Antarctic Science 18 (3), 291–301 (2006) © Antarctic Science Ltd Printed in the UK
DOI: 10.1017/S0954102006000344
Lichenicolous fungi described from Antarctica and the subAntarctic islands by Carroll W. Dodge (1895–1988)
DAVID L. HAWKSWORTH1* and TERESA ITURRIAGA2
1
Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, Plaza Ramón y Cajal, Ciudad Universitaria,
ES-28040 Madrid, Spain
2
Departamento Biología de Organismos, Universidad Simón Bolívar, Apartado 89000 Sartenejas, Baruta, Edo. Miranda, Venezuela
*Address for correspondence: MycoNova, The Yellow House, Calle Aguila 12, Colonia La Maliciosa, Mataelpino, ES-28492 Madrid, Spain
myconova@terra.es
Abstract: The identities of the 12 lichenicolous fungi described as new to science from the Antarctic and
sub-Antarctic islands by Carroll W. Dodge between 1938 and 1968 are reassessed on the basis of the original
publications, and where extant the original collections and slides in the Farlow Herbarium and Reference
Library (Harvard University, Cambridge, MA, USA). Typifications are discussed and lectotypes and
neotypes designated as necessary. Only one species is accepted in its original placement (Phacopsis usneae)
and one had already been transferred to an appropriate genus (Didymella cladoniae to Zwackhiomyces). Six
prove to be synonyms of previously described species, and four are combined into other genera here:
Arthonia parmeliae (syn. Diplonaevia parmeliae), Carbonea antarctica (syn. Alectoria antarctica),
Sphaerellothecium buelliae (syn. Orbicula buelliae), and Weddelomyces gasparriniae (syn. Phaeospora
gasparriniae) combs. nov. The generic name Thelidiola is also confirmed as a synonym of Muellerella. In
addition, as a result of earlier misinterpretations of Dodge’s taxa, two new species are described:
Endococcus matzeri, and Zwachkiomyces diederichii spp. nov. Intralichen lichenum is also reported from
the Antarctic for the first time. Descriptions and illustrations are included where appropriate.
Received 4 October 2005, accepted 30 January 2006
Key words: Ascomycota, discomycetes, hyphomycetes, lichens, pyrenomycetes, taxonomy
Introduction
published extensively on those from little-explored regions,
especially Africa, Antarctica, and South America until 1973,
aged 78 years. When he started to become interested in
lichens around 1912–15, there were only two active
lichenologists in the whole of the USA, and they were based
in the southern states. In consequence, he worked in
isolation, and relied largely on literature accounts for his
interpretation of previously published species. Although he
always produced lengthy descriptions including detailed
microscopical observations, his concepts of genera and
species were often idiosyncratic and re-evaluations of the
taxa he described as new are essential. The first author
corresponded with Dodge from 1967–74, mainly with
respect to alectorioid lichens, and was privileged to receive
various collections and slides from him on loan during a
period when he had ceased to communicate with many
lichenologists; this provided valuable insights into his ways
of working. For many years his collections were
deteriorating in a garage at his home, prior to their transfer
to the Farlow Herbarium and Reference Library (Harvard
University, Cambridge, MA, USA; FH) in 1985. A
staggering 75 000 collections were accessioned, including
those of non-lichenized fungi, and after laborious
conservation work they started to become available for
study in the early 1990s. Detailed biographical information,
and a full list of Dodge’s publications, is provided by
Fungi obligately occurring on lichens as parasites, saprobes
or commensals represent a major ecological niche for fungi.
There are over 1500 currently known lichenicolous species,
most of which belong to genera only known from lichen
hosts (Lawrey & Diederich 2003). In view of the
predominance of lichens in the Antarctic (Øvstedal & Smith
2001), numerous lichenicolous fungi are to be expected
growing on them, but while there are scattered reports, this
ecological niche has not yet been examined in depth. A
reappraisal of the already documented biota is a prerequisite
for further research, and this paper seeks to address the
identities of the lichenicolous taxa described as new to
science from the Antarctic and sub-Antarctic islands by
Carroll W. Dodge (1895–1988) over the period 1938–68.
Dodge was an enigmatic Vermont figure who made major
well-respected contributions to medical mycology (Dodge
1936), and hypogeous fungi (e.g. Dodge & Zeller 1934). He
held positions at Harvard University (1921–31) and the
Missouri Botanical Garden (1931–63), and as a Research
Professor in the University of Vermont (1963 on) after his
retirement; he also taught courses on medical mycology in
different South American countries from 1934–60.
However, he also had an absorbing interest in lichens which
went back to his undergraduate days, and these became his
main preoccupation after his retirement in 1963. He
291
http://journals.cambridge.org
Downloaded: 11 Dec 2013
IP address: 202.75.16.18
292
D.L. HAWKSWORTH & T. ITURRIAGA
Fig. 1. Carbonea antarctica.
Reproduction of the published
illustration by Dodge & Baker
1938: pl. 51, figs 209–216.
Fig. 209, habit sketch, x 6;
fig. 210, detail of thallus ends
with apothecium, x 35; fig. 211,
section of apothecium, x 55;
fig. 212, development of ascus
through maturity, x 1100;
fig. 213, paraphysis, x 1100;
fig. 214, mature spores, x 1100;
fig. 215a, hypha from medulla,
x 1100; fig. 215b, detail of cortex,
x 1100; and fig. 216, crosssection of thallus, x 177.
Rudolph (1990).
With respect to the lichens he described as new from
Antarctica, their identities have now mostly been resolved
based on a re-examination of the original collections (Hale
1987, Castello & Nimis 1995, Øvstedal & Smith 2001).
However, with his broad mycological background, it was
perhaps not surprising that he was also interested in the
fungi which grow on lichens. At the same time, he also
described as novel “lichens” some lichens which were
actually infected by a lichenicolous fungus. Hawksworth (in
Pegler et al. 1980) endeavoured to make some
reassessments of his Antarctic lichenicolous species, but at
that time Dodge’s material was not available for study.
This contribution provides a fresh evaluation of all the
lichenicolous fungi described from Antarctica and the subAntarctic islands by Dodge, whether as lichenicolous fungi
or erroneously as lichens, based on the original descriptions
and illustrations and where extant the original collections
and microscope slides now preserved in FH.
For more general information on the biology, ecology,
and systematics of lichenicolous fungi, see Lawrey &
Diederich (2003) and Hawksworth (2003, 2004).
Systematics
(1) Alectoria antarctica C.W. Dodge & G.E. Baker, Ann.
Mo. Bot. Gdn 25: 599 (1938).
Type: Marie Byrd Land, Edsel Ford Range, Mt Rea-Cooper,
on Pseudephebe minuscula, 1934, P. Siple, F.A. Wade, S.
Corey & O.D. Stancliff R-1 (FH – slide with gummed label,
lichenicolous fungus element – lectotypus hic designatus).
≡ Carbonea antarctica (C.W. Dodge & G.E. Baker) D.
Hawksw. & Iturr., comb. nov.
(Fig. 1)
http://journals.cambridge.org
Downloaded: 11 Dec 2013
Lamb (1964, p. 28) reports that he was sent a slide of this
taxon by Dodge, along with two other “authentic
specimens” from the same locality, but that only the slide
had ascomata and the other specimens were typical of the
species now called Pseudephebe minuscula. He considered
the ascomata either to be abnormal ones of the lichen or of a
“parasitic or parasymbiotic Ascomycete (perhaps a species
of Nesolechia)” and while listing the name as a synonym of
the lichen, as had Murrray (1963), no formal
lectotypification was made. Dodge also sent the first author
a single slide (perhaps the same one) on loan in 1967, from
which it was evident that the ascomata represented a
lichenicolous fungus. Three slides are now present in FH,
one with a gummed label giving abbreviated type locality
information and labelled “Rea-1”, and the two others with
“type” (also marked “Mt Rea. 7”) and “type 1” (also marked
“Mt Rea-1”) written on the glass, all in Dodge’s hand. The
slide labelled “type 1” only has microtome sections of the
thallus and none of the ascomata. In the absence of a
specimen bearing ascomata from which the slides could
have been made, we consequently designate the
lichenicolous element on the slide with the gummed label as
lectotype for the name here as that has the best preparations
showing the exciple structure and paraphyses as well as the
“Rea-1” annotation. We selected the lichenicolous fungus as
lectotype rather than the host lichen as it was the characters
of the fungus that Dodge regarded as distinctive in lacking a
thalline exciple (“ampithecium”) to the extent that “it might
be considered the type of a new genus and family in the
lecideine series of families” (Dodge & Baker 1938, p. 598).
As pointed out by Lamb, the ascomata definitely appear
to be lecideine and have a well-developed dark brown
exciple, without any trace of algae to suggest the thalline
exciple typical of Pseudephebe was present. This is
supported by the published illustrations of the apothecia and
IP address: 202.75.16.18
293
LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE
the section (Fig. 1, figs 210 & 211). Our studies of the slides
show the exciple to be carbonaceous and composed of
compacted angular thick-walled pseudoparenchymatous
cells 6–10 x 4–8 μm diameter, with walls 2.5–3 μm thick
and a small lumen, forming a textura angularis to textura
globulosa. The asci are lecanoralean, 38–35 x 7.5–11 μm,
and the ascospores are hyaline, narrowly ellipsoid,
somewhat curved in lateral view, with one end frequently
broader than the other, and 6–8 x 3–4.5 μm – much
narrower and more ellipsoid than in P. minuscula where
they measure 6.7–9 x 7.7–8 μm (Brodo & Hawksworth
1977 p. 141). Further, the paraphyses are repeatedly
anastomosed and with lumina only 1–1.5 μm thick. These
features all indicate that the ascomata are not deformed ones
of the host lichen, but of a lichenicolous fungus that is most
appropriately placed in Carbonea (Triebel 1989), a genus
including both lichen-forming and lichenicolous species; a
formal transfer to that genus is therefore made here.
Carbonea antarctica differs from the two other
recognized Antarctic species (Øvstedal & Smith 2001) and
others described in the genus in the sizes of the ascospores,
as well as being the first to be recognized growing on a
macrolichen. These authors also mention, but do not
illustrate or describe in detail an unnamed “Lecanora sp. D”
on Pseudephebe minuscula which we suspect might be the
same species; although placement in that genus suggests a
thalline exciple including algal cells was present, and the
asci and ascospores were given as c. 20 x 8 μm and c. 8 x
6 μm, respectively.
We also noted in the sections mounted on the lectotype
slide hyphae immersed in the hymenium which evidently
represent the hyphomycete Intralichen lichenum
(Diederich) D. Hawksw. & M.S. Cole (Hawksworth & Cole
2002) which appears to be the first report of this species
from Antarctica.
(2) Didymella cladoniae C.W. Dodge, BANZARE 1929–31,
series B, 7: 262 (1948).
Type: Macquarie Island, north end, Station 81, on
“Cladonia mawsonii” [C. cervicornis subsp. mawsonii],
BANZARE B540-1 (FH- holotype [two slides]).
≡ Zwackhiomyces cladoniae (C.W. Dodge) Diederich, in
Aptroot et al., Bibltheca Lich. 64: 210 (1997).
(Fig. 2)
Fig. 2. Zwackhiomyces cladoniae (FH - holotype). a. mature
ascospores, b. maturing ascus and c. anastomosing
paraphysoids. Bar = 10 μm.
http://journals.cambridge.org
Downloaded: 11 Dec 2013
Ascomata perithecioid, black, 300–425 μm diameter; walls
dark brown, pseudoparenchymatous, textura angularis,
individidual cells 4.5–19 x 4.5–15.5 μm, cell walls c. 2 μm
thick. Hamathecium persistent, composed of abundant
branched and anastomosting interascal filaments, width
rather uneven but mainly 0.5–1 μm thick. Asci bitunicate in
structure, subcylindrical, 70–80 x 8–10 μm, 6(–?8) spored.
Ascospores almost uniseriately arranged, slightly
overlapping, broadly fusiform, apices generally somewhat
pointed, 1-septate, hyaline, coarsely verrucose, the warts in
a gelatinous sheath, 12–19 x 3–5 μm.
Two slides were located in FH which have abbreviated
locality details and the collection number “B540-1”, but not
the specimen from which these were made. The two slides
are collectively treated as the holotype of Dodge’s name
here as they were clearly made from and so are part of the
original collection. As the slides had only squash mounts,
the above description is not as full as would have been the
case with sections or the original specimen.
The specimen is indicated as not seen by Grube &
Hafellner (1990), and nor was it located by Aptroot et al.
(1997) when the species was transferred to Zwackhiomyces.
Aptroot et al. examined a fungus on Cladonia yunnana
collected on Mount Wilhelm in Papua New Guinea which
they considered to be conspecific with Dodge’s taxon on the
basis of the original description. However, the perithecia in
their material were 75–125(–150) μm diameter, but could
reach 170 μm in water, and the asci were shorter (45–63 μm
tall); most other measurements they report were very close
to those published by Dodge. Sadly, in this case Diederich
IP address: 202.75.16.18
294
D.L. HAWKSWORTH & T. ITURRIAGA
(in Aptroot et al. 1997) was misled as Dodge´s published
measurements of the ascospores, given as “10–11 x 4–
5 μm”, uncharacteristically proved to be inaccurate (see
above). The Papua New Guinea material, with substantially
shorter asci and smaller ascospores evidently represents a
separate species, for which the name Zwackhiomyces
diederichii is introduced here.
Zwackhiomyces diederichii D. Hawksw. & Iturr., sp. nov.
and wider, about 15 x 4 μm” and the range given by
Hawksworth (in Pegler et al. 1980) is (12–)13–15(–16) x
(4–)4.5–6(–6.5) μm. The species is evidently frequent on
lobate and subfruticose species of Caloplaca species and
also on Xanthoria elegans in the Antarctic region and on the
subantarctic islands (Pegler et al. 1980).
(4) Didymosphaeria macquariensis C.W. Dodge,
BANZARE 1929–31, series B, 7: 263 (1948).
Ascomata semi-erumpescentia, 75–125(–150) μm in
diametro. Asci cylindrici, 45–63 x 6.5–7.5 μm, 6–8 spori.
Hamathecium ex hyphis ramosis et anastomosantibus, ca
1.5 μm latis. Ascosporae ellipsoideae vel soleiformes, cum
apicibus attentuatis, verruculosae, 10–12.5 x 3.5–4 μm.
Type: Macquarie Island, Featherbed Flat, on “Pyrenodesmia
inclinans”, BANZARE B533-7 (holotype – not located).
Typus: Papua New Guinea, Simbu, Mount Wilhelm,
Pindaunde Valley, near the hut, c. 3600 m, on Cladonia
yunnana, 1992, A. Aptroot 23663 (herb. Diederich –
holotypus; herb. Aptroot – isotypus).
This species was stated to differ from D. kuttlingeriae in the
“larger, thicker-walled perithecia, more numerous and
broader asci and slightly larger ascospores” (Dodge 1948,
p. 264). However, the perithecium size was not given in the
description, and the wall thickness can reach the c. 20 μm
cited near the ostiole in P. rugulosarium. Further, the
ascospores were given as 15–16 x 4–5 μm which is within
the range of the species (see under D. kuttlingeriae above).
However, the ascus size was given as “c. 35 x 15 μm”
which must either be an error or based on a very immature
ascus; were this correct, it is inconceivable that it could
accommodate the eight spores cited. On the basis of the
information available in the absence of any original
material, we see no reason not to treat the name as a
synonym of P. rugulosarium as suggested by Hawksworth
(in Pegler et al. 1980).
For a fuller description and illustrations of the asci and
ascospores, see Aptroot et al. (1997, 210–211).
(3) Didymosphaeria kuttlingeriae C.W. Dodge, BANZARE
1929–31, series B, 7: 263 (1948).
Type: Crozet Archipelago, Possession Island, American
Bay, on “Kuttlingeria crozetica” [? Xanthoria elegans],
BANZARE B20-22 (holotype – not located).
= Polycoccum rugulosarium (Linds.) D. Hawksw., in Pegler
et al., Kew Bull. 35: 513 (1980).
Also mentioned in the original account from “Blastenia
johnstonii” in the type locality (B20-19), and on
“B. keroplasta var. athallina” from Kerguelen, Mount
Wyville Thompson (B246-22). None of the three specimens
could be located in FH, but the very full description agrees
in almost all details with that of Polycoccum rugulosarium.
For example, the ascospores were given as “dark brown,
2-celled, long ellipsoid to oblong, upper cell slightly longer
= Polycoccum rugulosarium (Linds.) D. Hawksw., in Pegler
et al., Kew Bull. 35: 513 (1980).
(5) Diplonaevia parmeliae C.W. Dodge & G.E. Baker, Ann.
Mo. Bot. Gdn 25: 661 (1938).
Type: Marie Byrd Land, Edsel Ford Range, Skua Gull Peak,
on “Parmelia variolosa” [? Physcia caesia], P. Siple & S.
Corey 72W-3 (holotype not located).
= Arthonia parmeliae (C.W. Dodge & G.E. Baker) D.
Fig. 3. Arthonia parmeliae.
Reproduction of the published
illustration by Dodge & Baker
1938: pl. 62, figs 396–400. Fig.
396a, detail of hyphae of host, x
1100; fig. 396b, detail of hyphae of
parasite, x 1100; fig. 397,
paraphysis, x 1100; fig. 398, mature
spores, x 1100; fig. 399, section of
apothecium, the host surface
indicated by dotted lines, x 385;
and fig. 400, development of ascus,
x 1100.
http://journals.cambridge.org
Downloaded: 11 Dec 2013
IP address: 202.75.16.18
295
LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE
Hawksw. & Iturr., comb. nov.
(Fig. 3)
Castello & Nimis (1995) stated that they did not locate the
type of the name Parmelia variolosa, a species name
introduced by Dodge & Baker (1938), but indicated that it
was Physcia caesia on the basis of a specimen they said was
an isotype numbered “Siple et al. R-3”. However, no “R-3”
is mentioned in the original account, even amongst several
paratypes, so the true status of the specimen they studied is
unclear. The holotype was stated to be Siple et al. R-1, and
we were able to examine two packets with this number in
FH (Marie Byrd Land, Mt Rea-Cooper, 77º07'S 145º30'W,
on coarse-grained leucogranite, 1934, P. Siple, F.A. Wade, S.
Corey & O.D. Stancliff R-1); we consider the packet with an
inner envelope name in ink in Dodge’s hand to be the
holotype and the other an isotype. An annotation slip
indicates that the holotype was examined by Mason E. Hale
in 1965, and intriguingly both the holotype and isotype
packets have annotation slips by Castello & Nimis from
1994. We confirm that both specimens represent Physcia
caesia.
We found no lichenicolous fungus in the packets of “R-1”.
Diplonaevia parmeliae was described as forming black
convex emarginate ascomata up to 150 μm diameter, and
having interascal filaments that were straight with slightly
enlarged and darkened tips, asci 21–24 x 8–10, and hyaline
“apparently 2-celled” ascospores somewhat pointed at one
end and 7–8 x 2–2.5 μm. This species is illustrated by
Dodge & Baker (1938, pl. 62, figs 396–400), whose
drawing is reproduced here as Fig. 3, and shows a vertical
section with a dark hypothecium and a thick carbonaceous
epithecium arching over the hymenium; in addition,
ramifying brown hyphae are seen to penetrate rhizoid-like
into the host tissue, and the ascospores figured are narrowly
ellipsoid, somewhat narrowed towards the apices, and
definitely 1-septate. Asci in different developmental stages,
a paraphysis, and hyphae of the fungus and host lichen are
also figured.
We could find no report of an identical fungus on Physcia
caesia, but the details supplied suggest that the species may
well belong to Arthonia, although the paraphyses figured
are not as anastomosed as would be expected for that genus.
There is a species A. epiphyscia Nyl. 1875 which is wellknown on Physcia species, including P. caesia, but that has
larger ascospores (10–14 x (3–)3.5–5 μm; Purvis et al.
1992). We also considered a possible placement in Scutula,
which would accord more closely with the figured
paraphyses, but the dense epithecial tissue figured over the
hymenium mitigates against that possibility.
In the absence of the holotype specimen, one option
would be to designate the published illustration as lectotype
but we refrain from doing that here as the specimen may yet
be located in FH or fresh material suitable for
neotypification may be discovered.
A formal transfer to Arthonia is made here, even in the
absence of any material, as the fungus should be readily
recognizable if recollected and it can evidently be separated
Fig. 4. Sphaerellothecium buelliae (FH - lectotype). a. developing ascomata on the surface of the lichen host showing the superficial hyphae,
and b. a more mature ascoma. Bars = 10 μm.
http://journals.cambridge.org
Downloaded: 11 Dec 2013
IP address: 202.75.16.18
296
D.L. HAWKSWORTH & T. ITURRIAGA
from A. epiphyscia by the much shorter and narrower
ascospores.
(6) Orbicula buelliae C.W. Dodge, BANZARE 1929–31,
series B, 7: 261 (1948).
Type: Queen Mary Land, Possession Nunatak, c. 66º45'S
98º30'E, on “Buellia muscicola”, 15 Dec. 1912, C.T.
Harrisson A.A.E. 12 (FH – element with spherical ascomata
on a superficial mycelium, lectotypus hic designatus).
≡ Endococcus buelliae (C.W. Dodge) Matzer, Cryptogamie,
Mycol. 14: 14 (1993).
≡ Sphaerellothecium buelliae (C.W. Dodge) D. Hawksw. &
Iturr., comb. nov.
(Fig. 4)
Ascomata superficial, arising on a weft of superficial
hyphae on the surface of the host lichen, globose, dark
brown to black, (20–)30–55 μm diameter, ostiole not
apparent; exciple of 2–3 layers of pseudoparenchymatous
cells. Superficial hyphae ramifying over the surface, brown
to dark brown, septate, often swollen between the septa,
3–4.5 μm thick, walls 0.5–1 μm thick. Asci and ascospores
not seen.
The original collection includes two lichenicolous fungi,
one described by Dodge with the ascomata occurring on a
network of superficial hyphae (this is the only one present
on the only slide under this name), and the other with
erumpent ascomata not associated with superficial hyphae
of which slides were made by Matzer (also preserved in FH)
and led him to make the new combination into Endococcus
(Matzer 1993). The ascomata and ascospores of the
Endococcus were reported by Matzer as 100(–200) μm and
c. 18–13 x 6–8 μm respectively, quite different from the
measurements of Dodge who gave the ascomata as up to
50 μm diameter and the ascospores “about 6 x 4 μ”. We can
only presume that Matzer never studied Dodge’s original
slide, and find it inconceivable that Dodge would have been
so far out with his microscopic measurements in this
description; our experience with other species in the same
paper is that his measurement were generally in the same
range as those we made.
The fungus with small spherical ascomata on the
superficial mycelium is clearly the species described by
Dodge and that has to be selected as the lectotype element
as it is the part that agrees with the protologue. We consider
this species to be best accommodated in Sphaerellothecium
and therefore make the new combination here. We did not
see any mature asci or ascospores, and it seems likely
Dodge did not either as he describes them as “hyaline
becoming brownish” and does not mention the presence of a
septum; we would expect the ascospores to be brown and
1-septate at maturity. Most species in the genus have a
narrow host range, none is currently recognized as
http://journals.cambridge.org
Downloaded: 11 Dec 2013
Fig. 5. Endococcus matzeri (FH – holotype). a. maturing ascus,
and b. ascospores. Bar = 10 μm.
occurring on Buellia, and the ascospore sizes given by
Dodge are smaller than those published for other accepted
species in the genus. We therefore make the transfer here,
recognizing that a full description will have to await the
collection of further material.
The second fungus, as Matzer (1993) points out, appears
distinct from the known Endococcus species and so we
describe this as new below.
The host name Dodge used for the host lichen, Buellia
muscicola, was a species described by Dodge & Baker
(1938) from King Edward VII Land, but this name is not
mentioned by Lamb (1968), Castello & Nimis (1995), nor
Øvstedal & Smith (2001). We can confirm that it represents
a Buellia as apothecia belonging to that genus are present in
Harrisson’s collection; the thallus is creamy to whitish,
areolate, and the ascospores are 1-septate and 12–15.5 x
7–9 μm, rather short for many of the Antarctic species; the
material is so fragmentary that it was impossible to pursue
the identity of the host further, but it did not key out clearly
in Øvstedal & Smith (2001).
Endococcus matzeri D. Hawksw. & Iturr., sp. nov.
(Figs 5 & 6)
Similis Endococco propinquo, sed ascosporis ornamentatis,
asperulatis, 9.5–11.5(–12) x 5–6.5 μm.
Typus: Queen Mary Land, Possession Nunatak, c. 66º45'S
98º30'E, on “Buellia muscicola”, 15 Dec. 1912, C.T.
Harrisson A.A.E. 12 (FH – element with ellipsoid erumpent
ascomata not on a superficial mycelium, holotypus).
Ascomata perithecioid, subglobose, ostiolate, immersed at
IP address: 202.75.16.18
297
LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE
(7.5–)8.5–12.5(–13) x 5–7(–7.5) μm by Triebel (1989), and
we did not find any ascospores as wide as the 8 μm he
reported. The length:breadth ratios of the ascospores are
also in the same range, to judge from numerous collections
documented in the folders of notes and drawings on the
genus (including the neotype collection in K) of the first
author. The key difference is the ascospore ornamentation
seen only at high magnifications with quality research
microscopes, and most strikingly visible in maturing
ascospores where the full wall pigmentation has not
developed. This is something not reported before, and,
taken with the different host genus, is our reason for
recognizing this species here. Previous reports of
E. propinquus from Buellia species need to be reassessed to
determine if they also have the asperulate ascospores of
E. matzeri.
(7) Phaeospora gasparriniae C.W. Dodge, BANZARE
1929–31, series B, 7: 262 (1948).
Fig. 6. Endococcus matzeri (FH – holotype). a. detail of exciple,
the central cavity to the right, b. mature ascus with eight
ascospores, and c. immature ascospore showing the ornamented
walls. Bar = 10 μm.
first but becoming erumpent, black, 140–185 μm diameter;
exciple 23–31 μm thick, composed of 8–10 layers of thickwalled angular radially compressed and pseudoparenchymatous cells, textura angularis, individual cells
5–10 x 2–4 μm; lined internally by 3–4 layers of colourless
pseudoparenchymatous cells, individually 4–6 x 2.5–4 μm;
ostiole not papillate, gaping, 37–75 μm. Hamathecium
absent at maturity. Asci bitunicate in structure, broadly
clavate, thick-walled, c. 40 x 14 μm, 8-spored. Ascospores
irregularly overlapping within the asci, broadly ellipsoid,
rounded at the ends, 1-septate, not or only slightly
constricted at the septum, cells ± equal in size, appearing ±
smooth at lower magnifcations, but faintly but regularly
asperulately ornamented at 1500x, without a distinct
gelatinous sheath, 9.5–11.5(–12) x 5–6.5 μm,
length:breadth ratio 1.5–1.6–2.2.
This species is very close to Endococcus propinquus
(Körb.) D. Hawksw. 1979 which occurs mainly on lichens
in the Porpidiaceae (e.g. Amygdalaria, Porpidia) but is also
reported from a variety of other crustose lichens, including
Buellia buellioides (Triebel 1989). Matzer (1993) stressed
the relatively smaller size of the perithecia, ascospore size,
and ascospore ornamentation. We found the ascomata to be
in the range of E. propinquus, given as
(100–)150–200(–400) by Triebel (1989); we suspect that
Matzer's belief that they were mainly to 100 μm may have
been due to superficial measurements made on the
Sphaerellothecium which he did not recognise as also being
present. The ascospore range he gave, “c. 8–13 x 6–8 μm”,
is also pretty well in the range of E. propinquus, given as
http://journals.cambridge.org
Downloaded: 11 Dec 2013
Type: Macquarie Island, north end, on “Gasparrinia
maquariensis” [? Caloplaca sp.], BANZARE. B-540-16
(holotype not located).
≡ Weddellomyces gasparriniae (C.W. Dodge) D. Hawksw.
& Iturr., comb. nov.
The original description indicated that this fungus formed
spherical black ostiolate perithecia c. 180 μm diameter,
which were semi-immersed when mature. It was described
as having both periphyses and discrete unbranched slender
paraphyses, cylindrical to narrowly fusiform 8-spored asci
c. 65 x 9 μm, and brown thick-walled 3-septate ascospores
constricted at the septa and measuring c. 22 x 5 μm. The
host was presumably a lobate species of Caloplaca.
The presence of interascal filaments precludes a
placement in Phaeospora, and that they were described as
“slender” suggests they were paraphysoids even though no
anastomoses were noted. This feature, along with the
thick-walled brown 3-septate ascospores constricted at the
septa, indicates that the fungus was almost certainly a
species of Weddellomyces. Three species of that genus are
known to grow on Caloplaca species in Europe, but all
have longer and broader ascospores than the fungus Dodge
described: (21.5–)24–31(–38) x (9–)9.5–12(–13.5) μm in
W. epicallopisma (Wedd.) D. Hawksw. 1986, (28.5–)
30–40(–45) x (10–)11–15 μm in W. erythrocarpae
Nav.–Ros. & Roux 1995, and (24.5–)27–32.5(–39) x
(10.5–)11–13.5(–14–5) μm in W. protearius Nav.-Ros. &
Roux 1997 (Hawksworth 1986, Navarro-Rosinés & Roux
1995, 1997, Calatayud & Navarro-Rosinés 1998).
While we hesitated over making a new combination for
this fungus without seeing any material, as the probability
that this is the correct genus is high, this decision will
ensure that the name is taken into consideration when
additional species of the genus on Caloplaca are discovered
IP address: 202.75.16.18
298
D.L. HAWKSWORTH & T. ITURRIAGA
Fig. 7. Phacopsis usneae (FH – neotype). a. apothecia clustered
on a branch of the host lichen, b. paraphyses apices showing
granules and the apical caps, c. asci in different stages of
development, and d. mature ascospores. Bars = 0.5 mm (a), and
10 μm (b–d).
and material suitable for neotypification is found.
(8) Phacopsis usneae C.W. Dodge, BANZARE 1929–31,
series B, 7: 264 (1948).
Type: Kerguelen, Mount Wyville Thompson, 1000–1500 ft,
on Usnea trachycarpa, BANZARE B246-21 (holotype not
located); Melchior Archipelago, Lystad Island, north face,
alt. 20–30 ft, on U. antarctica [as “U. granulifera”], 14
March 1941, McCoy & P.A. Siple 331 (FH – neotypus hic
designatus [specimen and slide]).
(Fig. 7)
Ascomata apothecioid, erumpent at first but soon becoming
superficial, arising singly or more usually aggregated into
dense mounded clumps, black, the clumps occupying and
extending around the host branches and to 2–3 mm in
length; individual ascomata strongly convex, circular from
above, 0.25–0.75 mm diameter, emarginate, in vertical
section 60–90(–100) μm tall; hypothecium dark brown,
composed of thick-walled, intertwined hyphae forming
angular pseudoparenchymatous tissue in parts, cells mainly
7–10 μm diameter and the hyphae 5–7 μm thick, together
forming a compact cushion-like layer varying in thickness;
thecium colourless, 45–65 μm tall; epithecium welldeveloped, 12–20 μm tall, dark blue-grey, composed of a
mucilaginous gel surrounding and interspersed with the
paraphyses tips, binding them together, homogenous but
granular on the paraphyses tips. Hamathecium of densely
packed paraphyses, repeatedly septate, often somewhat
swollen between the septa, not or more commonly sparsely
http://journals.cambridge.org
Downloaded: 11 Dec 2013
branched and sometimes anastomosed, thick-walled,
mainly 3–4 μm thick, but becoming gradually swollen,
elongate–clavate, and to 9.5–10 μm wide at the tips where
the cell walls can be 3–4 μm thick, the tips with dark bluegrey granules and then capped with a dark blue-grey
mucilaginous sheath that can extend 5–6 μm above the tips.
Asci broadly clavate, Lecanora-type, with a strongly
thickened apex that can occupy to a third of the ascus length
when mature, 40–47 x 14–25 μm, 8-spored. Ascospores
irregularly overlapping in the asci, ellipsoid, rounded at the
ends, simple, smooth-walled, without an evident sheath,
9.5–10.5 x 5.5–6.5 μm.
No chemical reagent tests were carried out in view of the
limited amount of material available on the neotype
specimen. The distinctive blue-grey pigmented mucilage
was evident in water mounts and in lactofuschsin. Our
measurements and observations agree closely with those of
Dodge except that he gave the ascospores as “10–12 x 4 μ”,
but this may be due to our finding few mature ascospores;
he also described them as “fusiform”, but the apices are not
at all pointed.
Although the holotype could not be located, McCoy &
Siple 331 is annotated as this species by Dodge, and is
represented by a slide. Although collected prior to the
description of the fungus, it was not mentioned in the
original account and Dodge’s annotations on the packet and
slide are undated; it is therefore selected as a neotype rather
than a lectotype here. There are also two microscope slides
labelled as “Phacopsis usneae?” from different localities on
other Usnea species: on “Usnea aurantiaca-atra” from
Refuge Island (Bryant 32) and “Usnea siplei” from George
VI Sound (Eklund 860). In addition there are two
collections not named to species: as “Phacopsis” with
“U. fasciata” from Lambda Island (3 March 1941,
G. Frazier, Bailey & P.A. Siple 383a-1), and as “Nesolechia”
(a name regarded as a synonym of Phacopsis by Triebel
et al. 1995) on “U. frigida” from Eights Coast (Jones
Mountains, Lichen Point, 73º32'S 94º00'W, ? Jan. 1961, G.F.
Webers 61-1-1B-B).
This name was evidently overlooked in the monograph of
the genus by Triebel et al. (1995), who described one
species on Usnea as new, P. falcispora Triebel & Rambold
1995 on U. maculata from South Africa. That species is
clearly distinct from P. usneae and was figured as having
crescent-shaped ascospores, often attenuated at the apices,
and measuring (11.5–)13.5–15(–16) x 5–5–5(–6) μm (var.
falcispora) or (12–)14–16(–19) x 5–5.5(–6) μm (var.
sipmanii).
(9) Thelidiola eklundii C.W. Dodge, Nova Hedwigia 15:
300 (1968).
Type: Antarctic Peninsula, Nunatak, 3 miles south-east of
Mount Wilcox, north wall of McClary glacier, on
“Catillaria eklundii” [? Buellia sp.], 68º00'S 67º47'W,
IP address: 202.75.16.18
299
LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE
C.Eklund B-6-A [56] pro parte (FH – holotype).
= Muellerella pygmaea (Körb.) D. Hawksw., Bot. Notiser
132: 289 (1979).
This is the type species of Thelidiola, a new generic name
introduced by Dodge (1968, p. 299) for this species and
Thelidium caloplacae (see below). Two packets were
located in FH under this name, one with slightly fuller
locality information, but both with the collector´s number
“B-6-A”. However, only one has the US Antarctic Service
Expedition number 56, which was cited by Dodge and so
that is considered the holotype here. We suspect the second
collection was split from the first, perhaps as both also have
the species Catillaria eklundii Dodge 1968, described in the
same publication, and that this was done to enable a part to
be filed in the herbarium separately under the Catillaria
name. Castello & Nimis (1995, p. 89) give the correct “B-6A” reference in the entry under C. eklundii, which they
considered “by an educated guess, could be attributed to a
Buellia”, but under Thelidiola eklundii refer again to two
packets having both taxa, but give the reference as “Siple
et al. DW-4”; this number actually relates to the holotype of
T. caloplacae and not of T. eklundii; we therefore interpret
this as a simple lapsus for “B-6-A” which contains a
Muellerella, the ascospores of which “suggest
M. lichenicola”. However, the holotype specimen also has
an annotation label by Vagn Alstrup dated 23 March 1995
identifying the specimen as M. pygmaea. The perithecia
arise almost superficially and are rather large, reaching
260 μm diameter, and the ascospores are dark brown,
broadly ovoid, 1-septate, and the size range we measured
was 7–9(–10) x 3.5–4.5 μm. These observations and
measurements essentially agree with those of Dodge, who
stated that the perithecia were sessile, to 270 μm diam, and
the ascospores 10 x 4 μm. All these features conform to
M. pygmaea rather than M. lichenicola, more particularly
M. pygmaea var. pygmaea as understood by Triebel (1989).
In M. lichenicola the perithecia are generally immersed,
50–100 μm diameter, and the ascospores 5–7(–8) x 2–3
(–4) μm (Hawksworth 1979).
(10) Thelidium caloplacae C.W. Dodge & G.E. Baker, Ann.
Mo. Bot. Gdn 25: 525 (1938).
Type: Marie Byrd Land, Edsel Ford Range, Mt Donald
Woodward [as “Mt Rea-Cooper”; see below], on Xanthoria
elegans [as “Gasparrinia siplei and Kuttlingeria rufa”],
1934, P. Siple, F.A. Wade, S. Corey & O.D. Stancliff DW-4
(FH – holotype).
In addition to the specimen designated as type, Dodge &
Baker (1938, p. 526) also cited a collection from Skua Gull
Peak, F. Siple & S. Corey 72W-6. Specimens with the
numbers DW-4 and 72W-6 are both present in FH and
Castello & Nimis (1995, p. 88) considered these were
“probably” Muellerella pygmaea. There is, however, some
uncertainty over the typification as the locality name on
packet DW-4 is given as “Mt Rea-Cooper”; this is
presumably why Castello & Nimis state “type material not
located” even though they had studied DW4. Interestingly,
according to the route map published by Siple (1938,
p. 475) Donald Woodward Mount is very close to but
separated from Mount Rea and Mount Cooper (which are
separate peaks in the same mountain structure) by the Ames
Glacier. However, Siple (1938, p. 487) mentions this
species as present on Mount Donald Woodward, but not on
Mount Rea-Cooper (Siple 1938, p. 492–493). At the former
site the lichens are reported as growing on a schist, while at
the latter on a coarse-grained light pink granite. The
specimen does not appear to be on a granite, but rather a
homogenous schist, and the substrate indicated in the
protologue is schist not granite; we therefore suggest that
the locality name on the packet is an error of which Siple
was aware and so he cited the species only from the one
Mount Donald Woodward site. Dodge (1973, p. 14) also
does not mention Mount Rea Cooper as a locality for this
species.
The species was originally described as “parasitic on the
thalli of Gasparrinia siplei and Kuttlingeria rufa” in Dodge
& Baker (1938), while the packets “DW-4” and “72W-6” in
FH are labelled “on sterile Blasteniaceae”. The host in DW4 is actually a fragment of Xanthoria elegans, of which the
name K. rufa is a synonym according to Castello & Nimis
(1995). The last authors indicate that Gasparrinia siplei (as
K. siplei) is Caloplaca lucens but we did not note this on the
material we studied.
Santesson (1960, p. 508) was the first to suggest the
species was probably conspecific with M. lichenicola, as
noted by Hawksworth (in Pegler et al. 1980, p. 522). The
perithecia were immersed, erumpent only in the uppermost
part, and the largest we measured was 100 μm diameter and
the ascospores in DW-4 were 6–7.5 x 2–3(–4.5) μm, all
within the reported range for M. lichenicola (e.g.
Hawksworth 1979, Triebel 1989).
(11) Thelidium minutum C.W. Dodge, Nova Hedwigia 15:
301 (1968).
≡ Thelidiola caloplacae (C.W. Dodge & G.E. Baker)
Dodge, Nova Hedwigia 15: 299 (1968).
Type: Victoria Land, Walcott Glacial Area, south-east of
Lake Penny, on Rhizoplaca melanophthalma [as “Lecanora
lavae”], 22 February 1964, K.P. Rennell [E. Schofield AA135] (FH –holotype).
= Muellerella lichenicola (Sommerf.) D. Hawksw., Bot.
Notiser 132: 289 (1979).
= Muellerella pygmaea (Körb.) D. Hawksw., Bot. Notiser
132: 289 (1979).
http://journals.cambridge.org
Downloaded: 11 Dec 2013
IP address: 202.75.16.18
300
D.L. HAWKSWORTH & T. ITURRIAGA
According to Castello & Nimis (1995, p. 89) this was a
Muellerella, “perhaps M. pygmaea (Körb.) D. Hawksw.”
However, a later annotation slip by Vagn Alstrup (23 March
1995) identifies this is Muellerella lichenicola growing on
Rhizoplaca melanophthalma. The perithecia measure
120–250 μm diameter and the ascospores are broadly
ellipsoid, dark brown and 2-celled when mature, and
measure 6–8(–9) x 4.5 μm, which matches well with the
concept of M. pygmaea s. str. (Hawksworth 1979, Triebel
1989). The thallus consists of bullate areoles and is clearly
that of a Rhizoplaca and not at all like Lecanora flotowiana
which has no evident superficial thallus and of which the
name L. leave is a synonym (Øvstedal & Smith 2001,
p. 206).
(12) Thelidium parvum C.W. Dodge & G.E. Baker, Ann.
Mo. Bot. Gdn 25: 526 (1938).
Type: Marie Byrd Land, Edsal Ford Range, Skua Gull Peak,
growing over “Parmelia variolosa” [? Physcia caesia], P.
Siple & S. Corey 72W-4 (holotype not located).
= Stigmidium pumilum (Lett.) Matzer & Hafellner,
Bibltheca Lich. 37: 115 (1990).
The biological status of this species is obscure as the fungus
was described as forming hyphae which surrounded
“miscellaneous algae principally Scytonema and
Pleurococcus” with the whole “growing over the thallus of
Parmelia variolosa”. The host lichen named as P. variolosa
is Physcia caesia (see under Diplonaevia parmeliae above).
No specimen of Thelidum parvum was located by Castello
& Nimis (1995, p. 73), and nor did we find one, but in this
case an illustration was provided by Dodge & Baker (1938,
pl. 62, figs 393–395). This shows a section of a perithecium
with a pseudoparenchymatous cell wall, some suggestion of
interascal tissue; maturing asci (probably the contents
without the outer wall as a dimple appears at the apex), two
broadly soleiform ascospores, and hyphae ensheathing the
two kinds of algae. The perithecia were described as to 50 x
55 μm, with no paraphyses but “gelatinous filmy strands
among the asci”; the asci as elongate and somewhat clavate,
28–31 x 10–12 μm, and 8-spored; and the ascospores as
1-septate, hyaline, not constricted at the septum, with one
cell larger and blunt at the end and the other smaller and
more pointed, and 7.5–9 x 3–3.5 μm.
The maximum perithecium size given is much too small
for the lichen-forming genus Thelidium, the size range for
perithecia in the genus being 0.08–6.5 mm (Purvis et al.
1992). Critical to any placement is the nature of any
interascal tissues, but it seems unlikely that “gelatinous
filmy strands” were really some form of paraphyses as such
structures were carefully documented in many other species
treated in the same work. It seems most probable that Dodge
& Baker were dealing with a lichenicolous species of
Stigmidium, although they do not state if the perithecia were
http://journals.cambridge.org
Downloaded: 11 Dec 2013
immersed or superficial, and perhaps some other hyphal
fungi and cyanobacteria/algae. Indeed, the dimensions of
the ascomata and asci are in the ranges reported in
S. pumilum by Matzer & Hafellner (1990). Stigmidium
pumilum is confined to Physcia caesia and the closely allied
P. dubia and known from Chile and New Zealand as well as
the Canary Islands and Europe (Matzer & Hafellner 1990,
Wedin 1994). However, the ascospores in S. pumilum
measure (8–)9–13(–16) x 3–7 μm, embracing those
reported for T. parvum, but also becoming longer and
broader. Bearing in mind the coincidences as to dimensions
and possible host, however, we suggest that the species
belongs here and that Dodge & Baker simply measured
rather few and still maturing ascospores.
Acknowledgements
We are indebted to Donald H. Pfister, Scott LaGreca, and
especially Genevieve Lewis-Gentry, for diligently and
repeatedly searching for specimens and slides in the Dodge
collections at the Farlow Herbarium and Reference Library
(Harvard University) for us to study and to be sure elusive
collections were really no longer present. We are also
indebted to Josef Hafellner for his perceptive and helpful
comments on our original submission. This work was
undertaken while D.L.H. was supported by the Programa
Ramón y Cajal of the Ministry of Science and Technology
of Spain, and T.I. was on sabbatical in the Departamento de
Biología Vegetal II of the Facultad be Farmacia in the
Universidad Complutense de Madrid funded by the
Ministry of Education and Culture of Spain.
References
APTROOT, A., DIEDERICH, P., SÉRUSIAUX, E. & SIPMAN, H.J.M. 1997.
Lichens and lichenicolous fungi from New Guinea. Bibliotheca
Lichenologica, 64, 1–220.
BRODO, I.M. & HAWKSWORTH, D.L. 1977. Alectoria and allied genera in
North America. Opera Botanica, 42, 1–164.
CALATAYUD, V. & NAVARRO-ROSINÉS, P. 1998. Weddellomyces
xanthoparmeliae sp. nov. and additions to the chorology of other species
of the genus. Mycotaxon, 69, 503–514.
CASTELLO, M. & NIMIS, P.L. 1995. A critical revision of Antarctic lichens
described by C. W. Dodge. Bibliotheca Lichenologica, 57, 71–92.
DODGE, C.W. 1936. Medical mycology. London: Henry Kimpton, 900 pp.
DODGE, C.W. 1948. Lichens and lichen parasites. British and New Zealand
Antarctic Research Expedition 1929–1931, Reports, Series B (Zoology
and Botany), 7, 1–276.
DODGE, C.W. 1968. Lichenological notes on the flora of the Antarctic
continent and the sub-Antarctic islands. VII and VIII. Nova Hedwigia,
15, 285–332.
DODGE, C.W. 1973. Lichen flora of the Antarctic Continent and adjacent
islands. Cannan, NH: Phoenix Publishing, 399 pp.
DODGE, C.W. & BAKER, G.E. 1938. [Second Byrd Antarctic Expedition –
Botany] II. Lichens and lichen parasites. Annals of the Missouri
Botanical Garden, 25, 515–727.
DODGE, C.W. & ZELLER, S.M. 1934. Hymenogaster and related genera.
Annals of the Missouri Botanical Garden, 21, 625–705.
IP address: 202.75.16.18
301
LICHENICOLOUS FUNGI DESCRIBED BY C.W. DODGE
GRUBE, M. & HAFELLNER, J. 1990. Studien an flechtenbewohnenden Pilzen
der Sammelgattung Didymella (Ascomycetes, Dothideales). Nova
Hedwigia, 51, 283–360.
HALE, M.E. 1987. Epilithic lichens in the Beacon sandstone formation,
Victoria Land, Antarctica. Lichenologist, 19, 269–287.
HAWKSWORTH, D.L. 1979. Studies in the genus Endococcus
(Ascomycotina, Dothideales). Botaniska Notiser, 132, 283–290.
HAWKSWORTH, D.L. 1986. Notes on British lichenicolous fungi: V. Notes
from the Royal Botanic Garden Edinburgh, 43, 497–519.
HAWKSWORTH, D.L. 2003. The lichenicolous fungi of Great Britain and
Ireland: an overview and annotated checklist. Lichenologist, 35,
191–232.
HAWKSWORTH, D.L. 2004. Fungi living on lichens: a source of unexplored
diversity. British Wildlife, 15, 192–199.
HAWKSWORTH, D.L. & COLE, M.S. 2002. Intralichen, a new genus for
lichenicolous “Bispora” and “Trimmatostroma” species. Fungal
Diversity, 11, 87–97.
LAMB, I.M. 1964. Antarctic lichens I. The genera Usnea, Ramalina,
Himantormia, Alectoria, Cornicularia. British Antarctic Survey
Scientific Reports, No. 38, 1–34.
LAMB, I.M. 1968. Antarctic lichens II. The genera Buellia and Rinodina.
British Antarctic Survey Scientific Reports, No. 61, 1–129.
LAWREY, J.D. & DIEDERICH, P. 2003. Lichenicolous fungi: interactions,
evolution, and biodiversity. Bryologist, 106, 80–120.
MATZER, M. 1993. Beitrag zur Kenntnis der Ascomycetengattungen
Globosphaeria, Roselliniopsis und Synaptospora. Cryptogamie,
Mycologie, 14, 11–19.
MATZER, M. & HAFELLNER, J. 1990. Eine Revision der lichenicolen Arten
der
Sammelgattung
Rosellinia
(Ascomycetes).
Bibliotheca
Lichenologica, 37, 1–138.
http://journals.cambridge.org
Downloaded: 11 Dec 2013
MURRAY, J. 1963. Lichens from Cape Hallett area, Antarctica. Transactions
of the Royal Society of New Zealand, 2(5), 59–72.
NAVARRO-ROSINÉS, P. & ROUX, C. 1995. Le genre Weddellomyces
(Dothideales, Dacampiaceae) en Catalogne et en Provence. Mycotaxon,
53, 161–187.
NAVARRO-ROSINÉS, P. & ROUX, C. 1997. Weddellomyces protearius sp. nov.
et Lichenochora xanthoriae, champignons lichénicoles non lichénisés
parasites de Caloplaca proteus. Mycotaxon, 61, 433–440.
ØVSTEDAL, D.O. & SMITH, R.I.L. 2001. Lichens of Antarctica and South
Georgia: a guide to their identification and ecology. Cambridge:
Cambridge University Press, 411 pp.
PEGLER, D.N., SPOONER, B.M. & SMITH, R.I.L. 1980. Higher fungi of
Antarctica, the subantarctic zone and Falkland Islands. Kew Bulletin, 35,
499–562.
PURVIS, O.W., COPPINS, B.J., HAWKSWORTH, D.L., JAMES, P.W. & MOORE,
D.M. 1992. The lichen flora of Great Britain and Ireland. London:
Natural History Museum Publications, 710 pp.
RUDOLPH, E.D. 1990. Carroll William Dodge, 1895–1988. Mycologia, 82,
160–164.
SANTESSON, R. 1960. Lichenicolous fungi from northern Spain. Svensk
Botaniska Tidskrift, 39, 499–522.
SIPLE, P.A. 1938. The second Byrd Antarctic expedition – Botany I.
Ecology and geographical distribution. Annals of the Missouri Botanical
Garden, 25, 467–514.
TRIEBEL, D. 1989. Lecideicole Ascomyceten: Eine Revision der obligat
lichenicolen Ascomyceten auf lecideoiden Flechten. Bibliotheca
Lichenologica, 35, 1–278.
TRIEBEL, D., RAMBOLD, G. & ELIX, J.A. 1995. A conspectus of the genus
Phacopsis (Lecanorales). Bryologist, 98, 71–83.
WEDIN, M. 1994. New and noteworthy lichenicolous fungi from
southernmost South America. Lichenologist, 26, 301–310.
IP address: 202.75.16.18
http://journals.cambridge.org
Downloaded: 11 Dec 2013
IP address: 202.75.16.18