Botanical Journal of the Linnean Society ( 1988), 96: 45-55
Plate tectonics and the distribution of
cool temperate Southern Hemisphere
macrolichens
DAVID J. GALLOWAY, F.L.S.
Department
5BD
if Botany) British Museum
(Natural History)) Cromwell Road) London SW7
Received February 1987
GALLOWAY, D.]., 1988. Plate tectonics and the distribution of cool temperate Southern
Hemisphere macrolichens. Several macrolichen genera in the families Lobariaceae and
Pannariaceae speciate richly in cool temperate areas of the Southern Hemisphere between latitudes
35 and 55°S, being common in .Nothofagus forests, subalpine shrubland, bog and grassland habitats.
Affinities of southern cool temperate lichens are discussed in terms of plate tectonics and tectonostratigraphic terranes.
ADDITIONAL KEY WORDS:-Biogeography - lichens - Lobariaceae - Pannariaceae - plate
tectonics - terranes.
CONTENTS
45
Introduction .
Cool temperate Southern Hemisphere lichens
Australasian affinities .
Austral affinities .
Explanations of cool temperate distribution patterns.
Plate tectonics
Tectonostratigraphic terranes
Conclusions
Acknowledgements
References.
46
48
49
50
51
51
52
52
53
INTRODUCTION
The Southern Hemisphere, or austral, cool temperate zone between latitudes
35 and 55°S, comprises Tasmania and south-eastern Australia, New Zealand,
southern South America and the various subantarctic islands isolated in the vast
southern oceans. Climates in the austral cool temperate zone vary from oceanic
to hyperoceanic and are characterized by low seasonal temperature fluctuations,
high rainfall and humidity, frequent cloud cover and strong westerly winds. In
Tasmania, the South Island of New Zealand, and in southern South America,
45
0024-4074j8Bf0!0045+ II $03.00/0
© !988 The
Linnean Society of London
+G
D. J. GALLOWAY
there are marked east-west gradients of rainfall across the mountain chains,
with strongly oceanic climates and high rainfall to the west, and continental
climates east, of the mountains.
South of the cool temperate zone lies Antarctica. Although now heavily
glaciated, it was sufficiently warm and temperate in the Cretaceous and early
Tertiary to allow growth of diverse plant communities, principally conifers,
cycadophytes and ferns, with angiosperms appearing in the late Cretaceous. On
the Antarctic Peninsula the most common fossil angiosperm wood is Nothofagus
Blume, and the fossil remains of Nothofagus and the families Podocarpaceae and
Araucariaceae indicate forest compositions very similar to those found today in
South America and Australasia (Francis, 1986).
There are also affinities in climate, vegetation and soils between upper
timberlines of the austral cool temperate zone and the cool tropical highlands,
with the prevailing life forms in both the subantarctic and tropical high
altitudes being cushion plants, dwarfed shrubs, and tussock grasses (Troll,
1973).
COOL TD1PERATE SOCTHERN
HE~IISPHERE
LICHENS
The similarity of lichen vegetation in cool temperate regions of Tasmania,
Zealand and South America (Galloway, 1979, 1985, 1987; Jergensen,
1983) is most marked in the rainforests of these regions (Kantvilas, James &
Jarman, 1985; Kantvilas & James, 1987; Lamb, 1959; Red6n, 1973, 1974;
Runde!, 1980). Rainforests of the austral cool temperate zone as well as having
warm temperate hardwood elements of tropical or subtropical affinities, are
notable for the presence of Nothofagus, a Gondwanan genus which is a major
component of montane and southern forests in Tasmania, New Zealand and
Chile (Barlow, 1981; Thorne, 1986; Wardle, 1984). Nothofagus has long
fascinated biogeographers concerned with patterns of distribution in austral
biomes (Craw, 1985; Heads, 1985; Humphries, 1983, 1985) and it is interesting
also to note that an obligate ascomycete parasite, C)ttaria Berk., has coevolved
with }/othr?fagus in South America and Australasia (Rawlings, 1956; Wardle,
1984). There is also a close association between species of Nothofagus and several
macrolichen genera which have their richest diversity in the cool temperate
austral rainforests (Table 1). Two monotypic genera in the widely distributed
order Caliciales also appear to be obligate epiphytes of Nothofagus: Calycidium
Stirton known from }1/.fusca Oerst. in New Zealand (Galloway, 1985), and
N. dombe_vi Blume in Chile (Red6n, 1974); and Thysanophoron Stirton, a New
Zealand endemic, known only from }1/. solandri (Hook. fil.) Oerst. var.
diffortioides (Hooker fil.) Poole (Galloway, 1985).
As well as a rich diversity of lichen species, the great dominance and biomass
of foliose lichens is a characteristic of southern beech forests, species of
Pseudocyphellaria Vainio in the family Lobariaceae especially reaching a great
size. Other important austral cool temperate lichen habitats are subalpine
shrubland, and alpine grassland or moorland, with the beech forest--grassland
ecotone being rich in lichens, with terricolous species of Cladina Nyl.,
Pseudocyphellaria, Psoroma l\fichx. and Siphula Fr. often being welldeveloped. Several widely distributed macrolichen genera have distinctive
~ew
2
I
2
5
7
6
2
I
10
7
.Nothofagus Blume
Cyttaria Berk.
21
5
6
12
2
9
9
13
48
30
4
4
2-3
5
2
7
3
4
16
12
2
7
9
6
4
6
2
7
4
2
12
8
I
4
4
3
13
11
II
2
8
l
I
I
II
9
2
?4
5
3
I
4
9
2
10
4
2
15
I
6
16
3
8
6
5
22
4
NZ
8
10
6
9
14
8
TAS
I
AUS
I
5
6
19
52
20
I
10
3
2
7
12
10
2-3
I
2
7
2
5
SA
Cladina Nyl.
Co/lema Wigg.
Degelia Arvidsson & D. Galloway
Hypogymnia (Nyl.) Nyl.
Leioderma NyJ.
Lobaria (Schreber) Hoffm.
Menegau.ia Massal.
M elaniJlia Essl.
Metus D. Galloway & P. James
.Neofuscelia Essl.
.Nephroma Ach.
.Neuropogon Nees & flotow
Pannaria Del. ex Bory
Pannoparmelia (Miill. Arg.) Darb.
Parmelia Ach.
Parmeliella Mull. Arg.
Placopsis (Nyl.) Lindsay
Pseudocyphellaria V ainio
Psoroma Michx.
Psoromidium Stirton
Siphula Fr.
Sphaerophorus Pers.
Stereocaulon Hoffm.
Sticta (Schreber) Ach.
Lichen
36
10
30
22
125
80
2
8
80
4
?40
7
?80
45
37
3
70
27
15
?50
3
?25
40
35
110
60
WORLD
2
5
I
I
5
4
6
l
3
2
2
?2-3
I
2
I
2
3
Species
in
common
Humphries (1983)
Rawlings (1956)
P. W.James (pers. comm.)
Esslinger (1977, 1978)
Galloway & James (1987)
Esslinger (1977, 1978, 1986)
F. J. White & P. W. James (pers. comm.)
Walker (1985)
Galloway (1985);Jergensen (1978)
Galloway (1978)
Galloway & Elix (1983, 1984)
Galloway (1985);Jergensen (1978)
Lamb (1947)
Galloway (1986, 1987b)
Galloway ( 1985)
Galloway &James (1985)
Galloway ( 1985)
Ohlsson (1974); Tibell (1984)
Lamb (1977)
Malme (1899); Galloway (1985)
Filson (1981)
Degelius (1974)
Arvidsson & Galloway ( 1981)
Elix (1979)
Galloway & Jergensen ( 1987)
Reference
TABLE 1. A range of austral cool temperate macrolichen genera showing numbers of species in each of four areas: southern South America (SA); southeastern Australia (AUS); Tasmania (TAS) and New Zealand (NZ), with world totals and species common to all four areas. The angiosperm Nothofagus
and its obligate parasite Cyttaria, are included for comparison
......
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5z
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;
c:
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0
z
::r:
t"l
0
t-'
.....
0
~
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0
E:
t::l
[J1
z
0
>
z
0
0
>-l
t"l
>-l
t"l
~
t-'
...,
+8
D.
J.
GALLOWAY
austral species m one or all of these habitats and include Collema Wigg.,
H_vpogymnia (i\ yl. 1 :\' yl., .Yephroma Ach., Stereocaulon Hoffrn. and Sticta
i Schreber i Ach. 'especially stalked forms); and the lichenicolous ascomycete
Plectocarpon Fee (Hawksworth & Galloway, 1984) is a common parasite of cool
temperate species of Pseudocyphellaria and ]1/ephroma (White & James,
unpublished data) in South America and Australasia.
Two major distribution patterns or affinities are discernible m austral cool
temperate lichen yegetation.
Australasian affinities
The lichen floras of Tasmania and of south-east Australia share many species
with l'\ew Zealand (Table 2), particularly in commonly occurring genera such
as Cladonia Hill ex P. Browne, and several generic segregates of Parmelia Ach.,
most noticeably Xanthoparmelia (Vainio) Hale (Elix, Johnston & Armstrong,
1986), the latter being common in coastal sites and in arid, subalpine to alpine
habitats. There is an affinity between Xanthoparmelia floras of South Africa,
inland Australia and the limited areas of continental climate in South Island,
:\'ew Zealand, an affinity shown as well by distribution patterns in Neofuscelia
Essl. (Esslinger, 1977, 1986). Rainforest macrolichens ofTasmania and southern
:\ew Zealand are remarkably similar, with 80° 0 of the Tasmanian species being
common to l'\ew Zealand (Kantvilas & James, 1987) but with the diversity in
the ~ew Zealand flora being much greater (Kantvilas et al., 1985; see also
TABLE
2. Some Australasian macrolichen genera showing numbers of species from Australia and
:\"('w Zealand, and species common to both areas.
Australia
Hato!l~Vrf'\
Pcrs,
Canoparmelia Elix & Hale
Chondrapsis :\yl.
Uadonia Hill ex P. Browne
Dendri.scoraulon :\ yl.
( ;vmrwderma :\d.
ffttnndta :\yl.
lomghtiella :\Iilli. :\rg.
Paraparmelia Elix & J. Johnston
Hm.·oparmelia Hale
l'unrtelia Krog
Rnmaiea :\'vi.
Rama/ina Ach.
Sagmzdium Stirton
Tl'ioc~< hi.1tes Norman
7 hnanothmum :\font. & Berk.
{ ·.,nen Dill. ex Adans.
Xanthoparmelia Vainio, Hale
Xanthoria , Fr. Th. Fr.
3
6
I
56
2
I
2
I
18
7
4
I
20
I
0
2
?50
117
5
Common to
Australia and
New Zealand
~ew
z~aland
3
3
3
3
2
I
13
I
I
I
I
I
45
2
I
I
I
I
2
2
4
I
4
I
9
8
I
6
2
6
2
10
15
2
47
3
47
4
Taken from: A. \\·. Archer i personal communication),
J. A. Elix (personal communication), Elix, Johnston
0
& .-\rmstrong 11986;, Elix, Johnston & \"erdon :1986;, Filson (1986), Galloway (1985): Hale (1986), and
Stn-ens 1937 .
PLATE TECTONICS AND MACROLICHEN DISTRIBUTION
49
Table 1). Similarity at the species level in the families Lobariaceae and
Pannariaceae is most marked in the genera Degelia Arvidsson & D. Galloway
(Arvidsson & Galloway, 1981); Leioderma Nyl. (Galloway & j0rgensen, 1987);
Pseudocyphellaria (Galloway, 1988); Psoroma and Psoromidium Stirton (Galloway
& James, 1985). This, plus the persistence of monotypic relicts such as K nightiella
Miill. Arg. (Galloway & Elix, 1981) in Tasmania and New Zealand, suggests
that at least part of the cool temperate rainforest and/or grassland lichen floras
of these regions may have had a common and ancient origin in cool temperate
Panthalassic Gondwanaland prior to rifting and continental drift (Galloway,
1988; Galloway & j0rgensen, 1987). Although much of the New Zealand lichen
flora is of Gondwanan affinity, several lichens are of much more recent
provenance, Chondropsis Nyl. and several species of Xanthoparmelia for example,
probably reaching New Zealand from Australia (Galloway, 1981) via the West
Wind Drift, a mechanism responsible for the recent introduction of a variety of
viable biological particles (rusts, seeds, pollen, coccids, moths, aphids, etc.) to
the New Zealand biota (Close, Moar, Tomlinson & Lowe, 1978).
Austral affinities
Godley ( 1960) made a comparison of lowland forest at 41-42°S between the
island of Chiloe, in southern Chile, and the South Island of New Zealand, and
also of cushion bogs from the austral cool temperate zone. He found that at the
family level, New Zealand forests are more diverse than those in Chile, with a
more diverse shrub layer and a greater development of ferns. A comparison of
macrolichens in cool temperate rainforest in these two areas shows the Chilean
rainforest to be less diverse lichenologically than its New Zealand counterpart
(Table 1), although the biomass of foliose lichens in both areas is of a similar
order.
Platismatia Culb. & C. Culb., a Northern Hemisphere genus is a common
epiphyte in Chile, but is not known from Australasia. Lobaria, the sister group of
Pseudocyphellaria is entirely absent from the Chilean rainforest; there are very few
species of Pannaria, and the diversity of Psoroma species seems less in Chile than it
does in New Zealand. Stalked species of Sticta in the Chilean rainforest have
cyanobionts, while their Australasian counterparts have green photobionts.
Species of Nephroma and Pseudocyphellaria, commonly dominant epiphytes in the
Chilean rainforest, have more cyanobiont-containing species than one finds in
the Australasian rainforest, where a rich development of green photobiont
species occurs. Canopy species of Usnea Dillen. ex Adans. seem less common in
Chile than they are in New Zealand where they are often visually striking at
treeline, although in some habitats in Chile Usnea is replaced by species of
Protousnea (Mot.) Krog (Krog, 1976), a genus unknown from Australasia.
Araucaria araucana (Molina) C. Koch is notable for the copious development of
species of Protousnea.
Species of Baeomyces Pers. and Placopsis (Nyl.) Lindsay and Stereocaulon which
commonly occur on roadside banks and stream beds in wetter parts of southern
New Zealand, are much more scarce in corresponding habitats in southern
Chile. The small-sorediate Teloschistes velifer F. Wilson, formerly regarded as an
Australasian species is now known to occur in Valdivia.
0
D. J. GALLOWAY
Cushion bogs are an antarctic-montane--tropical type and are not known in
the Northern Hemisphere. Main centres for these bogs and their characteristic
genera lie between latitudes 4l-56°S in Tasmania, the South Island of New
Zealand, the subantarctic islands to the south, southern Chile and Tierra del
Fuego. Phanerogams of Chilean bogs are closely related to those of Tasmania
and New Zealand, especially the genera Centrolepis Labill., Donatiaj. R. & J. G.
Forster, Gaimardia Gaud., Oreobolus R. Br. and Pkyllachne J. R. & J. G. Forster
i Godley, 1960, 1978), and there is also a strong correlation with lichens of these
habitats, with species of Cladina, Cladonia, Psoroma and Siphula being common to
all areas, but with different species of Cladia Nyl. and Pseudocyphellaria being
found in Chile and in Australasia.
EXPLA:\ATIO:\'S OF COOL
TE~IPERATE
DISTRIBCTION PATTERNS
The history of amphi-Antarctic distribution patterns has long been one of the
great problems of historical biogeography, and several generations of
biogeographers have pondered the question: how is it that so many southern
plant and animal groups are represented by related populations in southern
South America, Tasmania and New Zealand, areas that are now widely
separated by the southern oceans and an ice-covered Antarctica? Joseph Hooker
who visited these areas on the Antarctic Expedition of 1839-1843, formed the
opinion that there was once a primary southern centre of evolution, and that
disjunct populations that he saw and collected in scattered austral lands were
members of a once more extensive southern biota that had been dislocated by
geological and climatic causes. In a letter to Darwin in November, 1851 he
wrote " ... recent discoveries rather tend to ally the N .Zeald.Flora with the
.-\ustralian--though there is enough affinity with extratropical S.America to be
very remarkable and far more than can be accounted for by any known laws of
migration. I am becoming slowly more convinced of the Southern Flora being a
fragmentary one---ali that remains of a great Southern Continent . . . "
i Skottsberg, 1960).
Hooker's notion of the fragmentation of a once common ancestral biota
received support from \Vegener's theory of continental drift, and over the past
20 years from the theory of plate tectonics. The fragmentation of a Mesozoic
Gondwanan biota, which no doubt included ancestors of the lichen families
Lohariaceae and Pannariaceae for example, into vicariant Southern
Hemisphere populations isolated by geological and climatic events is now widely
accepted 1e.g. Brundin, 1966, 1975; Cracraft, 1975, 1980; Craw, 1982, 1984;
Croizat, 1958; Croizat, Nelson & Rosen, 1974; Galloway, 1987; Humphries,
1983). Southern Hemisphere biogeography is concerned with past and present
distributions of the austral biota and the processes which have given rise to these
distributions. Philosophical and methodological approaches recognized in
contemporary biogeography include ( 1) dispersal from centres of origin, a
narrative explanation of usually ad hoc assumptions and lacking a predictive
component (see Croizat, et al., 1974; Humphries & Parenti, 1986); (2) analytical
framevvorks with a predictive component such as vicariance or cladistic
biogeography (Humphries & Parenti, 1986); and (3) panbiogeography (Craw
& \\'eston, 1984). In all of these frameworks the concepts ofplate tectonics, and
PLATE TECTONICS AND MACROLICHEN DISTRIBUTION
51
of the amalgamation and accretion of terranes have considerable importance in
relating biotic distribution patterns and geological areas.
Plate tectonics
Plate tectonic theory (e.g. Hallam, 1973) views the upper 100 km
(lithosphere) of the earth as being divided into 6 major, and c. lO minor, rigid
plates which are in motion at speeds of several centimetres a year. Where two
plates move apart, new ocean floor is formed at a spreading ridge; where two
plates collide, one is consumed beneath the other with the production of a deep
ocean trench and a volcanically and seismically active volcanic arc; and where
two plates slide laterally past each other a major transcurrent or transform fault
system develops. Plate tectonics portrays continental blocks as passive passengers
riding on the backs of plates which may consist of both dry land and ocean
floor.
At the end of the Palaeozoic era, it is believed that there was one
supercontinent Pangaea, surrounded by superocean Panthalassa. In the early
Mesozoic the Tethys Sea divided Pangaea into northern Laurasia, and southern
Gondwanaland. The cool temperate, Panthalassic or proto-Pacific margin of
Gondwanaland is thought to have provided ancestors of the austral biota now
seen disjunctively distributed from southern Chile to Australasia and the
subantarctic islands. Fragmentation of Gondwanaland and the subsequent
movement of its component plates is now well known (e.g. Craddock, 1982;
Grindley & Davey, 1982). Initial Mesozoic rifting led to the separation of West
and East Gondwanaland, followed by the opening of the southern Atlantic and
the moving apart of South America and Africa, and the separation of India
from Australia-Antarctica. The New Zealand microcontinent separated from
the Panthalassic margin of Gondwanaland some 80 million years ago, taking
with it a southern cool temperate flora, no doubt including lichens, related to
that in southern South America by way of a vegetated, cool temperate West
Antarctica. Final stages in the break-up of Gondwanaland were the separation
of Australia from East Antarctica, the opening of the Drake Passage between
Antarctica and South America with the initiation of the circum-Antarctic
current and the West Wind Drift (see Galloway, 1988).
However, it is simplistic to view tectonic plates as Noah's Ark, transporting
relict vicariant remnants of a Mesozoic Gondwana biota, since recent geological
evidence challenges this simple view by showing that most of the circum-Pacific
continents and continental fragments such as New Zealand, are composite
geological formations (e.g. Craw, 1982, 1985). This has come about in the past
I 0 years by geologists recognizing tectonostratigraphic terranes, geological
formations which have important consequences for austral biogeography.
Tectonostratigraphic terranes
Terranes (correctly tectonostratigraphic terranes) are fault-bounded
geological entities varying in size from small fragments of crust to large
continental blocks and characterized by a geological history distinct from that of
neighbouring terranes (Howell, Jones & Schermer, 1985). They can be
displaced from a few hundred, to more than 6000 km. Accretion of terranes
0
D.
J
GALLOWAY
refers to the collision and welding of a terrane onto a continent, thus
introducing a differing geological structure into a pre-existing geological
pattern. ~ew Zealand's South Island, for example, is composed of nine terranes
(Bishop, Bradshaw & Landis, 1985; Howell, 1980); Andean South America
south of Ecuador, of at least three terranes accreted during the Palaeozoic
(Dalziel & Forsythe, 1985; Ramos et al., 1986); and the Pacific margin of
Antarctica is at least four discrete or semidiscrete microcontinents (Dalziel &
Elliot, 1982; Dalziel & Grunow, 1985). Nur & Ben-Avraham (1983) postulate
that circum-Pacific accreted terranes may have had an origin in a former
landmass, Pacifica, a presumed eastward extension of Gondwanaland beyond
Ne>v Zealand and Australia, the fragmented remnants of which are found
scattered in northern and western circum-Pacific continents. Terrane analysis
has therefore considerable implications for palaeogeography and explanations of
regional relationships in the Southern Hemisphere. Croizat's three generalized
tracks of relationship between Australasia and South America, viz. transAntarctic, trans-Pacific and Gondwanic via the Indian Ocean and Africa
Croizat, 1958), now have an added significance in austral biogeography
(Craw, 1985) in the light of plate tectonics and terrane accretion, which may
pro\'ide suitable geological explanations for the distribution of lichen genera in
such families as the Lobariaceae and Pannariaceae.
CO:'\CLCSIO:\'S
Distinctive relationships exist between the lichen floras of both New Zealand,
the subantarctic islands and Tasmania; and between Australasia and southern
South America at the level of narrative comparisons of cool temperate forests,
shrublands, subalpine bogs and grasslands between latitudes 35 and 55°S.
Australasian lichens have close links at both genus and species level, whereas
austral affinities are primarily at the generic level. Vicariance of a presumed
.Mesozoic cool temperate Gondwana lichen vegetation, with dispersal around
the Pacific in response to tectonic change as envisaged by plate tectonics, is an
over-simplification, since recent terrane analysis has shown both New Zealand
and southern South America, and Antarctica to be polyphyletic geological
composites.
The families Lobariaceae and Pannariaceae are widely represented in all
parts of the Pacific area, subantarctic, austral and palaeotropical; and with
increasing knowledge of Pacific Basin taxa, these families and their genera are
ideal subjects for biogeographical analysis. However, a prime requirement is still
a wide-ranging taxonomic study of Southern Hemisphere lichen genera and
species for, as Humphries ( 1983) points out, " .... biogeography is dependent
on the theoretical foundations of systematics since it can only be as good as the
taxonomy it uses to establish historical distribution. Biogeography must
therefore be regarded as an integral part of systematics; indeed, changes in
biogeographical explanations follow changes in taxonomic theory".
ACK:\'0\\'LEDGE~IE:\'TS
I am grateful to :\'lr P. W. James and Professor P. M. j0rgensen for
comments on the ideas expressed in this paper, and to the British Museum
/Natural History) for the provision of funds for fieldwork in southern Chile.
PLATE TECTONICS AND MACROLICHEN DISTRIBUTION
53
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