Lichenologi~t 27(6):
THE
463-471
LICHEN
Ulrik
(1995)
GENUS CALOPLACA
REGIONS
SQ)CHTING*
and Maria
IN POLAR
OLECH$
Abstract:
Extensive
material
of Calopluca
from Arctic
and Antarctic
regions has
been critically
examined.
A list of 49 species is presented
for Arctic regions. They
are presumed
to have a more or less circurnpolar
distribution.
Twenty-two
species
are listed from the Antarctic
region,
but about ten more, probably
undescribed
species, are present there. Abour one-third
of the species in the Antarctic
region are
bipolar
or widespread
in cold regions;
these include
mainly
terricolous
and
muscicolous
species and none of them are maritime.
It is assumed that migration
of
the bipolar
or cosmopolitan
species has taken place along the Andean
mountain
chain, whereas
the maritime
polar species have evolved
separately
in the two
hemispheres.
The Culoplaca
species of the Antarctic
region
are provisionally
assigned
to the following
distribution
types:
continental
Antarctic,
western
Antarctic,
insul-Antarctic
and sub-Antarctic.
Culoplucu
exrecucu, C. suxicolu and
C. phaeocarpellu
are recorded
as new to the Antarctic
region. Culoplucu johnstonii
(Dodge)
Sochting
& Olech,
comb. nav., is established
as the correct
name of
C. tenuis Ovstedal.
Q 1995 The British
Lichen
Society
Introduction
Arctic and alpine regions are known to have a large number of Caloplaca
species. However,
there are few comprehensive
treatments
covering such
regions (Hansen et al. 1987; Poelt & Hinteregger
1993). Phytogeographical
studies are hampered by the sparse collecting of the more subtle species
and the difficulties in identifying the species due to lack of suitable floras. In
recent years more information
has accumulated due to focused collecting in
Greenland,
Svalbard, Siberia and northern
Fennoscandia.
Although many
taxonomic
problems
are still unsolved and a number of species are still
undescribed
it is now possible to get a broad overview of the distribution
of a
considerable number of Caloplaca species.
Recent papers on the Caloplaca flora of the Antarctic region (see below) and
our own continuing studies of extensive collections from Antarctica
and the
sub-Antarctic
islands have made it possible to compare the polar floras of the
two hemispheres. The present paper is based on published data and a number
of as yet unpublished
records from polar regions, which will eventually be
treated more thoroughly.
Phytogeographical
compilations
should be made
with extreme care, as published data are severely burdened by misidentifications. Most data here are accordingly based on material we have studied
ourselves.
*Department
DK-1353
SBotanical
of Mycology,
Botanical
Institute,
University
of Copenhagen,
0. Farimagsgade
Copenhagen
K, Denmark.
Institute,
Jagellonian
University,
Ul. Lubicz 46, F-3 1-5 12 Krakow,
Poland.
0024-2829/95/060463+09
$12.00/O
0
1995 The
British
Lichen
2D,
Society
464
THE
Vol. 27
LICHENOLOGIST
Delimitation
of the Polar
Floras
Arctic and sub-Arctic
regions are found in Alaska and Canada, Greenland,
Svalbard, northern Fennoscandia,
European Russia, northern Siberia and the
Russian far North-East.
We have not attempted to draw a southern border for
these regions, as the ranges of the species are still insufficiently
known.
Instead, the floras of Greenland and Svalbard, both well studied by us, have
been used to define the northern polar element. Similarly, the southern polar
element (hereafter referred to as Antarctic)
is defined as the species occurring
on the Antarctic continent and on the South Shetland Islands, areas that have
recently been intensively studied by us. Species unknown
from those areas,
but recorded from other peri-Antarctic
islands are not included.
The Arctic
Caloplaca
Flora
The Greenland Caloplaca flora has been thoroughly studied by Hansen et al.
(1987). The data in Table 1 are based on this source, but with some
modifications
based on later studies (unpublished).
The records of Caloplaca
species from Svalbard are based on Sochting (1989 and unpublished records)
and Elvebakk & Hertel (1995). The lichen flora of northern Fennoscandia
is
documented
in Santesson (1993) and the species included here are those
occurring in the provinces of Firmmark,
Troms and Tome Lappmark.
Table
1 also includes unpublished collections made by Sachting. At least 13 species
not known from Greenland or Svalbard occur in those provinces, but are not
included in Table 1. Records from North America are derived from the North
American checklist (Egan 1987) and only in a few cases have been verified or
changed by us. Accordingly,
the occurrences
in North America must be
considered provisional. Records from the northernmost
parts of the former
Soviet Union were recently published by Sechting et al. (1992). Older records
from those regions have largely been omitted if specimens have not been
checked.
The Antarctic
Caloplaca
Flora
The data on the Antarctic
species as presented in Table 1 are based on
specimens studied by ourselves, as the literature
on Antarctic
lichens is
often unreliable. Several of the records presented here have not previously
been published,
and we are presently studying about 10 more, probably
undescribed,
species from the Antarctic area. In the present paper we also
include data from a number of islands in the sub-Antarctic
that have a very
similar flora. We are, however, aware that these islands can support additional
species with a more northern distribution.
Discussion
of Polar
Distributions
Even though some areas of the Arctic, for
northern
Asia are insufficiently
studied, it
Caloplaca flora is more or less circumpolar.
of saxicolous species, is highly influenced
example North America and
appears that the high Arctic
The distribution,
particularly
by the occurrence
of suitable
1995
Polar Caloplaca-Gchting
& Olech
465
substrata. Muscicolous
and terricolous
species seem to be more uniformly
distributed.
The proximity
of large, only slightly separated, continents
is
believed to cause the rather uniform flora, and the richness in species has
been attributed to the ability of the flora to migrate southwards
and survive
there during the late Cenozoic glaciations. Sub-Arctic
species may be more
restricted in their range, but our knowledge
about their distribution
is so far
insufficient for final conclusions.
Twenty-one
species are listed from the Antarctic
region and with our
present knowledge we estimate that about 10 more undescribed
and possibly
endemic taxa are present. About one-third of the listed Antarctic species also
occur in the Arctic, consisting
mainly of species growing on detritus and
mosses, namely: C. ammiospila, C. citrina, C. phaeocarpella, C. tetraspora and
C. tiroliensis. The species in this group are also recorded from the Central
European Alps, apart from C. phaeocarpella, which may have been overlooked.
They probably represent an Arctic-Antarctic-Alpine
element, which migrated
along the Andean mountain range, or the Malaysian-Papuan
link (Lindsay
1977). Long-distance
dispersal from the Arctic is another possibility, which,
however, seems less likely (see below).
Other non-maritime
lichens in the more mesic parts of the Antarctic region,
such as Cladonia species, have been shown to have a mostly wide distribution
in cool regions (Stenroos
1993). This pattern has been attributed to longrange dispersal, which was, however,
facilitated by occurrence
in other
continents of the Southern Hemisphere.
Surprisingly
the most common species on detritus in the Arctic, C. cerina,
has so far not been found in the Antarctic
region. There is presently no
explanation for this. Further studies on alpine Caloplaca in the tropics would
serve to clarify the interpretation
of the ‘bipolar’ distributions
and possibly
show this element to be cosmopolitan.
None of the maritime species of the Northern
Hemisphere have been found
in the Antarctic region, and the majority of Caloplaca species endemic to the
region are more or less maritime. It appears that maritime species have had
little opportunity
to migrate across the tropics and subtropics and must thus
have evolved separately in the specific polar regions.
The ‘endemism’ of polar maritime Caloplaca species is so striking that it can
hardly be attributed to differences in the ecological conditions of the polar sea
shores. Accordingly
it appears that viable diaspores of the maritime species
have not been capable of long-range dispersal between the two polar regions.
We believe that there are historical
reasons for the apparently
isolated
evolution of the floras of the two hemispheres.
The antiquity
of many
Southern Hemisphere
species was pointed out by KHmefelt (1990) based on
distribution
patterns of more northern species. Further speculation on the
origin of the maritime Antarctic Caloplaca species must await critical study of
related species on the Southern Hemisphere
continents.
Distribution
Patterns
in the Antarctic
Region
The Caloplaca species in the Antarctic region can be grouped provisionally
according to their distribution,
which reflects their ecological requirements
as
well as probably their history of dispersal.
THE
466
TABLE
LICHENOLOGIST
1. Caloplaca
Vol. 27
species occurring
in polar
regions
Polar
Caloplaca
species
Arctic
species
alcaruln Poelt
approxirnata
(Lynge)
H. Magn.
arenaria (Pers.) Miill. Arg.
borealis
(Vain.)
Poelt
caesiorufella (Nyl.) Zahlbr.
cascellana (Rashen)
Poelt
celata Th. Fr.
cerina (Ehrh.)
Th. Fr.
concilians (Nyl.)
H. Olivier
decipens (Am.)
Blomb. 81 Forss.
diphyodes (Nyl.) Jatta
epiphyca Lynge
epithallina
Lynge
Javovirescens
(Wulf.)
Dalla Terre & Samth.
fraudans
(Th. Fr.) Oliv.
jidvolutea
(Nyl.) Jatta
insularis Poelt
invadens Lynge
jemclandica
H. Magn.
jungemzanniae
(Vahl) Th. Fr.
lacrea (Massal.)
Zahlbr.
leptocheila H. Magn.
lithophila
H. Magn.*
livida (Hepp)
Jatta
tnagni-jlii
Poelt
nivalis (Koerb.)
Th. Fr.
noeisii Sechting
ad interim
paulii Poelt
psorocida Hansen,
Poelt & Sechting
pyracea (Ach.) Th. Fr.4
saxifrogonon
Poelt
scopularis (Nyl.)
Lenau
scoroplaca (Nyl.)
H. Magn.
sibirica H. Magn.
sinapisperma
(Lam. & DC.) Maheu
& Gillet
soropelta (Hansen,
Poelt & Sochting)
Sechting
spicsbergensis H. Magn.
zonlinii Savicz
tomoksis
H. Magn.
rrachyphylla
(Tuck.)
Zahlbr.
verruculifera
(Vain.) Zahlbr.
‘Bipolar’
species
annniospila
(Ach.) Oliv.
anchon-phoeniceon
Poelt & Clauzade
cinina (Hoffm.)
Th. Fr.§
exsecuta (Nyl.)
Dalla Tome & Sam&t
locations*
Gr
Sv
NA
Si
ELI
+
+
+
+
+
+
+
-I+
+
+
+
+
+
+
+
+
+
-I+
+
+
+
+
+
+
+
+
+
+
f
+
-I-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-I-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-I+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
An
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-I-
+
+
+
+
+
+
-I-
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
-I-
+
+
+
+
+
-I+
+
+
+
?
+
+
+
+
+
+
+
+
+
Continued
+
+
+
+
1995
Polar Caloplaca-Smhting
TABLE
467
& Olech
1. Continued
Polar
locations*
species
Gr
Sv
NA
Si
Eu
An
‘Bipolar’
species
continued
phaeocarpella
(Nyl.)
Zah1br.t
saxicola (Hoffm.)
Nordint
tetraspora
(Nyl.)
Oliv.
riroliensis Zahlbr.
+
+
+
+
+
+
+
+
?
+
+
+
+
?
+
+
+
+
+
+
+
+
+
+
Caloplaca
Antarctic
species
arhallina
Darb.
austrosheclandica
(Zahlbr.)
Olech & Sochting
buelliae Olech & Sochting
cirrochrooides
(Vain.)
Zahlbr.
coralhgera (Hue) Zahlbr.
iomma Olech & Sechting
isidioclada Zahlbr.
johnstonii
(Dodge)
Sochting
& Olechf
lucens (Nyl.)
Zahlbr.
millegrana
(Mull.
Arg.) Zahlbr.
psoromatis Olech & Sochting
regalis (Vain.)
Zahlbr.
siphonospora
Olech L? Sochting
sublobulata
(Nyl.)
Zahlbr.
+
+
+
+
+
+
+
+
+
+
+
+
+
+
*Gr: Greenland;
Sv: Svalbard;
NA; North America;
Si: Siberia; Eu: Europe; An: Antarctic
region.
*Species belonging
to the C. holocarpa group are here separated into saxicolous
forms, C. lirhophila
and corticolous
and lignicolous
forms, C. Dracea.
This separation
must be regarded
as highly
provisional.
Specimens
of the C. holocarpa group growing
in the Antarctic
region probably
belong
to an undescribed
taxon.
§Caloplaca cirrina is variable and badly understood.
Antarctic
material may belong to several taxa,
probably
including
the Northern
Hemisphere
taxon C. cimna.
TCaloplaca johnsronii
(Dodge)
Sochting
& Olech, comb. nav. Blastenia johnsronii
Dodge,
British,
Australian,
New Zealand Anrarcric
Research Expedition
1929-31.
7: 223 (1948).
Type:
Crozet
Islands,
Possession
Island, American
Bay. 3 November
1929, BZO-19
(FH)-Syn.:
Caloplaca
tenuis 0vsteda1,
Norsk Polarinzitutt
.Skr$cer 185: 44 (1985).
+&Species new to the Antarctic
region.
1. Continental Antarctic
Continental Antarctic species are widespread on all sides of the continent
where suitable substrata are available, but are present also on the Antarctic
Peninsula and on the South Shetland Islands (Fig. 1A). This group includes
C. athallina
and C. citrina, two species confined to often eutrophicated
cushions of mosses. Caloplaca
citrina
in the Antarctic region is poorly
understood and it is possible that it includes several taxa, at least on the
Antarctic islands.
2. Western Antarctic
Western Antarctic speciesare absent from the continent. They are restricted
to the Antarctic Peninsula, South Shetland Islands, South Orkney Islands,
THE
468
Vol. 27
LICHENOLOGIST
Tristan
b”lh ocoas
Prince
CamphIl
7
. Tristan
1..
Da Cunho
Aucklnnd
I.
LJl&a//
Edward
- South
\
Ds
w
Cunho
Sandwich
1.
Prince
I.
Edwnrd
1.
Tasm=i”
.
D
Trisrnn
Dn Cunhn
AftiC”
outh
Georgia
A-
..So”lh
:
Snndwich
I.
Prince
Edwnrd
I
alkl
a South
‘..
SAt
1.
Shcllond
1.
sit,
I.‘..
Aucklnnd
FIG.
1 Distribution
of four species
A, C. athallina
(continental
Antarctic);
(insuL4nrarctic);
of Calopluca
representing
different
B, C. regalis (western
Antarctic);
D, C. sublobulacu (SubAntarctic).
1.
‘bma”is
distribution
types.
C, C. cirrochrooides
Tierra de1 Fuego and Patagonia (Fig. 1B). Caloplaca regalis, representing this
group, is a very conspicuous and much collected lichen. Accordingly,
its range
is very reliably known. Calopluca austroshetlundica is only recorded from the
Peninsula, South Shetland Islands and South America. The western Antarctic
area is considered to have been ice-free for the longest period of time (Lindsay
Polar Caloplaca-Stichting
1995
& Olech
469
1977) and possibly the location where evolution of the species occurred. The
islands further east of South America may have a similar climate, but have a
shorter history and may not yet have been colonized by all the species with a
potential ability to grow there. Furthermore,
there may be fewer suitable
niches due to their smaller size (Engelsksjon
& Jorgensen 1986).
Caloplaca regalis is largely found on shores fertilized by penguins, whereas
C. austroshetlandica is a pioneer species on rocks in late melting sites.
3. Insul-Antarctic
Insul-Antarctic
species occur in the oceanic climate of the sub-Antarctic
islands and have spread far to the East, even to Kerguelen. However, they have
not yet been found in South America, South Africa or Australia (Fig. 1C). It
may be significant that these species are spreading probably mainly by rather
large diaspores, such as soredia, isidia and isidia-like thallus branches. Similar
observations
were made when comparing
macrolichens
from the South
Orkney Islands with those that had reached the South Sandwich
Islands
(Lindsay 1977).
Caloplaca cirrochrooides is a typical representative
of this group, but a similar
pattern is shown by C. isidioclada, which is found as far North as Tristan da
Cunha (37”15’S,
12”13’E)
and Juan Femindez
Island (345,
78-81”W).
Caloplaca coralligera, also belonging to this group, has been recorded as far
south as Ross Island (785).
Caloplaca cirrochrooides and C. coralligera are
maritime species, and all three species grow mainly on heavily fertilized rocks.
4. Sub-Antarctic
Sub-Antarctic
species are found on the sub-Antarctic
islands, but do not
occur on the Antarctic continent or on the Antarctic peninsula. They have a
more northern
distribution
ranging even to the southern shores of South
America, Africa, Australia or New Zealand (Fig. 1 D) . The northern limits of
distribution
of these species are in need of investigation.
Most widespread
is
C. sublobulata, reaching all the three continents. Caloplaca hens reaches South
America and C. millegrana is recorded from Australia. All three species are
maritime, growing mainly on eutrophicated
rocks.
5. Endemic
Endemic species are so far known from the South Shetland Islands, namely
C. buelliae, C. iomma, C. psoromatis and C. siphonospora. They may subsequently be found in other parts of Antarctica, so it is premature to discuss their
distribution.
Discussion
of Antarctic
Distribution
Patterns
Phytogeographical
zones of the Southern Hemisphere
were discussed
by
Galloway
(1991). He recognized
an Antarctic
realm including
an insulAntarctic subrealm with a very oceanic climate and a continental Antarctic
subrealm
consisting
of the Antarctic
continent
with very arid climatic
470
THE
LICHENOLOGIST
Vol. 27
conditions.
The original circumscription
of the insul-Antarctic
sub-realm
(Fleming
1987) excluded the South Shetland Islands and the Antarctic
Peninsula, two regions with an intermediate position which, at least according
to the Galoplaca
flora, do have strong affinities to the sub-Antarctic
islands
included in &he insul-Antarctic
subrealm. Hertel (1984) studied the saxicdlous
lecideoid lichens of the sub-Antarctic.
He concluded provisionally
that the
continental
flora consists of rather widespread
species, whereas the subAntarctic islands have a more specific flora. The present study indicates that
species not confined to maritime habitats are more widespread.
Species
associated with maritime climates, particularly
on the sub-Antarctic
islands,
are largely specific to the Antarctic area and have probably evolved there. It
may be speculated that they have survived on some of the western islands
during the late glaciations and subsequently
spread towards the East onto
more recently developed islands. Such a process is supported by the studies of
lecideoid lichens on Prince Edward Islands (Hertel 1984). Engelsksjan
&
Jrargensen (1986) emphasized the importance of the strong westerly winds that
are able to transport diaspores from the western Antarctic region towards the
more eastern and younger islands. Sea currents in the region have the same
direction and may also be able to transport diaspores.
We acknowledge
loan of material
from the following
herbaria:
FH, H, S and UPS, and wish to
thank Professor H. Hertel and Dr C. Wetrnore
for letting us study their collections.
Dr R. Seppelt
revised the language. He and Professor P. M. Jorgensen are thanked for valuable commems
on the
manuscript.
Ruth Bruus Jakobsen and Else Meier Andersen
prepared
the maps.
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T. &Jsrgensen,
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D. J. (1991)
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hemisphere
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I. (1990) Evidence of a slow evolutionary
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D. C. (1977)
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U. (1993) Four new species of Caloplaca from Antarctica.
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S. (1993)
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Sachting
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Accepted for publication
24 May
1995