; apices rounded, sometimes
enlarged to 5 µm thick, at times developing
a cell-like, gelatinous cap, 5–20 µm wide.
Ascospores simple or very rarely 1-septate
(usually ± spuriously), ovate to broadly ellip-
soid, 36–43·1–56(–61) ×19–24·5–34(–36)
µm; wall 1·5–5(–7) µm thick.
Pycnidia immersed, black, speck-like;
conidia bacilliform 5–6·5 ×1 µm.
Chemistry. Perlatolic acid, hyperperlatolic
acid (±), caperatic acid (±), norcaperatic
acid (±), rangiformic acid (±), norrangifor-
mic acid (±) and/or roccellic acid (±); thallus
K−, C−, KC−, P−, UV+ whitish.
Notes on the type specimens. There are two
specimens collected by Claude Gay of
Lecidea dissimulans Nyl. in the Nylander
Herbarium in Helsinki; both are very
fragmentary. One (no. 70905) contains
perlatolic acid and is here designated the
lectotype. The second (no. 70904) has
apothecia typical of the species as deﬁned in
this paper, but the thallus contains atranorin
and lichesterinic acid. This is an unusual
chemistry and may derive from the thallus of
an adjacent or underlying lichen.
The type specimen of Lecidea hypomelina
Stirt. could not be located in either GLAM
or BM, despite Galloway (1985) citing it as
being in the former repository. The syn-
onymy given above is therefore tentative
and is based on interpretation of the compre-
hensive description of this taxon by
Galloway (1985) and specimens under that
name in MSC determined by H. A. Imshaug.
It is curious that none of the type speci-
mens of the four taxa synonymized above,
nor any previously published description,
makes any reference to the presence of
fucatus-violet pigment, despite this pigment
occurring rather frequently, albeit mostly in
collections from Tasmania.
Remarks. Mycoblastus dissimulans is an
extremely variable species, both with respect
to the morphology of the thallus (thick or
thin; smooth or variously uneven) and the
intensity and nature of the pigmentation of
its apothecia. Specimens from soil may be
particularly variable, with the thallus some-
times consisting of dispersed granules,
areoles or papillae over an effuse, rather
gelatinous prothallus. Such specimens have
been distributed by MSC under the unpub-
lished herbarium name ‘Mycoblastus falk-
landicus Imsh.’ The species is characterized
by the combination of an esorediate thallus
that invariably contains perlatolic acid, the
presence of either or both cinereorufa-green
and fucatus violet pigments in the apothecia,
and by the two-spored asci. In some respects,
it is the non-sorediate counterpart of M.
coniophorus, although fucatus-violet pigment
has never been observed in that species. The
thallus of M. dissimulans may become very
abraded and scurfy, exposing the white
medulla, but it is never truly sorediate. It is
also related to M. bryophilus (see above),
which is truly sorediate. Also superﬁcially
similar is the Northern Hemisphere’sM.
afﬁnis, but that species has Mycoblastus-type
168 THE LICHENOLOGIST Vol. 41
asci and differs chemically, containing atra-
norin and planaic acid. In the New Guinea
highlands, there is a superﬁcially similar,
undescribed species that is identical to M.
dissimulans in every respect, except that it has
incrementally larger ascospores that occur up
to 4 per ascus.
Mycoblastus dissimulans has been a most
enigmantic species to delimit. The diagnostic
apothecial pigments can vary dramatically.
In the extensive South American, Juan
Fernandez and New Zealand material
examined, no fucatus-violet pigment was
detected, yet in specimens from Tasmania
and the subantarctic islands, this pigment is
commonly present and mostly (at least in
Tasmania) dominant. Indeed, it is the
Tasmanian material that caused the greatest
difﬁculties, and specimens with no pigmen-
tation at all, with only cinereorufa-green or
only fucatus-violet, or with cinereorufa-
green in various relative concentrations
together with fucatus-violet, were all
observed. In general, the most heavily pig-
mented apothecia are either the largest (and
presumably oldest) or those from specimens
growing in exposed habitats. Similarly the
degree of inspersion with crystals and oil
droplets also varies greatly, and is also prob-
ably related to age or habitat. The concen-
tration of fucatus-violet tends to increase
with exposure, but there seems no consistent
pattern, and every hypothesis formed during
this study needed to be discounted with
almost every succeeding observation.
Perlatolic acid is the constant, major sub-
stance present in all specimens, but any com-
bination of a range of fatty acids (see above) is
usually also present. No pattern in the occur-
rence of the fatty acids could be discerned,
and these appear to offer no assistance in
delimiting any taxonomic groups. After large
numbers of repeated extractions, I suspect
that the concentration of these compounds
may even vary across a single thallus.
Distribution and ecology. Mycoblastus dis-
simulans is very widespread in the cool tem-
perate Southern Hemisphere and, in this
study, has been recorded from Juan
Fernandez, southern Chile, New Zealand,
Auckland Island, Campbell Island, the
Falkland Islands, Staten Island and
Tasmania. It is a common epiphyte in forest
and scrub, occurring on a very wide range of
host plants and bark types. In treeless envi-
ronments, such as alpine altitudes or the high
latitudes, it may often occur on peaty turf or
terricolous bryophytes and herbs.
In Tasmania, M. dissimulans is one of the
most common epiphytic crustose lichens in
higher rainfall areas in rainforest, scrub and
heathland, especially in western parts of the
island. It inhabits a wide range of bark types
and host species, although canopy branches
and young limbs of hosts such as Nothofagus
cunninghamii, N. gunni, Phyllocladus asplenii-
folius and Anodopetalum biglandulosum appear
to be particularly favoured. The species is
typically a component of a very rich
assemblage of crustose lichens that includes
Coccotrema cucurbitula, Austroblastenia pauci-
septata, Leiorreuma exaltata (Mont. & Bosch)
Staiger, Mycoblastus coniophorus, Pertusaria
novaezelandiae, Megalospora lopadioides
Sipman, Miltidea ceroplasta (C. Bab.) D. J.
Galloway & Hafellner and Thelotrema
Selected specimens examined. Falkland Islands: East
Falklands: cliffs on rock dome at summit of Mt Kent,
[450 m], 1968, H. A. Imshaug 40452 & R. C. Harris
(HO, MSC); summit ridge of N peak of Two Sisters,
1968, H. A. Imshaug 40403 & R. C. Harris (MSC).—
Argentina: Isla de los Estados: Puerto Vancouver, sum-
mit ridge E of Monte Tres Puntas, 54°47'S 64°06'E,
1971, H. A. Imshaug 52217 & K. Ohlsson (MSC); Puerto
Cook, summit of peak S of Monte Orejas de Burro,
54°45'S 64°01'W, 470 m, 1971, H. A. Imshaug 51595 &
K. Ohlsson (MSC); Bahia Capitan Canepa, 54°49'S
64°27'E, 120 m,. 1971, H. A. Imshaug 53126 & K.
Ohlsson (MSC); Puerto Celular, summit of mountain on
S side of cove, 54°48'S 64°19'W, 430 m, 1971, H. A.
Imshaug 52469 & K. Ohlsson (MSC); Puerto Roca,
54°45'S 64°12'W, 360 m, 1971, H. A. Imshaug 51168 &
K. Ohlsson (HO, MSC).—Chile: Brunswick Peninsula:
Puerto Cutter, 1967, H. A. Imshaug 39367 & R. C.
Harris (MSC). Prov. Magellanes: W side of Puerto Alert,
49°52'S 75°14'W, 1969, H. A. Imshaug 43887 & K.
Ohlsson (MSC); B. Fortescue, 53°42'S 72°01'W, 1969,
H. A. Imshaug 44968 & K. Ohlsson (MSC); NE side of
Puerto Gallant, 53°41'S 72°00'W, 1969, H. A. Imshaug
45114 & K. Ohlsson (HO, MSC); E shore of Puerto
Bueno, 1969, H. A. Imshaug 44562 & K. Ohlsson
(MSC); E side of Isla Pilot, 1969, H. A. Imshaug 43754
&K. Ohlsson (MSC).—Juan Fernandez: Mas a Tierra:
Cordón Salsipudes, 670 m, 1965, H. A. Imshaug
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 169
38096B (MSC); El Yunque, 430 m, 1965, H. A.
Imshaug 37790 (MSC); Cordón Salsipuedes, 670 m,
1965, H. A. Imshaug 38111 (MSC); Portezuelo de
Villagra, ridge towards El Pirámide, 580 m, 1965, H. A.
Imshaug 38275B (MSC); main ridge near junction with
Cordón Escarpado, 640 m, 1965, H. A. Imshaug 37967
(MSC); Valle Colonial, trail to Portezuelo de Villagra,
450 m, H. A. Imshaug 37679I (MSC). Mas a Fuera: Los
Innocentes, 1000 m, 1965, H. A. Imshaug 37430
(MSC); Quebrada de las Vacas, near summit of ridge on
N side of canyon beneath Cordón Barril, 700 m,1965,
H. A. Imshaug 37315G (MSC).—New Zealand:
Campbell Island: head of Garden Cove toward Filhol
Peak, 1970, R. C. Harris 5190 (MSC); near western
summit of Mt Lyall, [390 m], 1969, R. C. Harris 4781
(HO, MSC); between Garden Cove and Mt Filhol,
1970, H. A. Imshaug 46815 (HO, CHR, MSC); summit
of Mt Honey, 1969, H. A. Imshaug 46415 (HO, MSC);
summit of Filhol Peak, 1970, R. C. Harris 5072 (HO,
MSC). Auckland Island: between Smith Harbour and
Norman Inlet, 1973, H. A. Imshaug 57226 (MSC); head
of Tandy Inlet, 1973, H. A. Imshaug 57571 (MSC);
between Smith Harbour and Norman Inlet, 1973, H. A.
Imshaug 57226 (MSC); head of Tandy Inlet, 1973,
H. A. Imshaug 57571 (MSC). South Island: just below
Mount Arthur Hut, 41°12'S 172°43'E, 1260 m, 1993,
L. Tibell 19624 (AK, HO); near start of Punchbowl
Creek Track, Arthurs Pass NP, 1971, H. A. Imshaug
48199 (MSC); Buller Gorge, 1972, H. A. Imshaug
55814 (MSC).—Tasmania: Lake Skinner, 42°56'S
146°41'E, 970 m, 2008, G. Kantvilas 198/08 (HO);
Algonkian Mountain, 42°24'S 146°03'E, 940 m, 1990,
G. Kantvilas 136/90 (HO); W of Tahune Bridge,
43°06'S 146°40'E, 90 m, 2003, G. Kantvilas 64/03
(HO); Mt Sprent, 42°47'S 145°59'E, 500 m 1987, G.
Kantvilas 15/87 (hb. Veˇzda, HO); Flinders Island,
Strzelecki Peaks, 40°12'S 148°04'E, 750 m, 1997, G.
Kantvilas 295/97 (HO); c. 3 km S of Teepookana,
42°13'S 145°26'E, 220 m, 1990, G. Kantvilas 652/90
(HO); Weindorfers Forest, 41°38'S 145°56'E, 960 m,
1984, G. Kantvilas 371/84 & P. James (BM, HO); Mt
McCutcheon helipad, 42°22'S 145°40'E, 400 m, 1984,
G. Kantvilas 257/84 & P. James (BM, HO); Five Road,
42°44'S, 146°25'E, 450 m, 1981, G. Kantvilas 267/81
(BM, HO); Tornado Ridge, Cape Pillar, 43°11'S
147°57'E, 280 m, 2005, G. Kantvilas 260/05 (HO);
Mt. Freycinet, 42°13'S 148°18'E, 600 m, 1995, G.
Kantvilas 147/95 (HO).
Mycoblastus kalioruber Kantvilas sp.
Thallo esorediato, typice olivaceo-griseo vel olivaceo-
brunneo, acida perlatolicum roccellicumque continenti,
ascis bisporis, ascosporis 36–68 µm longis, 18–36 µm
latis, et praecipue pigmento aurantiaco, kalio rubro, in
hypothecio subhypothecioque locato recognita.
Typus: Tasmania, Pine Lake, 41°45'S 146°42'E,
1200 m alt., on Athrotaxis cupressoides in open montane
rainforest, 4 June 1989, G. Kantvilas 191/89 (HO—
Thallus dull olive-grey to olive-brown,
rather waxy when well-developed, somewhat
patchy to continuous, smooth to uneven to
verruculose, becoming deeply cracked and
areolate, sometimes rather ﬂaky, rarely
scurfy-abraded but never sorediate, mostly
80–500 µm thick, rarely very thin, bluish grey
to inapparent; cortex absent; medulla mostly
white but sometimes locally yellow-orange
beneath the apothecia; photobiont cells
± subglobose, 6–14 µm diam.
Apothecia 0·5–1·6 mm diam., roundish
and scattered, or crowded, deformed rhom-
boid and conﬂuent, usually convex to sub-
globose and markedly constricted at the
base, occasionally rather adnate and sunken
into the thallus, black, rarely brownish, dark
bluish green or piebald, smooth and glossy
when young, becoming uneven, scabrid
or in part eroded, immarginate. Exciple in
section very soon excluded, 20–50 µm thick,
colourless to pale yellowish within and
with a diffuse band of cinereorufa-green
pigment at the outer edge, composed of
branched and anastomosing, radiating hy-
phae 2·5–3 mm thick. Hypothecium 120–
250 µm thick, vivid yellow, K+ intense
blood-red, N+ yellow-orange, very densely
inspersed with minute granules that mostly
dissolve in KOH and golden oil droplets to
18 µm diam. Hymenium 90–160 µm thick,
not or only sparsly inspersed with granules
and oil droplets, in the uppermost part
thickly to diffusely cinereorufa-green pig-
mented, in the lower part colourless to pale
yellow, K+ ﬂeetingly golden yellow, colour-
less in N. Asci 2-spored, approximating the
Biatora- or Lecidella-types, broadly ellipsoid,
80–110 ×30–50 µm. Paraphyses numer-
ous, highly branched and anastomosing,
separating readily in K, rather knobbly to
moniliform, unevenly 2·5–4·5 µm thick;
apices not expanded, colourless or with pale
cinereorufa-green caps. Ascospores ovate to
broadly ellipsoid, simple, rarely spuriously
1-septate when over-mature, (36–)40–49·5–
60(–68) ×(18–)20–28·1–36 µm; wall 2–5 µm
Pycnidia not found.
170 THE LICHENOLOGIST Vol. 41
Chemistry. Perlatolic acid, hyperperlatolic
acid (minor) and roccellic acid; thallus K−,
KC−, C−, P± very faint orange, UV+ white.
The major pigment in and beneath the
apothecia is hybocarpone (determined by
Etymology. The speciﬁc epithet refers to
the characteristic pigment which turns a vivid
blood-red (ruber) in KOH (kalium).
Remarks. With its esorediate thallus con-
taining perlatolic and fatty acids, and two-
spored, Biatora- or Lecidella-type asci, M.
kalioruber is most closely related to M. dis-
simulans, and is recognized unequivocally by
the presence of the orange-yellow sub-
hypothecial pigment (hybocarpone) that
turns vivid red with the addition of KOH.
So strong is the reaction that in microscope
sections, the red coloration ‘bleeds’ into the
entire preparation. Even macroscopically,
the pigment is often visible beneath the
apothecia. The pigment is not the same as
that found in the Northern Hemisphere’sM.
sanguinarius, nor in the Tasmanian taxon, M.
sanguinarioides, both of which differ further
by having single-spored, Mycoblastus-type
asci and larger ascospores. A further differ-
ence between M. kalioruber and M. dissimu-
lans is that whereas the former contains only
cinereorufa-green pigment in the epithecium
and hymenium, the latter contains either or
both of fucatus-violet and cinereorufa-green
pigments. Although anatomical sections are
required to unequivocally separate these
species, M. kalioruber tends to have a thicker
thallus that is olive-tinged; the thallus of M.
dissimulans tends to be whitish to dull grey
and generally thinner.
Distribution and ecology. Mycoblastus kal-
ioruber is known only from Tasmania, where
it occurs in heathland, scrub, eucalypt forest
and rainforest. Although it is widespread
across higher rainfall areas of the island, in-
cluding coastal peaks of the East Coast and
Bass Strait islands, it is most abundant in
subalpine and alpine vegetation. A very typi-
cal vegetation type for the species is open
montane rainforest where it inhabits the bark
and wood of the dominant tree, Athrotaxis
cupressoides, as well as the trunks and twigs
of various undershrubs. There it grows in a
rich lichen association that includes Austro-
blastenia pauciseptata,Coccotrema cucurbitula,
Hypogymnia lugubris (Pers.) Krog, Leiﬁdium
tenerum (Laurer) Wedin, Megalospora lopa-
dioides,Menegazzia testacea P. James & D. J.
Galloway, M. pertransita,Mycoblastus dis-
simulans,Tasmidella variabilis and Usnea
inermis Motyka. Although primarily an epi-
phyte, it has been collected on quartzitic
rocks and dead wood.
Selected specimens examined.Tasmania: Lake
Osborne Track, Hartz Mountains, 43°13'S 146°45'E,
820 m, 1981, G. Kantvilas 528/81 & P. James (BM,
HO); Lonely Tarns, 42°58'S 146°27'E, 910 m, 2000,
G. Kantvilas 479/00 (HO); Lake Cygnus, 43°08'S
146°14'E, 880 m, 2006, G. Kantvilas 493/06 (HO);
Mt Freycinet, 42°13'S 148°18'E, 600 m, 1995, G.
Kantvilas 146/95 (HO); Lake Emmett, 42°38'S
146°33'E, 980 m, 1990, G. Kantvilas 154/90 (HO);
Lake Mackenzie, 41°41'S 146°23'E, 1100 m, 2005, G.
Kantvilas 354/05 (HO); Mt Sprent, 42°47'S 145°59'E,
500 m, 1987, G. Kantvilas 14/87 (hb. Veˇzda, HO);
Sentinel Range summit, 42°52'S 146°15'E, 880 m,
2007, G. Kantvilas 335/07 (HO); 4 km N of Precipitous
Bluff, 43°25.5'S 146°36.5'E, 730 m, 1990, G. Kantvilas
112/90 (HO). Flinders Island: Mt Strzelecki, 40°12'S
148°05'E, 710 m, 2006, Kantvilas 47/06 (HO).
Mycoblastus leprarioides Kantvilas &
Elix sp. nov.
Species apotheciis destituta, thallo toto granulari-
sorediato, pallide ﬂavido, acida perlatolicum proto-
cetraricumque continenti recognita.
Typus: Australia, Victoria, Mt Baw Baw National
Park, Mt Erica car park, 37°53'S 146°21'E, 1050 m
altitude, on base of eucalypt in cool temperate rainforest,
13 April 2008, G. Kantvilas 185/08 (HO—holotypus;
Thallus pale grey-green, initially composed
of minute (to c. 0·5 mm), inconspicuous
areoles, becoming sorediate and soon con-
sisting entirely of a contiguous, pale yellow-
ish, sorediate crust to c. 1 mm thick; soredia
coarsely granular, with individual granules
to c. 0·08 mm wide; prothallus effuse, ±
gelatinous, pale greyish, evident only in the
youngest, peripheral parts of the thallus;
photobiont cells ± globose, 5–10 µm diam.
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 171
Chemistry. Perlatolic acid (major), pro-
tocetraric acid (minor), virensic acid (trace,
hyperperlatolic acid (trace), anziaic acid
(trace), 4-O-methylolivetoric acid (trace);
thallus K−, KC−, C−, P+ red, UV± whitish.
Etymology. The speciﬁc epithet refers to
the superﬁcial resemblance of this taxon to a
species of Lepraria.
Remarks. This is an enigmatic species,
ascribed tentatively to the genus Mycoblastus
on the basis of its chemical composition
(dominated by perlatolic acid) and chloroc-
coid photobiont that is the same as that
found in other species of the genus. Initially
the thallus is comprised of tiny areoles but
these soon dissolve entirely into coarse sore-
dia. The thallus of M. leprarioides resembles
some forms of M. coniophorus which may also
be entirely granular sorediate, but that taxon
is invariably fertile and lacks protocetraric
acid. No fertile material of M. leprarioides has
been found to date, and in the ﬁeld, it could
easily be mistaken for a coarse species of
A clearly related but chemically different
taxon, also collected in Victoria (Elix 38675),
remains undescribed. Like M. leprarioides, it
has an entirely granular sorediate yellowish
thallus, but it differs chemically in contain-
ing perlatolic acid and hybocarpone. This
chemical composition is similar to that of
Distribution and ecology. The new species is
known only from the type collection. It grew
on thick bark on the buttress of a mature
eucalypt in cool temperate rainforest domi-
nated by Nothofagus cunninghamii.
Also examined: Australia: Victoria: Bemm River
Scenic Reserve, 37°37'30$S 148°53'12$E, 65 m, 2008,
J. A. Elix 38675 (CANB).
Mycoblastus sanguinarioides Kantvilas
AMycoblasto sanguinario thallo tenuiore et acidum
aliphaticum diversum contineti differt.
Typus: Tasmania, Pelion Plains, 1 km W of Pelion
Hut, 41°50'S 146°02'E, 890 m altitude, on eucalypt
stump in Eucalyptus delegatensis open forest, 11 March
1992, G. Kantvilas 267/92 (HO—holotypus; BM—
(Figs 1A, 2A, 4H)
Thallus whitish, continuous, rather waxy,
smooth to verruculose, often becoming
cracked and areolate, 120–250 µm thick, not
sorediate; cortex absent; medulla white
except occasionally with discrete specks of a
red pigment beneath the apothecia, K+
intensifying orange-red, N+ orange-yellow
and slowly dissolving; photobiont cells
globose, 5–11 µm diam.
Apothecia 0·5–2 mm diam., roundish, scat-
tered, superﬁcial, generally rather adnate,
strongly convex to ± hemispherical, glossy,
black, immarginate, occasionally fused in
clusters to 3 mm wide. Exciple in section ±
excluded from the ﬁrst, in very young
apothecia to c. 40 µm thick and heavily
cinereorufa-green pigmented, composed of a
reticulum of short-celled hyphae, typically
with adhering thalline tissue. Hypothecium
60–160 µm thick, colourless, sparsely in-
spersed with scattered oil droplets 6–18(–50)
µm diam. and minute granules that ﬂuoresce
bluish white in polarized light and dissolve in
K, subtended by a pale straw coloured sub-
hypothecium intensifying yellowish in K, un-
changed in N, sometimes with dilute patches
of cinereorufa-green pigment. Hymenium
110–160 µm thick, colourless in the lower
part, very heavily pigmented cinereorufa-
green in the upper part, sparsely inspersed as
the hypothecium. Asci single-spored, of the
Mycoblastus-type, elongate-clavate, 110–130
×40–52 µm. Paraphyses numerous, very
heavily and persistently gelatinized, branched
and anastomosing, 1·5–2 µm thick, of ± even
thickness throughout but with apices some-
times swollen to 3–4 µm and cinereorufa-
green pigmented. Ascospores broadly ellipsoid
to ± oblong, simple, (60–)64–93·1–104 ×
28–46·1–52(–55) µm; wall 5–12 µm thick,
consisting of two distinct layers.
Pycnidia immersed in the thallus, scat-
tered, resembling minute, black, apothecial
initials; conidia bacilliform, 6–8 ×1 µm.
172 THE LICHENOLOGIST Vol. 41
Chemistry. Atranorin and caperatic acid;
thallus K+ faint yellow, KC−, C−, P−, UV−.
The red pigment that is occasionally found
beneath some apothecia appears to be similar
to rhodocladonic acid but differs by fading to
a reddish brown in herbarium specimens.
Etymology. The speciﬁc epithet refers to
the similarity between this taxon and the
Northern Hemisphere M. sanguinarius.
Remarks. Within the austral lichen ﬂora,
this species is readily distinguished from all
other known species of Mycoblastus by its
one-spored, Mycoblastus-type asci, relatively
large ascospores and distinctive chemical
composition; all the other taxa have Biatora-
to Lecidella-type, typically two-spored asci
and contain perlatolic acid, often in combi-
nation with other compounds. The species is
also readily recognizable in the ﬁeld by its
whitish thallus and glossy, black, rather
adnate apothecia. The occurrence of the red-
dish pigment beneath the apothecia is very
sporadic and unstable: relatively few speci-
mens studied display this character, and of
these, many apothecia lack the pigment
anyway (see below).
Separation of the new species from M.
sanguinarius is more problematic, although
well supported by a detailed study of large
numbers of specimens of both taxa. The
thallus of M. sanguinarius is generally thicker,
coarser and more rugose-verrucose. The red-
dish pigments in both species display the
same reactions in K and N but, on the basis
of qualitative observations, appear to be dif-
ferent. In M. sanguinarius, the pigment is
almost always present, more extensive and
persistently bright red. In M. sanguinarioides
on the other hand, the pigment is only occa-
sionally present, and although it appears
bright red in freshly collected specimens, it
soon turns brownish and ultimately fades
altogether. Many specimens of the latter
taxon which, when ﬁrst collected, displayed
distinctive patches of red were found, on
re-examination several years later, to lack
the red pigment almost completely. The key
difference between the two taxa, however,
is the thallus chemistry. Both taxa contain
atranorin, but the accompanying fatty acid is
clearly different, at least when observed in
solvent C of Culberson & Johnson (1982).
Literature concerning M. sanguinarius (e.g.
Watson & James 1992) states that the fatty
acid in that species is caperatic acid but this
does not appear to be the case, and the fatty
acid present appears to have a Rf value more
similar to that of bourgeanic acid. Two speci-
mens from the United Kingdom contained
both bourgeanic and caperatic acids. Myco-
blastus sanguinarioides, however, does contain
caperatic acid, usually together with nor-
caperatic acid; it never contains bourgeanic
Also similar is M. japonicus Müll. Arg.,
which also has single spored, Mycoblastus-
type asci, large, ellipsoid to cylindrical asco-
spores and a bright red sub-apothecial pig-
ment. This species is easily distinguished
from both M. sanguinarius and M. sangui-
narioides by its thallus chemistry which com-
prises atranorin together with fumarproto-
Distribution and ecology.Mycoblastus san-
guinarioides is known from Tasmania where it
is relatively abundant, and from a single
locality in mainland Australia (eastern
Victoria). It typically occurs on the lignin of
various trees, especially Eucalyptus and
Athrotaxis, although it has also been observed
on the bark of Banksia marginata. It grows
mostly in wet forests or in wet heathlands and
grassy woodlands with emergent, mature
trees, usually in open situations on logs.
Not infrequently it has also been observed
on dead wood fallen from the forest canopy.
The species ranges from lowland to alpine
Selected specimens examined.Tasmania: Three
Thumbs, 42°36'S 147°52'E, 480 m, 1989, G. Kantvilas
203/89 (HO); Lake MacKenzie, 41°41'S 146°23'E,
1100 m, 2005, G. Kantvilas 353/05 (HO); Travellers
Rest Lake, 42°03'S 146°15'E, 950 m, 2003, G.
Kantvilas 10/03 (HO); Tanina Bluff, 42°39'S 147°02'E,
890 m, 2005, G. Kantvilas 204/05 (HO); 0·5 km SW of
Narcissus Hut, 42°01'S 146°06'E, 740 m, 2004, G.
Kantvilas 59/04 (HO); c. 7 km E of Lake Leake,
42°01'30$S 147°55'E, 400 m, 1996, G. Kantvilas s.n.
(HO); W of Tomalah Creek, 43°05'S 146°39'E, 270 m,
1999, G. Kantvilas 399/99 (HO).—Australia: Victoria:
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 173
Errinundra Saddle, Rainforest Walk, 37°19'03$S
148°50'19$E, 910 m, 2008, G. Kantvilas 167/08 & J.
Elix (HO, MEL).
Other Southern Hemisphere names
pertaining to Mycoblastus
Several additional Mycoblastus names have
been recorded for the Southern Hemisphere,
but are not treated above. These are dealt
Mycoblastus austroshetlandicus C.W.
Instit. Antarctico Chileno 6: 6 (1965).
The type specimen of this species (in FH)
was examined by A. M. Fryday who deter-
mined it as Farnoldia dissipabilis (Nyl.) Hertel
(A. M. Fryday, pers. comm.). It has eight-
spored asci with relatively small ascospores,
22–25 ×10 µm (Dodge 1965), characters
inconsistent with Mycoblastus.
Mycoblastus hookeri C.W. Dodge
Nova Hedwigia 19: 477 (1970); type: New Zealand,
North Island, Kawakawa, on branch of Trophis, J. D.
Hooker, ex hb. Taylor no. 642 (FH!— holotypus).
This species is based on a specimen of
Megalospora gompholoma (Müll. Arg.)
Sipman subsp. gompholoma.
Mycoblastus implicatus (Stirt.) Müll.
Bull. Herb. Boissier 2, app. 1: 56 (1894) —Lecidea impli-
cata Stirt., Rep. Trans. Glasgow Soc. Field Nat.1: 19
(1873); type not seen.
Galloway (1985) records this as a synonym
of Ochrolechia pallescens (L.) A. Massal., but
did not locate the type specimen. The orig-
inal description (Stirton 1873) refers to
eight-spored asci and pale fruiting bodies,
characters not consistent with Mycoblastus.
Mycoblastus perustus (Nyl.) C. W.
B.A.N.Z.A.R.E. Rep., ser. B, 7: 102 (1948).—Lecidea
perusta Nyl. in Cromb., J. Bot. 13: 334 (1874); type:
Kerguelen, Royal Sound, February 1875, A. E. Eaton
(BM!— lectotypus, ﬁde Hertel 1984).
This widespread austral taxon is now
known as Poeltidea perusta (Nyl.) Hertel &
Hafellner (Rambold 1989).
Mycoblastus stephanodes (Stirt.) C. W.
B.A.N.Z.A.R.E. Rep., ser. B, 7: 103 (1948).—Lecidea
stephanodes Stirt. in Cromb., J. Linn. Soc. Bot. London
16: 221 (1877); type: Kerguelen’s Land, January 1875,
H.N. Mosely (BM!— lectotypus, ﬁde Hertel 1984).
This subantarctic taxon is now known as
Porpidia stephanodes (Nyl.) Hertel.
Salient features of other taxa studied
Mycoblastus afﬁnis (Schaerer) Schauer
Thallus. Grey to greenish grey, usually
thick and verruculose-warty, not sorediate.
Pigments. Cinereorufa-green in the epi-
hymenium, usually appearing very deep
Asci.Mycoblastus-type (?); ascospores (1–)2
per ascus, broadly ellipsoid, (40–)47–70
(100) ×(25–)30–42 µm (James 1971).
Chemistry. Atranorin, planaic acid, roccel-
lic acid (±), norcaperatic acid (±).
Distribution. Temperate Northern Hemi-
Selected specimens examined. USA: Alaska: vicinity of
Juneau, Mt Roberts Trail, 1962, H. A. Imshaug 28226B
(MSC). Washington: near Mt Adams, 1994, G.
Kantvilas & J. Davis s. n. (HO). Oregon: Multnomah
Co., Larch Mt, 1962, S. Sushan sl-1899A (MSC).—
Canada: British Columbia: 15 miles NE of Stillwater, S
of Powell River, 1969, K. E. Ohlsson 997 (MSC).—
Switzerland: Val Bregaglia, 46°20'N 9°34'E, 1999, G.
Kantvilas (HO).—Austria: Klingelbergalm, 1540 m,
1989, R. Türk 8644 (A. Veˇzda: Lich. Sel. Exsicc. 2362)
Mycoblastus alpinus (Fr.) Kernst.
Thallus. Grey, verruculose-warty, soredi-
ate; soredia granular, yellowish, initially in
punctiform soralia, then spreading to ± cover
the entire thallus. Only sterile specimens
seen and hence pigments, asci and asco-
spores not observed.
Chemistry. Atranorin and planaic acid,
with usnic acid in the soralia.
174 THE LICHENOLOGIST Vol. 41
Distribution. Temperate Northern Hemi-
Specimens examined. Poland: Rysianka Mountains
near Sopotnia Wielka, 1250 m, 1970, J. Nowak (Lich.
Pol. Merid. Exsicc. 38) (MEL).—USA: Alaska: Atqasuk,
dry ridge to W of ITEX site, 2001, A. M. Fryday 8096
Mycoblastus caesius (Coppins &
P. James) Tønsberg
Thallus. Grey with a purplish tinge due to
underlying prothallus, areolate-warty, sore-
diate; soredia granular, ± concolorous with
the thallus, scattered.
Pigments. Predominantly cinereorufa-
green in the epihymenium, sometimes with
traces of fucatus-violet, with the combination
appearing very deep indigo bluish.
Asci. Approximating the Biatora- or
Lecidella-types; ascospores 2 per ascus, ovate
to broadly ellipsoid, (30–)40–50 ×20–30
Chemistry. Perlatolic acid, frequently with
traces of several fatty acids.
Distribution. Chieﬂy in temperate oceanic
areas of the Northern Hemisphere.
Selected specimens examined. USA: Washington: E of
La Push, 47°54'N 124°33'W, 30 m, 1994, T. Tønsberg
21265 (BG, HO, MSC); Olympic Peninsula, N of Forks
Village, 47°57.6'N 124°23.5'W, 70–80 m, 1999, T.
Tønsberg 27132 (BG, HO); Olympic Peninsula, SE
of Forks, 1999, T. Tønsberg 27212a (BG); Olympic
Peninsula, between Sappho amd Clallam Bay, 48°11'N
124°13'W, 50 m, 1991, T. Tønsberg 13941 (BG).
Alaska: Tongass Nat. Forest, Mitkof Island, 2001, T.
Tønsberg 30241 (BG).
Mycoblastus fucatus (Stirt.) Zahlbr.
Thallus pale to dark grey or grey-brown,
smooth to areolate-warty, sorediate; soredia
granular, grey-green, in discrete soralia.
Pigments dominated by fucatus-violet in
Asci: not observed; ascospores (1–)2(−3)
per ascus, broadly ellipsoid, (25–)30–
48(−52) ×15–21 µm (James 1971).
Chemistry. Atranorin, fumarprotocetraric
acid (James 1971).
Distribution. Western Europe; a specimen
from China, distributed by A. Veˇzda (Lich.
Rar. Exsicc. 66) is of a different, distinct
Specimens examined.Germany: Regierungsbezirk
Trier, 50°18'N 6°26'E, 590 m, 1999, V. John (H.
Hertel: Lecideaceae Exsicc. 333) (HO); weg von
Aremberg nach Eichbach, 1990, H. T. Lumbsch & G. B.
Feige 7759a (hb. Lumbsch, HO).—Lithuania: Punia
Forest Nature Reserve, 2006, J. Motiejunaite 7625 (BI-
LAS, HO).—Belgium: Membach, 1984, E. Sérusiaux
6771 (HO, LG).
Mycoblastus glabrescens (Nyl.) Zahlbr.
Thallus whitish cream, smooth, esorediate.
Pigments. Cinereorufa-green in the epi-
Asci Mycoblastus-type; ascospores: 1–2 per
ascus, broadly ellipsoid; mature spores not
Chemistry. Atranorin, chloroatranorin,
Distribution. North-western North
When Nylander (1864) introduced this
species in his Prodromus Florae Novo-
Granatensis (ﬁrst published in the previous
year), he was mistaken in believing it origi-
nated from Colombia, South America. The
type specimen was collected by John Scouler,
who was active in the Columbia River area of
the Paciﬁc North-West of the USA (Scouler
1905). The label simply states ‘‘Columbia,
Scouler, ex hb. Hooker’’. The specimen is
a minute fragment with only one intact
apothecium. Whereas the chemistry is
unusual, all other characters suggest that this
name is a synonym of M. afﬁnis. However, no
other specimens examined from the same
region contained nephrosterinic acid, and
instead had the typical M. afﬁnis chemistry of
atranorin and planaic acid.
Specimen examined. [USA] Columbia, J. Scouler
Mycoblastus japonicus Müll. Arg.
Thallus brownish grey, thick, verrucose to
bullate, not sorediate.
2009 Mycoblastus in the Southern Hemisphere—Kantvilas 175
Pigments. Pale olive-green pigment
(possibly very dilute cinereorufa-green) in
epihymenium, bright red pigment (cf. rhodo-
cladonic acid) beneath the apothecia.
Asci. Mycoblastus-type; ascospores 1 per
ascus, broadly ellipsoid to cylidrical, 65–84 ×
26–32 µm (Müller Argoviensis 1891).
Chemistry. Atranorin, fumarprotocetraric
acid; an alternative chemistry, identical to
that of M. sanguinarius, has been recorded by
Huneck & Schmidt (1995).
Specimen examined.Japan: Mt Fuji, 1971, R. Filson
14017 & S. Filson (MEL).
Mycoblastus sanguinarius (L.) Norman
Thallus pale to dark grey, usually thick and
verrucose-papillate, occasionally sorediate.
Pigments. Cinereorufa-green in epi-
hymenium, usually appearing very deep
indigo bluish; bright red pigment (rhodocla-
donic acid) beneath the apothecia.
Asci. Mycoblastus-type; ascospores 1 per
ascus, broadly ellipsoid to cylidrical, 70–100
×25–45 µm (Brodo et al. 2001).
Chemistry. Atranorin, bourgeanic acid (±),
caperatic acid (±).
Distribution. Temperate Northern Hemi-
Selected specimens examined.USA: Washington: near
Mt Adams, 1994, G. Kantvilas & J. Davis s.n. (HO).—
Canada: Quebec: E side of Lac Jean Pere, 47°04'N
76°32'W, 1977, H. A. Imshaug 60308 (HO, MSC).—
Japan: Honshu: fourth steps of Mt Fuji, Subaru Line
Highway, 2100 m, 2004, G. Kantvilas, H. Kashiwadani
& K. Moon s.n. (HO).—Great Britain: Scotland:
Resipole, 1994, G. Kantvilas, B. J. Coppins & A. M.
O’Dare s.n. (HO).—Sweden: c. 10 km NE of
Kramfors, 63°00'N 17°56'E, 170 m, 1984, H. Hertel &
R. Moberg (H. Hertel: Lecid. Exsicc. 129) (HO).
This project was commenced whilst I was a visitor at the
Michigan State University Herbarium (MSC), East
Lansing, Michigan, USA; I gratefully acknowledge sup-
port from US National Science Foundation, Award No.
DBI–9808 735 (L. A. Prather, PI) that permitted me to
visit the herbarium to study H. A. Imshaug’s extensive
lichen collections. In particular, I thank A. M. Fryday for
his hospitality and support throughout this visit, and for
his help in various ways since. S. J. Jarman prepared the
photographs, and compiled and scanned the line draw-
ings for publication. I thank J. Elix for assistance with
chemical analyses and for alerting me to the relevance of
certain Australian collections pertinent to Mycoblastus.
T. Tønsberg and T. Spribille helped improve the manu-
script in the capacity of referees. The assistance of cura-
tors of the herbaria cited in the text is gratefully
acknowledged. In particular, I thank K. Watson
(GLAM) and S. LaGreca (formerly BM) for their
searches for authentic material of M. hypomelinus, T.
Tønsberg for supplying comparative material of M.
caesius, and D. O. Øvstedal and A. Aptroot for other
loans. O. Vitikainen is further acknowledged for assist-
ance with locating and interpreting several literature ref-
erences. I also take this opportunity to acknowledge with
pleasure the role of P. W. James, who as my ﬁrst teacher
and mentor in lichenology many years ago, kindled my
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178 THE LICHENOLOGIST Vol. 41