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© CSIRO 2001 10.1071/BT00023 0067-1924/01/03271
Aust. J. Bot., 2001, 49, 271–300
The breakup history of Gondwana and its impact on pre-Cenozoic
floristic provincialism
Stephen McLoughlin
School of Botany, University of Melbourne, Vic. 3010, Australia.
Email; s.mclo@unimelb.edu.au
Abstract. The concept of ‘Gondwana’, an ancient Southern Hemisphere supercontinent, is firmly established in
geological and biogeographical models of Earth history. The term Gondwana (Gondwanaland of some authors)
derives from the recognition by workers at the Indian Geological Survey in the mid- to late 19th century of a
distinctive sedimentary sequence preserved in east central India. This succession, now known to range in age from
Permian to Cretaceous, is lithologically and palaeontologically similar to coeval non-marine sedimentary
successions developed in most of the Southern Hemisphere continents suggesting former continuity of these
landmasses. Palaeomagnetic data and tectonic reconstructions suggest that the main assembly of Gondwana took
place around the beginning of the Palaeozoic in near-equatorial latitudes and that the supercontinent as a whole
shifted into high southern latitudes, allowing widespread glaciation by the end of the Carboniferous. From
Carboniferous to Cretaceous times the southern continents had broadly similar floras but some species-level
provincialism is apparent at all times. The break-up of Gondwana initiated during the Jurassic (at about 180 million
years ago) and this process is continuing. The earliest rifting (crustal attenuation) within the supercontinent initiated
in the west (between South America and Africa) and in general terms the rifting pattern propagated eastward with
major phases of continental fragmentation in the Early Cretaceous and Late Cretaceous to Paleogene. Gondwanan
floras show radical turnovers near the end of the Carboniferous, end of the Permian and the end of the Triassic that
appear to be unrelated to isolation or fragmentation of the supercontinent. Throughout the late Palaeozoic and
Mesozoic the high-latitude southern floras maintained a distinctly different composition to the palaeoequatorial and
boreal regions even though they remained in physical connection with Laurasia for much of this time. Gondwanan
floras of the Jurassic and Early Cretaceous (times immediately preceding and during break-up) were dominated by
araucarian and podocarp conifers and a range of enigmatic seed-fern groups. Angiosperms became established in
the region as early as the Aptian (before the final break-up events) and steadily diversified during the Cretaceous,
apparently at the expense of many seed-fern groups. Hypotheses invoking vicariance or long distance dispersal to
account for the biogeographic patterns evident in the floras of Southern Hemisphere continents all rely on a firm
understanding of the timing and sequence of Gondwanan continental breakup. This paper aims to summarise the
current understanding of the geochronological framework of Gondwanan breakup against which these
biogeographic models may be tested. Most phytogeographic studies deal with the extant, angiosperm-dominated
floras of these landmasses. This paper also presents an overview of pre-Cenozoic, gymnosperm-dominated, floristic
provincialism in Gondwana. It documents the broad succession of pre-angiosperm floras, highlights the distinctive
elements of the Early Cretaceous Gondwanan floras immediately preceding the appearance of angiosperms and
suggests that latitudinal controls strongly influenced the composition of Gondwanan floras through time even in the
absence of marine barriers between Gondwana and the northern continents.
Gon dwana break-up and pre-Cen o zoic floristic provincialism S. McLoughlinBT00023
Stephen McLoughlin
Introduction
This paper brings together palaeomagnetic, tectonic,
sedimentological, palaeontological and geochronological
data from a range of published sources to provide a summary
of the current state of knowledge concerning the breakup of
Gondwana and the changes in its terrestrial floras during
Palaeozoic and Mesozoic times. In the first part of the paper
the term Gondwana is defined. The second part deals with
the temporal constraints on the separation-age of various
landmasses that formerly constituted Gondwana. The third
section deals with pre-angiosperm floristic provincialism in
Gondwana using several key taxa, most notably Glossopteris
and Dicroidium for the Permian and Triassic periods,
respectively, to illustrate palaeoenvironmental controls on
the past distribution of plant groups. The final section
summarises the Cretaceous transition between gymnosperm-
and angiosperm-dominated floras of southern lands. This
272 S. McLoughlin
section also examines palaeoenvironmental factors affecting
the southern early angiosperm floras and highlights some
temporal problems for interpreting causal mechanisms that
resulted in the distribution of some key extant plant groups
(e.g. the Proteaceae, Nothofagaceae and Casuarinaceae) that
are traditionally regarded as Gondwanan.
Gondwana or Gondwanaland?
The term ‘Gondwana sediments’ was originally used to
denote a sequence of principally non-marine sedimentary
rocks exposed in a series of small, graben-type basins of
Peninsula India in unpublished work by Medlicott and
Oldham (staff of the Geological Survey of India) during the
1870s (Lele 1964; Maheshwari 1992). The term ‘Gondwana
floras’, therefore, related to the distinctive suites of fossil
plants preserved in those sediments. The earliest studies
mostly employed a two-fold subdivision of the Gondwanan
strata: the ‘Lower’ and ‘Upper’ Gondwanas for Permian
(Glossopteris-bearing) and Mesozoic (Ptilophyllum-
bearing) sequences, respectively (Oldham 1893; Cotter
1917; Fox 1931). Other workers (e.g. Feistmantel 1882;
Vredenburg 1910; Wadia 1957; Lele 1964) adopted a three-
fold subdivision of the Gondwanan sequence based primarily
on the recognition of distinctive Triassic, Dicroidium-
dominated, fossil floras in sediments stratigraphically
positioned between those bearing Glossopteris and
Ptilophyllum. This three-part scheme remains a useful
subdivision of the Indian Permian–Cretaceous succession
although geologists and palaeontologists now recognise
numerous distinct litho- and bio-stratigraphic units within
these sequences (Sukh-Dev 1987; Maheshwari 1992).
The terms ‘Gondwana’, ‘Gondwanan’ and ‘Gondwa-
naland’ derive from the ancient Kingdom of the Gonds
(a people inhabiting the area south of the Narmada River in
central India: Wadia 1957), where the typical Gondwanan
strata are well exposed. Gondwana has been translated from
Sanskrit to mean ‘land of Gonds’ (Maheshwari and Bajpai
1987) or alternatively ‘Gond forest’ (J. F. Rigby, pers.
comm.). Some workers regard this term as applicable only to
the region hosting the classical Permian–Cretaceous
sequences of India. They have employed the broader term
Gondwanaland (or ‘Gondwana Land’) to denote the
Southern Hemisphere supercontinent that existed from
Palaeozoic to mid-Mesozoic times (e.g. Wadia 1957;
Veevers 1984; Metcalfe 1991; Li and Powell 1993). Others
regard the name Gondwanaland as tautological and have
broadened the definition of Gondwana (and Gondwanan) to
apply to the entire supercontinent and the sediments and
fossil assemblages laid down during its existence
(Maheshwari and Bajpai 1987; Smith 1999). In this paper I
adopt the broader definition and use Gondwana throughout.
In early tectonic studies, the term Gondwana was applied
to the past assembly of the major modern Southern
Hemisphere landmasses (South America, Africa,
Madagascar, Australia, New Zealand and Antarctica)
bearing the distinctive sedimentary facies and fossil biotas
characteristic of the traditional Indian Gondwana sequence
(e.g. du Toit 1937). More recent geological studies have
suggested that several small continental terranes and
microplates that now make up parts of southern and eastern
Asia, southern Europe and Florida were at various times in
the past directly connected to, or closely associated with, the
core Gondwanan provinces. However, they do not always
bear characteristic ‘southern’ fossil biotas or sedimentary
facies (Sengör and Hsü 1984; Scotese and McKerrow 1990;
Metcalfe 1991). It is estimated that 90 or more continental
fragments (recognisable at the 2000-m bathymetric contour)
have been part of Gondwana at one time or another, and in
total these fragments cover an area of about 132 million km2
(Smith 1999). Additionally, it is now clear that some
Gondwanan terranes have extremely complex tectonic
histories. In some cases, they have undergone substantial
rotational or translational movements with respect to their
neighbouring landmasses (e.g. West Antarctic and New
Zealand terranes). In other cases (New Guinea, New
Caledonia and parts of New Zealand) these landmasses
represent compound tectonic entities comprised partly of
Gondwanan continental crust, partly of obducted ophiolitic
material and partly of accreted exotic terranes (Stevens
1980; Hall 1997).
Despite the complex histories of several marginal
terranes, the term Gondwanan has remained useful because
of the clear similarities recognised across the Southern
Hemisphere landmasses in many sedimentary successions,
volcanic episodes and tectonic events. Most significantly, the
high latitude occupied by Gondwanan terranes for much of
the late Palaeozoic and Mesozoic resulted in the distribution
of similar biotas across these landmasses. Although varying
degrees of provincialism are evident in the fossil biotas of
Gondwana through time, major extinction events are broadly
correlative across the ancient supercontinent and broad
similarities are evident in the fossil floras and faunas for each
geological period between the Carboniferous and
Cretaceous. Subsequent fragmentation of Gondwana
permitted divergence amongst progressively more isolated
populations and led to a greater degree of provincialism in
the biotas. Although the major stages in the breakup of
Gondwana occurred between 180 and 35 million years ago,
biogeographers dealing with extant biotas apply the term
Gondwanan to animal, plant and fungal groups having a
southern distribution with origins (cladogenesis) before or
during continental segregation. Evidence for a Gondwanan
origin around this time may be in the form of a fossil record
extending back to the Cretaceous. Alternatively, organisms
with extant distributions restricted to the Southern
Hemisphere are commonly suggested to have had an origin
in this region before continental dispersal. However, the
fossil record indicates that not all groups with an extant
Gondwana break-up and pre-Cenozoic floristic provincialism 273
Southern Hemisphere distribution need necessarily have
been restricted to that region through time. Araucarian
conifers are essentially restricted to southern landmasses at
present but in the Mesozoic they had a global distribution
(Stockey 1982). Their present distribution is probably
relictual following extensive extinctions associated with
Cenozoic climatic upheavals and perhaps competition from
angiosperms.
The broad geological framework of Gondwana
The formation of Gondwana
Fragmentation and amalgamation of continental blocks and
supercontinents is a continuing and cyclic process (the
Wilson Cycle: Dalziel 1997) involving lateral translation of
crustal blocks related to mantle convection and heat
exchange within the Earth. Gondwana was a composite
supercontinent formed by the fusion of at least five major
cratons (stable continental blocks) during the Neoprotero-
zoic to Cambrian (Li and Powell 1993; Unrug 1993). Its
formation followed the breakup of Rodinia, a late
Mesoproterozoic to early Neoproterozoic supercontinent
incorporating parts of what were later to become North
America, southern China and parts of Gondwana (Li et al.
1995). Debate continues with regard to the precise timing of
accretion of the various components of Gondwana and
particularly with regard to the structure and geographic
position of eastern and western Gondwanan terranes relative
to one another during the process of accretion. Nevertheless,
most workers agree that amalgamation of the core
Gondwanan provinces had occurred by the Early Cambrian
and that much of the supercontinent at that time was located
in equatorial latitudes (Scotese and McKerrow 1990;
Fig. 1a). Subsequently, Gondwana rotated into high southern
latitudes and by the Ordovician–Silurian (Fig. 1b) parts of
north-western Gondwana (northern Africa) experienced
extensive glaciation (Hambrey 1985). Gondwana continued
to rotate over the southern pole during the late Palaeozoic
with parts of central South America experiencing glaciation
in the Devonian and Carboniferous (Fig. 1c). Very extensive
glaciation affecting southern South America, southern
Africa, Arabia, India, Antarctica and Australia developed
during the latest Carboniferous and Early Permian (Crowell
1995; Fig. 2a). Glaciation appears to have persisted longest
in Australia where glendonites (cold-water carbonate
mineral pseudomorphs) and ice-rafted dropstones occur in
marine sediments of the Sydney Basin well into the Late
Permian (Crowell and Frakes 1971; Shi and McLoughlin
1997). This late Palaeozoic rotation over the South Pole left
a lasting signature on the southern landmasses in terms of the
prevailing sedimentary facies, the climatic regime for the
subsequent 200 million years and the morphological and
physiological adaptations of the biota occupying these high-
latitude environments.
Gondwana as part of Pangea
Gondwana and its Northern Hemisphere counterpart
Laurussia (comprising the Laurentian, Baltic, Khazakstan
and Siberian cratons) became welded together near the close
of the Palaeozoic to form the megacontinent Pangea rimmed
by a very broad ocean designated Panthalassa (Fig. 2a, b).
During the Permian–Jurassic additional terranes (e.g. the
North and South China blocks) accreted to the eastern
margin of Laurussia to form the large Northern Hemisphere
landmass Laurasia (Sengör 1984; Sengör and Hsü 1984).
During this time Laurasia and Gondwana were connected in
the west but separated in the east by a broad oceanic
embayment—the Tethys Sea (Fig. 2a–c). Although the core
cratonic provinces retained their relative positions, it is now
clear that several small continental blocks rifted away from
the northern margin of Gondwana during this interval
(Sengör and Hsü 1984; Fig. 2b, c). These microplates
eventually collided with the southern flank of Laurasia to
form what are now the heavily deformed terranes of the
southern Asian rim. Contemporaneously, several small
exotic terranes were accreting to the convergent
Panthalassan margin of Gondwana to form parts of eastern
Australia, New Zealand and West Antarctica (Wilson et al.
1989; Veevers et al. 1994). While much of Gondwana was
relatively stable from Cambrian to Jurassic times, minor
accretion and foreland basin rifting events led to intermittent
mountain-building, especially along the Panthalassan
margin (Veevers et al. 1994). From the Permian to mid-
Mesozoic, Pangea represented a very large landmass
extending from high northern to high southern latitudes
providing land-based pathways for biotic interchange
between most of the continental landmasses. However,
substantial climatic gradients existed between latitudinal
belts allowing the persistence of strong provincialism in the
Pangean biota. ‘Gondwanan’ has become almost
synonymous with the Southern Hemisphere or with an
isolated southern landmass. However, it is pertinent to note
that the classical and distinctive Indian Gondwana (Permian
to Early Cretaceous) sedimentary sequences and fossil biotas
were deposited while the southern continents were united to
those of the north as part of Pangea.
Breakup and dispersal of Gondwana
Assessment of vicariance or long-distance dispersal
hypotheses to explain Gondwanan floristic relationships
requires precise age constraints on both cladogenesis and
isolation of landmasses. Resolution of the timing of
cladogenesis requires detailed analysis of the fossil record, or
in the absence of a palaeobotanical record, molecular
divergence rates may in some cases provide a reliable proxy
for clade divergence. Where geographic isolation of
landmasses has occurred substantially earlier (on the order of
tens of millions of years) than differentiation of sister clades
274 S. McLoughlin
Siberia
Kazakhstania
North China
Australia
Antarctica
India
Africa
GONDWANA
RHEIC OCEAN
South
China
(C). Early Devonian
(390 Ma)
South America
PANTHALASSA
OCEAN
Laurentia
Baltica
EURAMERICA
PANTHALASSA
OCEAN
IAPETUS OCEAN
RHEIC
OCEAN
GONDWANA
(B). Middle Silurian
(425 Ma)
(A). Late Cambrian
(514 Ma) PANTHALASSA
OCEAN
IAPETUS OCEAN
Baltica
Siberia
Laurentia
India
Antarctica
Africa
South America
GONDWANA
Ancient land area
Modern
Suduction zone
Sea-floor spreading
Fig. 1. Early to mid-Palaeozoic continental reconstructions (after Metcalfe 1996; Scotese 1997). (A) Late
Cambrian reconstruction showing configuration of Gondwana, soon after amalgamation, extending between
low northern and high southern latitudes. (B) Middle Silurian reconstruction showing configuration of the
continents around the time of the oldest known vascular land-plant fossils. (C) Early Devonian
reconstruction showing the initial phase of rifting along the northern margin of Gondwana separating the
North and South China Blocks and opening the Palaeotethys Ocean.
Gondwana break-up and pre-Cenozoic floristic provincialism 275
Africa
South
America
India
Antarctica
Australia
PANTHALASSA
OCEAN
Sibumasu
Iran
Turkey
North
China
South
China
Indochina
Siberian Flora
(C). Early Triassic
(237 Ma)
PALAEOTETHYS
OCEAN
North
America
MESOTETHYS
OCEAN
Euramerican Flora
Cimmeria
East
Asian
Flora
Gondwana Flora
Middle Asian Flora
(A). Late
Carboniferous
(306 Ma)
PALAEOTHETHYS
OCEAN
PANTHALASSA
?
Land
Maximum Carboniferous-
Permian glaciation
(B). Late Permian
(255 Ma)
Cathaysian
Flora
Angaran
Flora
Southwest
United States
Flora
Transantarctic
Dulhuntyispora
Indo-
Lambert
G
o
n
d
w
a
n
a
n
F
l
o
r
a
West
Gondwanan
Subangaran
Flora
Euramerican
Flora
Patagonian
GONDWANA
PANGEA
LAURUSSIA
PANTHALASSA
Lhasa
Fig. 2. Late Palaeozoic to early Mesozoic continental reconstructions (after Scotese 1997). (A) Late
Carboniferous reconstruction showing the regional extent of the latest Carboniferous to Early Permian
Gondwanan glaciation (after Veevers and Powell 1987; Crowell 1995). (B) Late Permian reconstruction
showing global floristic provinces (modified from Meyen 1987; Cúneo 1996) and the initial stage of rifting
between the Cimmerian terranes and northern Gondwana opening the Mesotethys Ocean. (C) Early
Triassic reconstruction showing global floristic provinces (modified from Dobruskina 1994) and the
northerly translation of Cimmerian terranes closing the Palaeotethys Ocean.
276 S. McLoughlin
represented on those landmasses, then long-distance (trans-
oceanic) dispersal must be considered a possible factor in
explanations of the biogeographic history of the clades.
Alternative possibilities might include an incomplete fossil
record or inappropriate molecular substitution rates used to
assess the antiquity of the clade. Nevertheless, a precise age
framework for the breakup of Gondwana is necessary to test
alternative historical biogeographic scenarios. This section
briefly reviews the stages in Gondwanan breakup from the
Permian to Neogene.
Cimmerian terranes
South-east Asia is a composite landmass comprised of a
series of Gondwanan and Cathaysian continental and island–
arc fragments sutured together at various times between the
mid-Palaeozoic and Cenozoic (Hall 1996, 1997; Metcalfe
1996). The timing of the most recent continental fragments
to depart Gondwana, traverse the Tethyan embayment and
amalgamate with Asia, is potentially important to
biogeographic analyses of angiosperm and conifer groups
represented in the extant flora. Metcalfe (1996) identif ied
three major phases of terrane rifting along the Tethyan margin
of Gondwana. Some of these rift, drift and accretionary
events are reasonably well constrained by palaeomagnetic
and palaeontological data, others are inferential or
speculative (Metcalfe 1996). In all cases, sea-floor anomalies
have been destroyed by subduction during accretion to Asia.
Devonian rifting saw the separation of the North and South
China, Tarim, Indochina and Quidam blocks from north-
eastern Gondwana, creating the Palaeotethys Ocean. These
terranes drifted northwards and became sutured to northern
Asia during Late Devonian–Permian time (Metcalfe 1996;
Figs 1c, 2a, b). A second phase of rifting occurred during the
Late Carboniferous to Early Permian separating the
Cimmerian continent (Sengör 1984) from northern
Gondwana and opening the Mesotethys Ocean (Metcalfe
1996; Fig. 2b). The Cimmerian terranes (Sibumasu and
Qiantang) eventually accreted to southern Asia during the
Late Permian–Early Jurassic (Figs 2c, 3a). Permian–Triassic
sediments on the Cimmerian terranes host Gondwanan
marine invertebrate faunas but evidence for typical
Gondwanan floras is equivocal. A third phase of rifting
initiated in the Late Triassic–Late Jurassic separating the
Lhasa, West Burma (and possibly the Woyla) terranes from
north-eastern Gondwana and opening the Neotethys
(= Cenotethys) Ocean (Metcalfe 1996; Fig. 2a,b). After
northward drift and subduction of the Mesotethyan sea floor,
the Lhasa block sutured to southern Asia during the Early
Cretaceous and the West Burma and Woyla terranes were
accreted to South-east Asia by the Late Cretaceous.
Considering the early age for rifting of these terranes from
Gondwana (pre-Cretaceous), their potential role in ferrying
angiosperm taxa from Gondwana to Asia would appear to be
negligible. Their role as potential ‘stepping-stones’ for Asian
taxa to infiltrate Australia would also appear to be minimal
considering their wide separation from Australasia and
collision with Asia by the time of angiosperm diversification
in the Late Cretaceous.
East and We s t Gondwana
The major phase of Gondwanan breakup began during the
late Early Jurassic (about 180 million years ago). Breakup
was associated with the development of a series of deep-
seated mantle plumes beneath the extensive Gondwanan
continental crust (Storey 1995). Mantle plumes may be deep-
seated structures originating near the core–mantle boundary
or they may originate at relatively shallow depths in the
mantle as a response to thermal incubation beneath large
continents (Hawkesworth et al. 1999). Initial rifting between
East and West Gondwana was preceded by the emplacement
of extensive plume-related flood basalts in southern Africa
(at c. 182 million years ago) and the Transantarctic
Mountains (at c. 176 million years ago: Storey 1996). After
a short phase of intra-continental rifting, continental breakup
initiated with sea-floor spreading between Africa and
Madagascar in the Somali Basin during the Jurassic
magnetic quiet interval (c. 165 million years ago:
Rabinowitz et al. 1983). Madagascar drifted to the south-east
with respect to Somalia along the trend of the Davie Ridge
(Figs 3a, 4a). Spreading in the Somali Basin ceased during
the M9 magnetic interval (c. 130 million years ago) and since
that time Madagascar’s position has not changed
significantly with respect to Africa (Coffin and Rabinowitz
1988). Roughly contemporaneous with the opening of the
Somali Basin was seafloor spreading in the Mozambique
Basin and Weddell Sea, producing an en echelon fracture
system between East and West Gondwana. Although
geochronological constraints are not so well resolved in the
southern part of this fracture system, best estimates suggest
initial spreading in the Weddell Sea at c. 162 million years
ago (magnetic anomaly M29: Leitchenkov and Masolov
1997). The southernmost extension of this rift system may
have opened the Rochas Verdes Basin in southern Chile
(until about 130 million years ago; Fig. 4b) but rifting
apparently did not cause sufficient extension to develop fully
marine conditions between South America and the Antarctic
Peninsula at that time (Grunow et al. 1991; Storey 1996).
Africa–South America
The opening of the South Atlantic Ocean closely followed
the emplacement of the plume-related Parana–Etendeka
continental flood basalts (137–127 million years ago) in
Brazil and Namibia (Turner et al. 1994). Sea-floor
spreading began in the South Atlantic at about 135–130
million years ago (M11 magnetic anomaly: Jones 1987;
Barker et al. 1991; Grunow et al. 1991). However, physical
separation of the continents was probably not synchronous
along the line of rifting. Translational movement of Brazil
Gondwana break-up and pre-Cenozoic floristic provincialism 277
Antarctica
Africa
Australia
PACIFIC
OCEAN
North
America Europe
Asia
(C). Latest Cretaceous
(69.4 Ma)
India
South
America
NORTH
ATLANTIC
OCEAN
SOUTH
ATLANTIC
OCEAN
(B). Mid-Cretaceous
(94 Ma)
NEOTETHYS
OCEAN
Sibero-Canadian Flora
Palaeoequatorial
Flora
Gondwanan
Flora
European-Sinian Flora
Tasman
Patagonia-
Palmer
(A). Late Jurassic
(152 Ma)
Gondwana Flora
Palaeoequatorial Flora
European-Sinian Flora
Siberian Flora
West
East
Southern
African
Indo-Eromangan
MESOTETHYS
OCEAN
NEOTETHYS
OCEAN
PANTHALASSA
OCEAN
GONDWANA
LAURASIA
EASTERN
INDIAN
OCEAN
Fig. 3. Middle and late Mesozoic continental reconstructions (after Scotese 1997). (A) Late Jurassic
reconstruction showing initial rifting between east and west Gondwana, separation of Lhasa, West Burma
and Woyla terranes from northern Gondwana (Metcalfe 1996) and major Jurassic floristic provinces (after
Meyen 1987). (B) Early Late Cretaceous reconstruction shortly after isolation of Africa from other
Gondwanan landmasses, opening of eastern Indian Ocean and emplacement of Kerguelen Plateau basalts.
Cretaceous floristic provinces modified from Vakhrameyev (1984). (C) Latest Cretaceous reconstruction
shortly before eruption of the Deccan Traps showing progressive isolation of Gondwanan landmasses and
rapid northward migration of India.
278 S. McLoughlin
and equatorial Africa along the Guinea Fracture Zone may
have maintained low latitude connections between the
continents until 119–105 million years ago (early Aptian to
early Albian: Jones 1987; Fairhead and Binks 1991; Binks
and Fairhead 1992). Similarly, transform faulting between
southernmost Africa and the easterly extension of the
Falklands Plateau may have maintained continental
connections or close proximity of southern Africa and
South America until c. 105 million years ago (Barron and
Harrison 1980; Barron 1987; Figs 3b,4a).
Madagascar–India–Seychelles Block
At about 95–84 million years ago a new phase of rifting
initiated in the proto-Indian Ocean separating Madagascar
from the Seychelles–India block (Plummer and Belle 1995).
India (including its northern extension now probably
underthrust beneath Tibet) had previously separated from
Australia and east Antarctica by the Hauterivian
(c. 132 million years ago; M10 magnetic anomaly: Barron
1987; Veevers and Li 1991; Fig. 4a). Relatively rapid sea-
floor spreading in the southern Indian Ocean saw the
Seychelles–India block move northwards into middle
latitudes by Late Cretaceous (Barron 1987). Eruption of the
Deccan flood basalts about 65 million years ago
accompanied a repositioning of the western Indian Ocean
(Carlsberg) spreading ridge and resulted in separation of
India and the Seychelles block. The Seychelles block
subsequently became fixed with respect to Africa while
India continued its rapid northward migration reaching
equatorial latitudes by the Eocene and colliding with
southern Asia about 43 million years ago (Barron and
Harrison 1980; Lee and Lawver 1995).
Kerguelen Plateau
The Kerguelen Plateau represents a large (c. 460 000 km2),
mostly submerged (to depths of 1000–2000 m) plateau in the
Antarctic sector of the Indian Ocean (Schlich et al. 1994). Its
structure and origin has aroused considerable debate
particularly with respect to whether it represents a hot-spot
related mound of oceanic basalt (comparable to Iceland or
the Ontong Java Plateau) or whether it is floored by
continental basement rocks (Schlich and Wise 1992).
Extensive seismic and drilling surveys of the region by the
Ocean Drilling Program indicate that the plateau is floored
by a thick sequence of Neocomian–Albian (130–110 million
years ago) plume-related basalts that were erupted near to the
junction of the Australian, Indian and Antarctic plates
(Royer and Coffin 1992; Figs 3b, 4a). Significant parts of the
plateau experienced subaerial erosion in the Early and early
Late Cretaceous (Schlich and Wise 1992; Schlich et al.
1994) and probably supported a diverse terrestrial flora
during that time (Francis and Coffin 1992; Mohr and Gee
1992). Continual subsidence from the Cenomanian to
present saw inundation of almost the entire plateau and
deposition of a thick (up to 3000 m) sequence of marine
sediments (Schlich and Wise 1992). Despite its minor role in
supporting the present-day Southern Hemisphere terrestrial
flora, the Kerguelen Plateau was emplaced above sea level at
the time of the earliest radiations amongst angiosperms
(Crane and Lidgard 1989). Its large size and its
palaeoposition off north-eastern India (Royer and Coffin
1992) may have made it a crucial corridor for early
angiosperm dispersal to southern high latitudes by halving
the oceanic barriers between India and Australia–Antarctica.
We s t Antarctic terranes
West Antarctica consists of a series of structurally discrete
terranes with different geological histories. The principal
continental terranes include the Ellsworth–Whitmore
Mountains block, Antarctic Peninsula block, Thurston Island
block and Marie Byrd Land (Storey 1996). Of these areas
only the Antarctic Peninsula today supports a few small herb
and grass species. However, before mid- to late Cenozoic
glaciation these terranes may have played an important role
as a corridor for interchange of terrestrial vascular plants
between the Australian and South American regions. The
precise details of the tectonic development and relative
movements between the various West Antarctic terranes
remain poorly resolved owing to the difficulty in obtaining
geological data in an area of extensive ice cover.
Nevertheless, a broad understanding of the region’s history
can now be pieced together from the limited palaeomagnetic,
stratigraphic and geophysical data available. Most
controversy surrounds the position of the Ellsworth–
Whitmore Mountains block which contains a Cambrian–
Permian sedimentary succession that was deformed as part
of the Permian–Triassic Samfrau Orogeny (Collinson et al.
1994) but the deformational trend on this terrane is now at
approximately 90° to the structural trend of the adjacent
Transantarctic Mountains in East Antarctica. Recent
modelling suggests that the Ellsworth–Whitmore Mountains
block was originally positioned between the Falklands
Plateau and Queen Maud Land coast of East Antarctica as
part of the main Gondwanide Fold Belt before Gondwanan
breakup (Storey 1996; Fig. 4b). The 90° rotation of the
Ellsworth–Whitmore Mountains block probably occurred in
the later stages of the Samfrau Orogeny (c. 175 million years
ago) but before continental breakup (c. 165 million years
ago: Dalziel and Grunow 1992). Most of the remaining West
Antarctic microplates incorporate Mesozoic igneous suites
emplaced against, or intruded into, Proterozoic–Palaeozoic
metamorphic and igneous basement rocks. During the late
Palaeozoic to mid-Cretaceous they were all positioned along
a belt of convergence between East Antarctica and the
subducting Panthalassan oceanic plate (Collinson et al.
1994; Storey 1996). Although each of the West Antarctic
terranes has experienced independent rotation, deformation
and igneous processes, they have mostly retained their
Gondwana break-up and pre-Cenozoic floristic provincialism 279
Afr
EAnt
Mad
Ind
Aus
MBL
TI
SAm
FP
WS
RVB
162-130 Ma
AP
EMB
WM
MB
SB
CP
NZ NC
BSB
LHR
Collision with
Asia c. 43 Ma
c.130-
110 Ma
Drake Passage Powell Basin
DR
S
Afr
EAnt
Mad
Ind
Aus MBL
TI
SAm
FP
WS
RVB
AP
EMB
WM
SB
CP NZ
NC
BSB
LHR
KP
DR S
Southern Asia
LB
WB
WT
300-
280 Ma c. 210-
150 Ma
c. 260-
210
c. 145-
110 CT
CT
c. 110
-90
c.180-165 Ma
135-105 Ma
119-
105
Ma
165 Ma
162 Ma
165-
162 Ma
132 Ma
95-84 Ma
c. 132
Ma
65 Ma
KP
35-30.5 Ma
32-28 Ma
MB
Africa
Antarctic Peninsula
Australia
Byrd Subpolar Basin
Campbell Plateau
Cimmerian Terranes
Davie Ridge
East Antarctica
Ellsworth Mts Block
Falklands Plateau
India
Whitmore Mts
WT
Kergulen Plateau
Lord Howe Rise
Madagascar
Mozambique Basin
Marie Byrd Land
New Caledonia
New Zealand
Rochas Verdes Basin
Seychelles Block
South America
Somali Basin
Thurston Island Block
Wedell Sea
WB
Woyla Terrane
West Burma
Lhasa
LB
Guinea
Fracture
Zone
10 Ma
Subduction zone
Continental shelf
Rift/sea-floor spreading Time of separation
Time of collision
North America
Afr
SAm
Ind
Mad
EAnt
Aus
AP
S
0°
A
B
75°60°45°
Palaeoequator
Gondwanide Fold Belt
?
Fig. 4. Gondwanan breakup episodes. (A) Reconstruction of the South Atlantic–Indian Ocean–Neotethys Ocean regions during the late Early Cretaceous (110 million years ago)
showing the timing of separation and amalgamation of Gondwanan and Asian terranes (compiled from numerous sources—see text; base map modif ied from Scotese 1997).
(B) Polar projection of southern Gondwana at approximately 150 million years ago illustrating the relationships between west Antarctic terranes and showing the stages of initial
en echelon rifting between east and west Gondwana (after Grunow et al. 1991).
280 S. McLoughlin
relative positions with respect to one another since the
Triassic and have been tectonically locked to East Antarctica
since the Early Cretaceous (110 million years ago: Grunow
et al. 1991). The exception may be Marie Byrd Land, which
Grunow et al. (1991) suggested was separated from East
Antarctica and the remainder of West Antarctica during the
Jurassic and Early Cretaceous by rift and transtensional
embayments (e.g. Byrd Sub-polar Basin) several hundred
kilometres wide (Fig. 4b). At this time, however, Marie Byrd
Land was connected to Gondwana proper via southward
continuity of the Western Province terranes and Median Fold
Belt of New Zealand (Bradshaw et al. 1997). Convergence
between Marie Byrd Land and Thurston Island–East
Antarctica probably occurred during separation of the
former region from New Zealand and no significant relative
movement has occurred between Marie Byrd Land and West
Antarctica since 83 million years ago (C33 magnetic
anomaly: Weissel et al. 1977).
We s t Antarctica–South America
The earliest separation of South America and the Antarctic
Peninsula may have been in the Powell Basin area, which
developed in an area of back-arc extension from c. 35–30.5
million years ago (Lawver and Gahagan 1998). Subsidence
in the Powell Basin may have permitted a middle to deep
water opening between the continents slightly before sea-
floor spreading in the Scotia Sea saw establishment of the
deep marine Drake Passage during the Oligocene (c. 32–28
million years ago: Barker and Burrell 1977; Lawver and
Gahagan 1998; Fig. 4a). Final separation of West Antarctica
and South America permitted establishment of the South
Circumpolar Current and accompanied the development of
the first extensive ice sheets in Antarctica, although smaller
ice caps had probably been established in the region by the
Middle Eocene (Zachos et al. 1994; Lawver and Gahagan
1998). More intense glaciation probably occurred with
development of a more vigorous South Circumpolar Current
following closure of the Panama Isthmus and convergence of
Australia and South-east Asia in the Pliocene (Zachos et al.
1994).
East Antarctica–Australia
Crustal attenuation between Australia and East Antarctica
began during the Neocomian (c. 132 million years ago:
Ve ev e r s et al. 1991). Veevers and Li (1991) modelled the
inception of sea-floor spreading between Wilkes Land and
south-western Australia at c. 96 million years ago (within the
Cretaceous Magnetic Quiet Zone: Cenomanian; Fig. 5).
Rifting and sea-floor spreading propagated eastwards, and
crustal thinning on the South Tasman Rise had probably
allowed penetration of a shallow seaway between Australia
and Antarctica by the early Cenozoic (Lawver and Gahagan
1998). Complete separation of the continents by oceanic
crust was accompanied by dramatic falls in sea temperature
at about the end of the Eocene (c. 35.5 million years ago:
Shackleton and Kennett 1975; Veevers et al. 1991).
We s t Antarctica–Tasmantia
Tasmantia is the name applied to the large, but mostly
submerged, continental block incorporating New Zealand,
the Campbell Plateau, Chatham Rise, Lord Howe Rise,
Norfolk Ridge and New Caledonia (Fig. 5). Separation of
Marie Byrd Land and Tasmantia was preceded by a dramatic
shift from arc- to rift-related magmatism at about100 million
years ago (Storey 1996). The separation of these terranes is
well constrained to about 84 million years ago by the
identification of the C34 magnetic anomaly in oceanic crust
adjacent to the Campbell Plateau (Mayes et al. 1990; Storey
1996; Storey et al. 1999; Fig. 5).
Tasmantia–Australia
The sea-floor spreading patterns around Australia have been
thoroughly reviewed by Veevers et al. (1991) and Veevers
and Li (1991). The oldest tentatively identified pair of
magnetic anomalies in the Tasman Sea are A33y
(74.5 million years ago) located 110 km off the Tasmanian
continental–oceanic crust boundary. Veevers and Li (1991),
therefore, suggested sea-floor spreading initiated at
c. 80 million years ago. Evidence of a pulse of easterly-
sourced volcanigenic sediment in the Gippsland Basin
during the Barremian–early Aptian (c. 127–120 million
years ago) suggests that volcanism preceding Tasman Sea
breakup had initiated by the Early Cretaceous (Bryan et al.
1997). Tasman Sea spreading persisted until magnetic
anomaly A24y (55.5 million years ago: earliest Eocene) but
spreading rates were greater in the south than in the north.
This resulted in a wedge-shaped oceanic basin, wider
(c. 1500 km) in the south between Tasmania and the
Campbell Plateau than in the north (c. 370 km) between
central Queensland and the northern extremity of the Lord
Howe Rise (Veevers et al. 1991). Sea-floor structure is
poorly resolved in the region between the northern end of the
Lord Howe Rise and the Queensland Plateau but sea-floor
spreading in the adjacent Coral Sea has been constrained by
magnetic anomalies to 63.5–55.5 million years ago (Veevers
and Li 1991). Little geological data is available for the
basement or lower post-breakup sediments on the Lord Howe
Rise but crustal extension and thinning during breakup
(Bradshaw 1991) probably led to rapid marine inundation of
this terrane during the latest Cretaceous.
New Zealand–New Caledonia
New Zealand and New Caledonia are linked by attenuated
continental crust along the Norfolk Ridge (Sutherland 1999).
Both areas represent composite landmasses formed by
amalgamation of diverse tectonic terranes during Palaeozoic
to Cenozoic convergence along the Panthalassan/Pacific
margin of Gondwana. New Zealand incorporates at least
Gondwana break-up and pre-Cenozoic floristic provincialism 281
eight distinct basement terranes (Howell 1980; Cooper 1989;
Sutherland 1999). Early Palaeozoic terranes of the Western
Province are geologically akin to the Lachlan Fold Belt of
eastern Australia (Cooper and Tulloch 1992). Late
Palaeozoic–Mesozoic arc and accretionary wedge terranes
of the Eastern Province were sutured to the Western Province
along the line of the Median Tectonic Zone—a suite of
Carboniferous–Cretaceous arc-related igneous rocks
(Howell 1980; Sutherland 1999). Deformation along this belt
ceased at about mid-Cretaceous (c. 110-100 million years
ago) at the close of the Rangitata Orogeny (Knox 1982;
Bradshaw 1991; Chamberlain et al. 1995). The Permian–
Jurassic Teremba terrane of western New Caledonia is a
correlative of the Murihiku terrane of New Zealand
(Aitchison et al. 1995). The central axis of New Caledonia is
composed of the Permian–Jurassic metasedimentary Bohgen
terrane and Lower Triassic–Jurassic island-arc and deep
marine sedimentary rocks of the Koh terrane are emplaced
Aus
LHR
LHI
(6.9 Ma)
NIs
(3.05 Ma)
NR
TS
SFB
NFB
NC
V
F
Samoa
Sol
OJP
WB
CS
QP
NG NBNIr
Bor Sul
Banda
Sea
T
EAnt
STR
ETR
GSB
BT
CR
CHI
BI
AnI
CI
AuI CP
Louisville
seamount
chain
To
Argo Land
(W. Burma?)
TB
96 Ma
4 Ma
40-27
Ma
7 Ma 5 Ma
25 Ma
10 Ma
15 Ma
(emplaced 122-90 Ma)
25-4 Ma
35.5 Ma
c. 80 Ma
85-75 Ma
NCB
3.5 Ma
Greater
India
(132 Ma)
84 Ma
(from Marie
Byrd Land)
Sea-floor spreading (active)
Sea-floor spreading (extinct)
Subduction zone
Time of separation
Time of collision
Present land area
2000 m isobath
120E 140E 160E 180 0
20S
40S
60S
Sorong Fault
63.5- 55.5 Ma
158.6
Ma
Marine incursions
c. 120 Ma, 40 Ma, 22 Ma
Frigidity barriers
(< 35 Ma)
Photoperiod barriers
(>250 Ma)
NZ
LHS
TSC
South Circumpolar
Current established
c. 35-28 Ma
GB
20-5 Ma
Aridity barriers
c. 30 Ma
Fig. 5. Modern distribution of geological terranes in the Australasian region showing the time of initial
separation via sea-floor spreading, emplacement ages of selected basaltic islands and timing of terrane collisions
(base map adapted from Burrett 1991; Veevers and Li 1991; Sutherland 1999; age constraints from numerous
sources—see text). Abbreviations on map: AnI, Antipodes Islands; AuI, Auckland Island; Aus, Australia;
BI, Bounty Island; Bor, Borneo; BT, Bounty Trough; CHI, Chatham Islands; CI, Campbell Island; CP, Campbell
Plateau; CR, Chatham Rise; CS, Coral Sea; EAnt, East Antarctica; ETR, East Tasman Rise; F, Fiji;
GB,Gippsland Basin; GSB, Great South Basin; LHI, Lord Howe Island; LHR, Lord Howe Rise; LHS, Lord
Howe seamount chain; NB, New Britain; NC, New Caledonia; NCB, New Caledonia Basin; NFB, North Fiji
Basin; NG, New Guinea; NIr, New Ireland; NIs, Norfolk Island; NR, Norfolk Ridge; NZ, New Zealand;
OJP, Ontong Java Plateau; QP, Queensland Plateau; SFB, South Fiji Basin; Sol, Solomon Islands; STR, South
Tasman Rise; Sul, Sulawesi; TB, Taranaki Basin; To, Tonga; TS, Tasman Sea; TSC, Tasmantid seamount chain;
V, Vanuatu; WB, Woodlark Basin.
282 S. McLoughlin
along the island’s eastern flank. Similarities between the
New Caledonian and New Zealand basement terranes
suggest post-Permian structural continuity along the Norfolk
Ridge (Sutherland 1999) and this is supported by limited
data from sea-floor dredging (Mortimer et al. 1998).
Subduction-related volcanism occurred along the ridge at
least during the Late Oligocene and more recent (Pliocene)
basaltic volcanism led to the emergence of Norfolk and
Philip Islands. However, crustal thinning of the Lord Howe
and Norfolk Ridges occurred during the extensional tectonic
phase associated with opening of the Tasman Sea and New
Caledonia–Taranaki Basin in the Late Cretaceous (85–75
million years ago: Knox 1982; Veevers and Li 1991;
Sutherland 1999) and both ridges (now with crest depths
typically >1000 m below sea level) have probably been
covered by shallow seas since the end of the Cretaceous.
Marine sediments are known from the crests of both ridges
by Paleocene–Lower Miocene times (Coleman and Veevers
1971; Bentz 1974). New Caledonia underwent subduction
and/or accretion-related deformation along the convergent
Panthalassan margin of Gondwana during the Permian–
Jurassic (Aitchison et al. 1995). Upper Cretaceous coal
measures locally overlie basement rocks suggesting that
most of the area was emergent during the Mesozoic.
However, from the Late Cretaceous (Campanian) to Early
Eocene, a blanket of progressively deeper water marine
sediments was deposited over much of the island suggesting
substantial, if not total, submergence (Lillie and Brothers
1970; Aitchison et al. 1995). Collision between New
Caledonia and a north-easterly trench–island–arc system in
the Middle–Late Eocene saw obduction (overthrusting) of
two major ophiolitic nappes across the New Caledonian
basement rocks (Aitchison et al. 1995), although alternative
models involving subduction of New Caledonia Basin
oceanic crust to the south-west of the island have also been
proposed (Rawling and Lister 1999). Crustal thickening and
uplift associated with ophiolite emplacement has probably
maintained New Caledonia as a large island mass since the
Late Eocene.
Isolated continental shelf islands
Several small landmasses with Palaeozoic–Mesozoic
Gondwanan basement rocks (in some cases intruded and
capped by Cenozoic volcanics) exist, isolated by marine
environments, on the continental shelves of the major
Gondwanan continents. Examples of these isolated
continental islands include the Falkland Islands, Sri Lanka,
Tasmania and the Chatham, Campbell, Antipodes and
Auckland islands on the Campbell Plateau, south-east of
New Zealand. The Falkland Islands occur on an easterly
extension of the Patagonian continental shelf. The basement
geology of the Falklands has more in common with
deformed Palaeozoic rocks of the Cape Fold Belt in southern
Africa than it has with neighbouring areas of Patagonia.
About at the end of the Permian–Triassic Samfrau Orogeny,
the Falkland Islands underwent approximately 100° of
rotation and some south-westerly translation but since the
mid-Mesozoic its position has been fixed with respect to
South America (Dalziel and Grunow 1992). It has probably
been isolated from mainland South America since at least the
mid-Cenozoic, at which time marine sediments of the
Chenque Formation were deposited over most of Patagonia
(Bellosi 1990).
Sri Lanka is separated from India by the narrow and
shallow Palk Strait. Its close proximity to mainlaind India
and the shallow intervening continental shelf suggest that
these landmasses were probably in full connection as
recently as the last major sea-level rise 6000 years ago.
Similarly, Tasmania is separated from mainland Australia
by the shallow continental shelf of Bass Strait. These
landmasses may have been connected via the Bassian Rise
(a basement high only 15–20 m deep along the eastern
side of Bass Strait) as recently as 9000–6500 years ago,
prior to establishment of the post-glacial sea-level
maximum (Hope 1973; Blom 1988). The Queensland
Plateau, a detached region of northeastern Australian
continental shelf (Fig. 5) may have been exposed as an
extensive area of land during Quaternary sea level minima
but the area has been mostly inundated and separated from
the mainland since Oligocene subsidence and marine
sedimentation in the Queensland and Townsville Troughs
(Day et al. 1983).
Age constraints on isolation of the small islands of the
Campbell Plateau and Chatham Rise are more difficult to
quantify. Several of these islands represent the locations of
diachronous Late Cretaceous to Quaternary volcanic activity
(Hay 1978; Adams et al. 1979; Adams 1981, 1983;
Campbell et al. 1988) and each island group may have been
emergent since emplacement of the volcanics. However,
these areas have probably been isolated from southern New
Zealand by late Early Cretaceous–Miocene crustal thinning
in the area of the Great South Basin and Bounty Trough
(Carter 1988; Herzer and Wood 1992; Fig. 5). Marine
sedimentation in these areas was established by Paleocene–
Eocene times and they are presently represented by sea-floor
depths of > 500 m (Katz 1982) suggesting that even dramatic
sea-level falls of 100–175 m during the Pleistocene (Veeh
and Veevers 1970; Rohling et al. 1998) would not have
connected the outer shelf islands to mainland New Zealand.
The Chatham Islands may have undergone complete
submergence during an Early Oligocene transgression
(Wood et al. 1989) suggesting Neogene recolonisation of the
isolated landmass. Additionally, the main islands of New
Zealand were probably an area of relatively low relief during
most of the Cenozoic (prior to uplift of the Southern Alps in
the Pliocene, <5 million years ago: Chamberlain et al. 1995).
High sea-level stands, particularly during the Oligocene,
probably subdivided New Zealand into several smaller areas
Gondwana break-up and pre-Cenozoic floristic provincialism 283
intermittently separated by shallow seas (Stevens and
Suggate 1978).
Northern New Guinea
Following separation from Antarctica in the Cretaceous and
Paleogene, Australia has moved northwards into middle
latitudes (Veevers et al. 1991), resulting in extensive
subduction of oceanic crust to the north of the continent.
Australia’s leading edge began collisions with exotic island–
arc terranes of the Philippines Sea Plate during the
Oligocene (c. 25 million years ago: Crowhurst et al. 1996;
Hall 1997; Fig. 5). However, deformation, metamorphism
and obduction of local ophiolitic complexes had been
occurring along the New Guinean margin since at least the
Paleocene (Hill and Raza 1999). Translational movements
along the Sorong Fault system in northern New Guinea may
have subsequently (20–5 million years ago) transferred
slivers of the northern Australian continental margin
westward as far as Suluwesi (Hall 1996; Fig. 5). Australia
has progressively closed on the Sundaland (South-east
Asian) continental crust since the Oligocene so that the
direct separation is now only about 1000 km. That distance
is further reduced by a series of intervening island–arc
terranes so that, during periods of Quaternary eustatic sea-
level lowstands, the maximum marine separation of
terrestrial environments between the continents was
probably not more than 150 km.
Melanesian terranes
The complex geology of the Melanesian region incorporates
mostly island–arc terranes, plume-related basaltic plateaux
and small back-arc basins initiated by interactions between
the Pacific and Indo-Australian plates during the Cenozoic.
The Solomons, Vanuatu, Fiji and Tonga island–arc systems
may have initiated as early as the Late Cretaceous with
extension in eastern Gondwana and collision with the Pacific
Plate (Petterson et al. 1999) but exposed rocks on these
terranes are in most cases no older than Late Eocene (Colley
and Hindle 1984). Island–arc development in this region
occurred in two main phases. First, an Eocene–Middle
Miocene stage was associated with a simple arc–trench
convergent system (Colley and Hindle 1984). Second, a
Middle or Late Miocene to Holocene stage of arc
reorganisation was associated with collision of the Ontong
Java Plateau in the Solomons area, substantial transform
faulting, relocation of subduction zones and major phases of
back-arc spreading (Colley and Hindle 1984; Burrett et al.
1991; Petterson et al. 1997, 1999; Hill and Raza 1999;
Fig. 5). Throughout the Cenozoic these arc-related terranes
probably represented a chain of isolated islands and at no
time did they form a continuous land bridge between New
Guinea and the continental fragments of New Caledonia and
New Zealand to the south-east.
Oceanic islands
Predominantly basaltic oceanic islands are chiefly generated
by hot-spot related volcanism sourced deep within the
mantle. Oceanic crust over-riding hot spots may host a trail
of basaltic islands having progressively greater ages with
distance from the currently active volcano. With age, many
of these islands are reduced to submarine seamounts and
guyots owing to subaerial erosion of the crests, isostatic
equilibration and thermal subsidence of the surrounding
oceanic crust. The Hawaiian–Emperor seamount chain in the
north-western Pacific represents the best-studied chain of
this type, but equivalent island chains are represented in all
major ocean basins (Morgan 1971). The islands produced by
hot-spot volcanoes are typically small and in most cases
geologically short-lived, although the hot-spot magmatic
source may persist for >70 million years (in the case of the
Hawaiian chain: Claque and Dalrymple 1987). Landmasses
of this type lack a granitic chemistry and in a genetic sense
are unrelated to continental terranes even where, in cases like
Lord Howe and Norfolk islands, they may have been
emplaced through attenuated continental shelves. They are
not herein considered to be elements of Gondwana sensu
stricto but should be regarded as discrete oceanic island
entities. Of pertinence to this study are the north–south
orientated Tasmantid and Lord Howe seamount chains in the
Tasman Sea (Fig. 5), which represent southerly-younging
hot-spot related volcanics emplaced from >24 to 6.4 million
years ago (McDougall and Duncan 1988; Eggins et al.
1991). Many of these seamounts had peaks above sea-level
for periods shortly after their emplacement (judging from
their eroded crests) but most are now submerged to depths of
c. 1000 m and at no time during the Cenozoic would they
have provided continuous land bridges or even close-spaced
island ‘stepping stones’ for the interdispersal of organisms
across the Tasman Sea. Norfolk Island, composed mostly of
basalts emplaced at 3.05–2.3 million years ago, has similarly
been isolated from other landmasses throughout its history.
Other south Atlantic, Indian and Southern Ocean Islands
such as Ascension, Saint Helena, Tristan da Cunha, Gough,
Bouvet, Prince Edward, Crozet and Amsterdam islands are
geologically young islands located on active crustal
spreading ridges and have not been associated with
continental landmasses since opening of the respective ocean
basins.
Significance
From a biogeographic perspective the Jurassic–Holocene
history of Gondwana has commonly been viewed as a simple
sequence of diverging terranes, hosting progressively more
isolated and distinctive biotas through time. However, the
breakup history of Gondwana should be considered as a
reticulate and multidimensional pattern of separating and
amalgamating landmasses, fluctuating climates, emergence
284 S. McLoughlin
of terrane-linking island arcs, intermittent orogenesis,
sporadic marine transgressions and regressions and a
changing mosaic of soil substrates. Biogeographic studies of
extant organisms aiming to erect or test models for
Gondwanan terrane dispersal should take these factors into
account and assess the reticulate history of this province
rather than simple dichotomous models of terrane
separation. The following section examines the broad pattern
of floristic provincialism that developed in relation to
Gondwanan and Pangean amalgamation and breakup
through the late Palaeozoic and Mesozoic.
Pre-angiosperm Gondwanan floras
The pre-angiosperm floras are broadly similar across
Gondwana for each period before the Cretaceous largely as a
consequence of the united landmass occupying high
southern latitudes for much of its history. At most times, the
Gondwanan floras had a composition distinct from those of
Northern Hemisphere landmasses. Two key plant groups,
glossopterids and corystosperms, are used as case studies to
illustrate strong latitudinal controls on the distribution of
Gondwanan plant groups under different global
palaeoclimatic regimes in the Permian and Triassic. Despite
the existence of broadly similar floras across Gondwana
during much of the Palaeozoic and Mesozoic, varying
degrees of intra-Gondwanan floristic provincialism are
evident from the Devonian to Cretaceous. The Gondwanan
floras also show short intervals of extinctions separating
longer intervals of radiations and generic uniformity. These
extinction events probably had multiple causes but most
appear to have incorporated major climate changes.
Fundamental restructuring of the vegetation following
extinction events permits categorisation of the floras into
five major phases of development before the appearance of
the angiosperms. The principal plant groups characteristic of
these floristic phases and the palaeoenvironmental factors
influencing their distribution are summarised below.
Cosmopolitan early land plant floras (Silurian–Early
Devonian)
The oldest vascular land plant fossils occur in Lower Silurian
rocks (Edwards and Fanning 1985), although some dispersed
spores, cuticles and tracheid-like tubes have occasionally
been reported from older rocks (Richardson 1985; Taylor
1988). Upper Silurian to Lower Devonian floras
incorporating simple rhyniophyte, zoosterophyll, trimero-
phyte and lycopod fossils occur sporadically throughout
Gondwana (Tims and Chambers 1984; Anderson and
Anderson 1985; Morel et al. 1995). The geographic
distribution of the earliest land plant fossils suggests that
they arose or were most prevalent in low latitudes. Several
early land plants (especially Zoosterophyllum and the
lycopod Baragwanathia) are represented on both Southern
and Northern Hemisphere continents indicating little
provincialism in the early floras (Tims 1980; Tims and
Chambers 1984; Meyen 1987; Morel et al. 1995). Silurian
and Early Devonian plants were free-sporing, herbaceous
forms occupying moist habitats (coastal areas, river flats,
delta swamps, lake margins) as a requirement for motile
gamete exchange.
Arborescent lycopod and seed-fern floras in a cooling
climate (Middle Devonian–Carboniferous)
By the Late Devonian the north-western part of Gondwana
had rotated into polar latitudes (Fig. 1c) and parts of central
and northern South America were affected by glaciation
(Veevers and Powell 1987). Eastern parts of Gondwana
remained in middle latitudes during this time, allowing
development of extensive stromatoporoid–coral–cyano-
bacterial reef systems (Playford 1984). This continental
movement and strong latitudinal gradient across the
landmass saw the initial development of a distinctive
Southern Hemisphere flora (Meyen 1987) and also the first
expression of intra-Gondwanan floristic provincialism.
Subsequently, in the Carboniferous, the rotation of eastern
parts of Gondwana into polar latitudes and the return of parts
of western Gondwana into mid-latitudes (Figs 1c, 2a) helped
maintain a distinctive southern flora but with a different
profile of intra-Gondwanan floristic provincialism.
Middle Devonian floras were mostly dominated by
herbaceous to shrub-sized lycophytes attributed to
Haplostigma, Leclercqia, Archaeosigillaria and Proto-
lepidodendron. The Middle and Late Devonian also saw the
emergence of the first arborescent plants: lepidodendralean
lycophytes and progymnosperms. By the end of the
Devonian (Famennian) lycophytes such as Bumbudendron
and Leptophloeum had attained the stature of tall trees
(perhaps up to 20 m) and were important colonists of coastal
floodplains (Gould 1975). Late Carboniferous sedimentary
sequences of North America and Europe incorporate thick
coal deposits derived largely from lycophytes, sphenophytes,
cordaitaleans and medullosan seed ferns adapted to growth
in extensive tropical mires. Some thin Carboniferous coal
seams exist in eastern Australia and South America
(Rattigan 1964; Azcuy 1985), but coal development at this
time in Gondwana was minor compared with the Northern
Hemisphere and relates to the unfavourable climates and
geological settings for the accumulation of thick mire
deposits.
With the advent of cooler climates over much of
Gondwana during the Carboniferous, seed-fern floras
progressively replaced lycophyte floras. The dominant early
seed ferns, such as Diplothema, Eusphenopteris,
Eonotosperma, Nothorhacopteris, Fedekurtzia, Austrocalyx
and Botrychiopsis (Césari 1987, 1997; Césari and Garcia
1988; Cúneo 1990; Vega and Achangelsky 1996) had
possible calamopityalean or buteoxylonalean affinities
(Taylor and Taylor 1993). Glaciation reached its acme in
Gondwana break-up and pre-Cenozoic floristic provincialism 285
Gondwana during latest Carboniferous to earliest Permian
times (Crowell 1995; Fig. 2a). Retallack (1980) likened the
Gondwanan low-diversity Botrychiopsis-dominated floras of
this time to tundra vegetation but few detailed
palaeoenvironmental studies have been carried out on these
fossil biotas.
The glossopterid flora (Per mian)
Glossopteris, the spathulate, reticulate-veined leaves
produced by an enigmatic group of extinct gymnosperms are
the principal diagnostic fossil of Gondwanan Permian
terrestrial deposits and its distribution was one of the first
pieces of palaeontological evidence used to support the early
notions of an ancient Southern Hemisphere landmass
(du Toit 1937). Glossopterids first appear in the fossil record
at about the end of the Carboniferous in West Gondwana and
disappear from all continents along with about 80% of the
world’s biota during the Permian–Triassic extinction (Erwin
1995). By Permian times, most of the world’s continental
blocks were united into the megacontinent Pangea
(Fig. 2a,b). However, the strong equator to pole climatic
gradients imposed by Gondwanan glaciation resulted in
pronounced provincialism in the world’s floras. Glossopteris
was distributed across all Gondwanan landmasses between
40° and 90° palaeolatitude in the Permian (Fig. 6). Only
northern African and northern South American parts of
Gondwana may have hosted non-glossopterid floras (Broutin
et al. 1995). Glossopterids are not represented in areas
outside Gondwana with the possible exception of Turkey and
Thailand (Kon’no 1963; Archangelsky and Wagner 1983),
although these may be Cimmerian terranes that rifted away
from Gondwana proper during the Permian (Sengör et al.
1984; Fig. 4a). Voltzialean conifers, cordaites and
peltaspermalean, medullosan and gigantopterid seed-ferns
dominated palaeoequatorial and Northern Hemisphere
Permian floristic provinces (Fig. 2b).
Glossopterid remains dominate the Permian floras at
virtually all fossil localities across Gondwana. Their success
was largely due to their deciduous habit and the development
of specialised root systems enabling them to colonise the
extensive high-latitude, lowland, lake-margin and mire
environments following the retreat of the Permian ice sheets.
Only in South America, which had rotated into lower (and
warmer) latitudes during the Early Permian, are there
significant assemblages dominated by lycophytes and
conifers (Cúneo 1996). Subsidiary elements of the flora
(sphenophytes, ferns, cycadophytes, conifers, cordaites) are
also relatively uniform at generic level across Gondwana in
the Permian (Rigby 1972). Although the floras show broad
similarities, differences at species level amongst most
groups indicate that significant provincialism was
maintained during the Permian (Cúneo 1996; Figs 2b, 6).
IRAN
ARABIA
CENTRAL EUROPE
IBERIA
SOUTH
AMERICA
INDIA
?
EAST
ANTARCTICA
AFRICA
AUSTRALIA
N.Z.
QIANTANG
SIBUMASU
FLORIDA
Late Permian Pole
?
??
TURKEY
LHASA
MAD
?
?
0°30°
60°
CIMMERIAN
TERRANES
?
Glossopteris
macroflora
Possible glossopterids
Limit of glossopterids
Dulhuntyispora
spores
Reworked
Dulhuntyispora
Permian basins
Continental shelf margin
Fig. 6. Palaeolatitudinal distribution of the Permian Glossopteris flora and distribution of the principal
occurrences of Dulhuntyispora (cheilocardoid fern spores) in Gondwana (base map from Lawver and
Scotese 1987; fossil data from numerous sources).
286 S. McLoughlin
The Dicroidium flora (Triassic)
Gondwanan floras were fundamentally restructured at the
end of the Permian. The Late Permian glossopterid-rich mire
floras were replaced in the Early Triassic by floras dominated
by newly diversified groups such as peltasperms,
corystosperms, pleuromeian lycophytes and voltzialean
conifers. Although a large proportion of Gondwana
remained at high latitudes, the new floras initially occupied
a drier and probably warmer climatic regime, indicated by
the absence of coals anywhere on Earth and the widespread
development of red-beds at this time (Retallack et al. 1996;
McLoughlin et al. 1997). Although the Permian–Triassic
floristic turnover occurred over a relatively short period of
time, there is some evidence that the transition was
diachronous across the supercontinent with the Glossopteris
flora declining earlier in lower latitudes and persisting latest
in southern Gondwana (McLoughlin et al. 1997).
Gondwanan Triassic floras are consistently more diverse
than Permian floras (Anderson et al. 1999). By the Late
Triassic, coal-forming mire communities had become re-
established in several Gondwanan basins. The floras are best
known from the extensive studies of the Molteno Formation
of South Africa (Anderson and Anderson 1983, 1985, 1989).
Although most of the Late Triassic floras are dominated by
Dicroidium (Corystospermales; Fig. 7), voltzialean conifers,
ginkgophytes, peltasperms, putative gnetaleans, bennetti-
taleans, pentoxylaleans and cycadophytes, together with a
range of seed ferns of uncertain alliance, are all well-
represented. Herbaceous elements of the flora include a
range of lycophytes and osmundacean, gleicheniacean,
dicksoniacean, dipteridacean and marattiacean ferns
(Walkom 1915, 1917a, 1917b; Retallack 1975, 1977;
Holmes and Ash 1979). Triassic macroplant suites show
distinct facies controls on taxonomic composition. Several
ecostratigraphic assemblages representing different
vegetation types associated with distinct palaeosols have
been recognised in fluvial tracts of Australia and South
Africa (Retallack 1977; Anderson et al. 1989). Provincialism
within the Gondwanan Triassic flora remains poorly
understood due to a dearth of studies from some regions and
difficulties in achieving precise age controls on some
terrestrial sequences. Nevertheless, some taxa (especially
Dicroidium species) were clearly widespread (Anderson and
Anderson 1983, 1989) signaling broadly comparable
vegetation types over most of the supercontinent. Dicroidium
occupied high latitude parts of Gondwana in the Triassic
comparable to the area occupied by Glossopteris in the
Permian (Fig. 7) suggesting similar adaptation to strongly
seasonal climates. The most significant variation within the
ARABIA
CENTRAL EUROPE
IBERIA
SOUTH
AMERICA
INDIA
EAST
ANTARCTICA
AFRICA
AUSTRALIA
N.Z.
FLORIDA
TURKEY
LHASA
MAD
0°
30°
60°
Triassic Pole
Dicroidium
macroflora
Ipswich palynoflora
Onslow palynoflora
Intermediate palynoflora
Triassic basins
Continental shelf margin
Limit of
Dicroidium
Fig. 7. Palaeolatitudinal distribution of the Triassic Dicroidium flora and the principal occurrences of
‘Onslow-type’, ‘Ipswich-type’ and ‘Intermediate-type’ palynofloras (base map from Lawver and Scotese
1987; Powell and Li 1994, with additional data from Anderson and Anderson 1983; Foster et al. 1994).
Gondwana break-up and pre-Cenozoic floristic provincialism 287
Gondwanan Triassic flora is evident in spore-pollen
assemblages. Late Triassic ‘Onslow-type’ palynofloras of
northern Australia, India, northern Madagascar and east
Africa contain Tethyan (equatorial) elements that are absent
in southern Australian, Antarctic, South African, southern
Madagascan and South American ‘Ipswich-type’
assemblages (Dolby and Balme 1976; Foster et al. 1994;
Fig. 7). Milder, maritime climates may have been associated
with the northern margin of Gondwana (flanking the
palaeoequatorial Tethys Ocean) allowing differentiation of
the floras from those occupying more seasonal and cooler
continental and polar climates in southern Gondwana.
Jurassic pteridosperm-conifer floras
Gondwana occupied mostly middle to high (30–80°)
latitudes in the Jurassic (Smith et al. 1981), although
northern South America and northern Africa extended to the
equator. This configuration probably resulted in a
considerable climatic gradient across the supercontinent.
Although good palaeoclimatic indicators are often lacking,
stable isotope studies, the absence of red-beds, the presence
of local coal deposits and the lack of evidence for glaciation
suggests that conditions were generally warm and, on the
whole, relatively humid in south-eastern Gondwana during
the Jurassic. Parts of South America and South Africa (then
in low and middle latitudes) experienced arid conditions
(Soares and Assine 1992). The global scarcity of glacial
deposits (Woolfe and Francis 1991) suggests that warm
conditions prevailed world-wide at that time.
In Gondwana, the end of the Triassic was marked by a
floristic turnover of a magnitude equivalent to that of the
end-Permian extinction. Conifer- and bennettitalean-
dominated communities replaced Dicroidium-dominated
floras across the Southern Hemisphere. The floristic
turnover coincided with a major global extinction event
affecting terrestrial vertebrates and marine invertebrates
(Stanley 1987). Jurassic floras have not been well studied in
Gondwana, compared with their Permian, Triassic, or
Cretaceous counterparts. Knowledge of the floras derives
chiefly from assemblages studied in Argentina and eastern
Australia. Plant groups having their origins in the Triassic
but only reaching quantitative importance in the Jurassic
include caytonialeans, bennettitaleans, pentoxylaleans and
pachypterid seed-ferns. Other elements in the Jurassic floras
include marattiacean, matoniacean, osmundacean,
dicksoniacean and dipteridacean ferns, equisetaleans,
herbaceous lycophytes, cycadaleans and some seed-ferns of
uncertain alliance (e.g. Komlopteris). Although fluctuations
in the abundance of some plant groups are evident through
time, the Jurassic floras show a remarkable conservatism at
genus-level and most genera persist into the earliest
Cretaceous.
Traditionally, Jurassic floras have been interpreted to be
more cosmopolitan than those of preceding and succeeding
periods. Araucarian, podocarp, cheirolepidacean and
taxodiacean conifers, caytonialeans, bennettitaleans and
ginkgoaleans dominated broad tracts of both the Northern
and Southern Hemispheres (Meyen 1987). This
cosmopolitan aspect is generally interpreted as a
consequence of more equable global climates, the assembly
of Pangea and a lack of physical barriers to plant migration
(or range-expansion). Jurassic floras certainly contain
several widespread elements at family and order level but
floristic provincialism is, nevertheless, evident between
Gondwana and the Northern Hemisphere and within
Gondwana during this time (Fig. 3a). Ginkgophytes were a
common element of northern floras but have a disjointed
fossil record in Gondwana. Cheirolepidacean conifers were
more prominent in drier and coastal parts of Pangea. The
dramatic increase in cheirolepidacean pollen in some parts
of Gondwana at the beginning of the Jurassic (Helby et al.
1987) may reflect drier conditions accompanying the demise
of the Dicroidium flora. Pinaceae were present in northern
Pangea but were apparently absent from Gondwana (Meyen
1987). Pentoxylaleans appear to have been a predominantly
Gondwanan group and in many parts of southern Gondwana
they are major components of leaf assemblages from this
time (Halle 1913; Edwards 1934; Gould 1975). More subtle
trends are evident in foliar physiognomy across Gondwana in
the Jurassic and Early Cretaceous. Bennettitalean
assemblages from the palaeoequatorial and European–
Sinian (low-latitude) belts appear to have a significantly
higher species diversity and include many forms with large
( >30 cm long) fronds (Harris 1969). Reticulate-veined
bennettitaleans are also more common at lower latitudes and
appear to be entirely absent from southeastern Gondwana
(at >60° S) in the Jurassic and Early Cretaceous. These
patterns suggest strong palaeoenvironmental controls
(largely associated with latitude) on the distribution of plant
groups regardless of the absence of marine barriers between
Gondwana and Laurasia.
The Cretaceous gymnosperm–angiosperm transition
accompanying Gondwanan breakup
Composition of the Early Cretaceous floras
A large number of modern vascular plant families with
predominantly Southern Hemisphere distributions had their
origins in the Cretaceous. Earliest Cretaceous floras differed
little from those of the Jurassic and were dominated by a
range of conifer and pteridosperm groups. Araucarian and
podocarp conifers were important components of
Gondwanan floras throughout the Cretaceous, although clear
distinctions are evident at specific and generic level between
different regions. Most of these forms are assigned to extinct
genera, although some forms show a suite of characters
suggestive of a close affiliation to Araucaria, Agathis,
Wollemia, Pod oca rp us and Microcachrys (Drinnan and
288 S. McLoughlin
Chambers 1986; Cantrill 1991, 1992; Dettmann 1994;
McLoughlin et al. 1995b, 2000). Cheirolepidacean conifers,
scarce as macrofossils, are represented by locally rich
concentrations of fossil pollen, especially in South America
and in coastal deposits of eastern Gondwana (Dettmann
1994), but this group became progressively less important
through the Cretaceous. Taxodiacean conifers are of
relatively low abundance in Lower Cretaceous assemblages.
Bennettitaleans and pentoxylaleans remained important in
Early Cretaceous (Neocomian–Aptian) floras (Douglas
1969; Baldoni and De Vera 1980; Bose and Banerji 1984;
Anderson and Anderson 1985; Drinnan and Chambers 1985;
Césari et al. 1998, 1999; Cantrill 1997a) but in most parts of
Gondwana these groups had declined to a very minor
component of the vegetation by the Cenomanian. Additional
seed ferns present in the Early Cretaceous, but disappearing
by the mid-Cretaceous, include Komlopteris, Archangelskya,
Ticoa, Pa chypteris and several undescribed taxa. In older
works on fossil floras, the Mesozoic is sometimes described
as the ‘era of cycads’. However, cycads have a very limited
fossil record in the Gondwanan Cretaceous (Douglas 1969;
Archangelsky and Baldoni 1972). Most of the cycad-like
foliar remains from this period represent the leaves of
Bennettitales (sometimes called cycadeoids) which are not
closely related to cycads (Crane 1985) and probably had
entirely different environmental tolerances. The
palynological record of cycads is also equivocal, as fossil
cycad pollen is difficult to distinguish from that of
ginkgophytes, bennettitaleans and pentoxylaleans.
Ginkgoaleans continued to have a disjointed fossil record in
the Early Cretaceous. They are locally abundant in some
Aptian deposits of south-eastern Australia, India and
Argentina, but otherwise they are minor elements of the
Gondwanan Cretaceous floras and probably became extinct
throughout most fragments of the supercontinent before the
end of the period. The fern macrofloras are predominantly
composed of the same groups that had been important
elements of the Jurassic floras: Dipteridaceae,
Gleicheniaceae, Osmundaceae, Schizaeaceae and
Matoniaceae. The fossil spore record indicates a greater
diversity of ferns than that represented in the macrofloras
and includes forms with possible affinities to the
Pteridaceae, Dicksoniaceae, Lophosoriaceae, Blechnaceae
and Marsileaceae (Dettmann 1994). It is likely that many of
the modern fern families had their origins in the Cretaceous,
and these groups underwent diversification in association
with the radiation of angiosperms (Dettmann 1994) at the
expense of some older groups such as the Osmundaceae and
Matoniaceae. Heterosporous aquatic ferns may have
originated in the Late Jurassic (Kovach and Batten 1993), but
they show a dramatic extension in their distribution, and
increase in diversity and abundance in the Aptian and Albian
based on dispersed megaspores (Arcellites and Molaspora).
Lycophytes have a relatively poor macrofossil record in the
Cretaceous. Most forms were probably herbaceous and did
not survive extensive post-mortem transport. However, the
few detailed studies of megaspore suites from the
Gondwanan Early Cretaceous show that most regions
contained diverse populations of isoetaleans and/or
selaginellaleans (Douglas 1973; Banerji et al. 1984; Kovach
and Batten 1989). Equisetaleans declined in importance
through the Early Cretaceous and probably suffered regional
extinctions (e.g. in Australia) by the latter part of the period.
Liverworts are locally well-represented in southern
macrofloras (Drinnan and Chambers 1986; Cantrill 1997b;
Dettmann and Clifford 2000), but to date few studies have
seriously investigated the Gondwana-wide diversity of this
group. Palynological studies suggest that bryophytes were
abundant and diverse components of the Australian
Cretaceous vegetation and the distributions of several
bryophytic spores are used as biostratigraphic markers in
various regions of Gondwana (Morgan et al. 1995;
Archangelsky and Villar de Seoane 1996; Vijaya 1999).
Origin of the angiosperms
By mid-Cretaceous times, angiosperms had diversified to
become important elements of the vegetation. Several
workers have postulated a palaeoequatorial origin for
angiosperms based on the distribution of the earliest known
fossil pollen (Muller 1981; Crane and Lidgard 1989;
Dettmann 1994). Other workers have suggested that high
latitudes may represent important sources of evolutionary
novelties for both animals and plants (Hickey et al. 1983;
Zinmeister and Feldman 1984; Case 1988, 1989; Askin
1989; Hill and Scriven 1995). The latter argument suggests
that the suite of structural characters defining angiosperms
may have included adaptations developed in response to the
harsh environmental conditions and strong selective
pressures of high-latitude environments. However, several
studies have shown successive first appearances of
angiosperm fossils at progressively higher latitudes through
the Cretaceous (Hickey and Doyle 1977; Crane and Lidgard
1989; Drinnan and Crane 1989). During the early radiation
of angiosperms, diversity levels were greatest at lower
latitudes (Crane and Lidgard 1989, 1990) giving support to a
palaeoequatorial origin for the group. The earliest
convincing angiosperm remains are pollen from the
Hauterivian (132–127 million years ago) of Israel and
England (Brenner 1984; Hughes and McDougall 1987).
Sporadic reports of older angiosperm fossils (Cornet 1989a,
1989b; Sun et al. 1993, 1998) have been disputed on the
basis of their taxonomic assignment and/or the age of the
host rocks. Nevertheless, phylogenetic analyses of seed
plants recognise that the sister group to angiosperms
(Gnetales) has a fossil record extending back to at least the
Triassic suggesting that ‘stem taxa’ leading to angiosperms
have an equivalent, if not yet detected, stratigraphic range.
The high diversity of charcoalified flowers ( >100 taxa) in
Gondwana break-up and pre-Cenozoic floristic provincialism 289
Barremian–Aptian deposits of Portugal (Pedersen and Friis
1999), together with the relatively low diversity of
angiosperm pollen and wood in the same deposits also
suggests that herbaceous, insect-pollinated angiosperms had
a considerable evolutionary history by that time. A pre-
Cretaceous origin for angiosperms is, therefore, a strong
possibility, although based on the fossil record any pre-
Cretaceous forms would seem to have had a relatively low
importance in the general vegetation.
Diversification and expansion of early angiosperms
If a palaeoequatorial origin of angiosperms in the mid-
Neocomian is accepted (Drinnan and Crane 1989; Dettmann
1992) then angiosperms required a relatively short period of
time to expand their range into the higher latitudes of
Gondwana. Early pollen and leaf fossils attributable to
angiosperms are reported from the Barremian–Aptian (127–
112 million years ago) of Patagonia (Romero and
Achangelsky 1986) and south-eastern Australia (Dettmann
1986a; Drinnan and Chambers 1986) and from the Albian
(112–99 million years ago) of Alexander Island, Antarctica
(Cantrill and Nichols 1996). The earliest (Neocomian)
angiosperm pollen are monosulcate grains typical of
magnoliids and monocots. The pollen is referable to
Clavatipollenites and is similar to pollen produced by extant
representatives of the Chloranthaceae (Dettmann 1994).
Triaperturate grains diagnostic of the ‘higher’ dicots are first
represented higher in the stratigraphic record at about the
Barremian–Aptian boundary, but in several parts of
Gondwana they first appear and rapidly diversify in the late
Albian associated with a major marine regression (Burger
1980). The rapid expansion of angiosperms corresponded to
an important phase in both the breakup history of Gondwana
and a series of major eustatic sea-level fluctuations.
Although impossible to prove, it is tempting to invoke a
causative link between these geological events and biological
processes. For example, Dettmann (1989) suggested that the
intra-Gondwana rift system and the margins of the newly
developed epicontinental seaways acted as corridors for the
dispersal of early flowering plants. Rift systems typically
incorporate areas of both high and low relief, variable soil
conditions over short distances and a rapidly changing
topography. Similar substrate and climatic variability would
be expected with the rapid marine transgressions and
regressions across the continents in the Early and mid-
Cretaceous. Some surviving angiosperm taxa regarded as
having an early Gondwanan origin (e.g. Nothofagus) are
strongly dependent on disturbance for colonisation of new
territory (Veblen et al. 1977, 1980). Early herbaceous
angiosperms may have been opportunistic species adapted to
colonisation of disturbed sites (Retallack and Dilcher 1981)
and might have taken advantage of the newly established rift
corridors and seaway margins. Alternatively, several studies
have suggested that early angiosperms may have entered
Australia from south-east Asia via a series of detached
Gondwanan or island–arc terranes (Burger 1981,1990,
Truswell et al. 1987). This hypothesis has not been strongly
supported by subsequent fossil or biogeographic studies
(Dettmann and Thomson 1987; Dettmann and Jarzen 1988,
1990; Dettmann 1989; Jarzen and Dettmann 1990; Dettmann
et al. 1992) and most of these microcontinents had probably
assembled as part of Asia before the origin or diversification
of angiosperms (Sengör and Hsü 1984). Palynological data
suggest that many elements of the south-eastern Gondwanan
angiosperm–conifer flora either evolved within the Austro–
Antarctic region or entered Australasia via East and/or West
Antarctic corridors by terrestrial range expansion (Dettmann
1981, 1989; Dettmann and Jarzen 1990).
Early Cretaceous terrestrial environments
In Gondwana, the Early Cretaceous transition from
gymnosperm- to angiosperm-dominated floras corresponded
to a unique combination of environmental conditions.
Through the Early Cretaceous, the middle to high latitude
Gondwanan landmasses experienced strong annual
photoperiod variation, an absence of ice-sheet development,
extensive rifting and continental fragmentation, high levels
of local substrate disturbance, major marine transgressions
and regressions, enhanced CO2 levels and a parent flora and
associated fauna that had changed little at higher taxonomic
levels for the previous 60 million years.
The extreme photoperiod regime experienced at high
latitudes must have placed severe physiological constraints
on the vegetation, particularly with respect to energy loss
through respiration during winter in the absence of
photosynthesis. Modern environments at these latitudes
experience extremely cold climates, but higher CO2 levels in
the Cretaceous (Barron and Washington 1985; Arthur et al.
1988, 1991; Berner 1990) and greater mixing of equatorial
and polar marine waters probably led to globally warmer
conditions (Spicer and Corfield 1992). Evidence for
extensive ice sheets is lacking for the Early Cretaceous,
although ice-rafted dropstones, cryogenic soil patterns,
glendonites and possible varved sediments indicate at least
seasonal freezing at latitudes >70°S in the Aptian (Frakes
and Francis 1988, 1990; Constantine et al. 1998; Tosolini
et al. 1999). Stable isotope data indicate average annual
palaeotemperatures of about –2° to +5°C for the south-
eastern Australian Otway Basin at c. 80°S (Gregory et al.
1989) and +12°C for the central eastern Australian
Eromanga Basin at c. 70°S (Stevens and Clayton 1971) in the
Aptian. Slightly higher Albian–Cenomanian mean annual
palaeotemperatures of about +16°C from the 60–70°S
Eromanga Basin and about +10°C for the c. 70–80°S New
Zealand South Island (Parrish et al. 1998) support a general
warming trend towards the Late Cretaceous corresponding to
increased atmospheric CO2 levels (Berner 1990). Relatively
low palaeotemperatures (about +10°C) estimated for the
290 S. McLoughlin
50°S Carnarvon Basin, Western Australia, at that time may
have been a consequence of local upwelling zones resulting
in anomalously low sea temperatures (Pirrie et al. 1995).
Most Early Cretaceous woods from high latitudes show
strong seasonal growth banding suggesting that many plants
underwent dormancy during the long polar winters
(Jefferson 1983; Francis 1986; Frakes and Francis 1990;
McLoughlin et al. 1995a). There is also some indication that
several plant groups were deciduous. Seed ferns, such as
bennettitaleans and pentoxylaleans, are often represented by
densely matted whole leaves along bedding planes,
suggesting a short period of deciduous leaf fall.
Ginkgophytes, although less common, may also have been
deciduous by analogy with the only extant species of that
group. Conifers such as Elatocladus may have shed entire
short shoots annually, much like the extant Taxodium
distichum which has a similar short-shoot morphology. Even
some ferns, such as Phyllopteroides, may have been
deciduous. Their pinnules are shortly petiolate and whole
pinnules are commonly found in dense accumulations
detached from the parent rachises. Other conifers, including
Brachyphyllum, Pag i o p h y l l u m , Tomaxellia and Athrotaxus
species, are represented by irregularly branched axes with
attached leaves showing rhythmic growth. Such plants may
have been evergreen but probably minimised metabolic
processes during winter in a manner similar to modern
conifers of northern high latitudes. Axelrod (1984)
suggested that light conditions at high latitudes are sufficient
for forest growth, provided that minimum temperatures are
not extreme. Creber and Chaloner (1984) also argued that
ambient temperature rather than light availability is the
limiting factor on the latitudinal distribution of modern
vegetation and that the total amount of available light energy
per annum beyond the Antarctic Circle is not significantly
less than that at temperate latitudes. Read and Francis (1992)
showed experimentally that a range of modern Southern
Hemisphere gymnosperms and angiosperms can survive
extended periods of darkness and that most taxa cope best
when the period of darkness accompanies moderately cold
(about 4°C) rather than warm conditions. Hill et al. (1999),
therefore, suggested that many high latitude Cretaceous
plants may have preferred cooler interior continental settings
rather than warmer coastal environments, where winter
respirational loss would have been greater and summer
cloudiness might have reduced photosynthesis potential.
Evidence from in situ fossil tree stumps and theoretical
models suggest that Cretaceous high-latitude forests
consisted of widely spaced trees. These trees may have
adopted conical canopies and even vertically orientated
leaves or leafy shoots to maximise low-angle light
interception (Creber and Chaloner 1985; Hill 1994). The
consequent open forest structure may have permitted
anemophily to be a viable method of pollination for many
understory shrubs and herbs (Hill 1994).
Floristic provincialism in Early Cretaceous Gondwanan
floras
Some earlier studies suggested a relatively uniform flora
existed across Gondwana in the Early Cretaceous. Meyen
(1987) recognised a single Gondwanan floristic province and
Vakhrameev (1991) identified two provinces in the
Cretaceous corresponding to low–middle and high latitudes.
Recent description of additional fossil assemblages and
revision of older systematic works necessitate a restructuring
of these simple models of provincialism. Although several
genera are distributed widely across the supercontinent,
eastern and western Gondwanan floras can be broadly
distinguished by the distribution of a range of unique taxa and
in some cases by specialised foliar physiognomies. The lack
of exposed Early Cretaceous sediments in East Antarctica
means that our knowledge of its vegetation from that time is
restricted to reworked palynomorphs in younger sediments
preserved around the periphery of the continent.
Nevertheless, East Antarctica held a central position in
Gondwana and was probably critical to understanding the
relationships of floras from different parts of the
supercontinent. Despite the lack of data concerning the East
Antarctic biota, about four subprovinces (essentially regions
of endemism) are recognisable within southern Gondwana
(Fig. 3b): Patagonia–Palmer (southern South American–
Antarctic Peninsula), South Africa, Indo-Eromanga (India–
northern Australia) and Tasman (southeastern Australia and
New Zealand). Differentiation of these floras was associated
with a suite of gradational environmental factors; the
principal ones being latitudinal or climatic differences,
tectonic segregation of formerly connected landmasses and
extensive marine incursions in the Barremian–Albian
(particularly in Australasia, South America and the Antarctic
Peninsula) causing further fragmentation of terrestrial
environments and promoting allopatric speciation. Fossil
assemblage composition can additionally be influenced by
local habitat differences associated with topography,
depositional settings, substrate types, moisture levels,
interspecific competition and fire (Dettmann 1986a, 1986b,
1994; Askin 1989; Dettmann et al. 1991; Douglas 1994). In
the Lower Cretaceous sediments of Australia, fern spores are
reportedly more diverse and abundant in coastal settings,
whereas homosporous lycopods are more common away from
marine influence (Dettmann 1994). Sphagnalean moss and
hepatic spores are more numerous in low-relief epicontinental
basins (Dettmann 1994). Unless these local habitat variations
can be accounted for and until broader palaeobotanical
sampling is available across Gondwana, studies using
multivariate analyses of the distribution of fossil plant genera
may lead to spurious biogeographic interpretations.
Nevertheless, there are some broad, Early Cretaceous,
floristic patterns evident across Gondwana at generic and
higher levels and some of these are outlined below.
Gondwana break-up and pre-Cenozoic floristic provincialism 291
A range of very small-leafed bennettitaleans (with
<5 mm2 pinnules) and an abundance of pentoxylaleans,
Phyllopteroides ferns and bryophytes characterise south-
eastern Australian Neocomian floras. Ginkgophytes are
common in the Aptian, but cheirolepidacean pollen is
generally in low abundance (Dettmann 1994). The presence
of seed-ferns such as Komlopteris sp., Pachypteris
austropapillosa and ‘Rienitsia’ variablis and an abundance
of heterosporous lycophytes (as evidenced by diverse
megaspore assemblages) is also characteristic of this
province in the Early Cretaceous (Douglas 1969; Drinnan
and Chambers 1986). New Zealand floras are also rich in
Phyllopteroides and small pentoxylalean leaves (Arber 1917;
Edwards 1934; Parris et al. 1995). Together with
bennettitaleans, these groups appear to persist slightly later
in the stratigraphic record than in Australia. Palissya, an
enigmatic gymnosperm fruit, is well represented in south-
eastern Australia and New Zealand but these fruits have not
been recorded elsewhere in Gondwana (Parris et al. 1995).
Indo-Eromangan floras are rich in bennettitaleans having
larger leaves (with commonly >100 mm2 pinnules) than
forms from the Tasman region. These floras are also rich in
gleicheniacean and dipteridacean ferns (Walkom 1928;
McLoughlin 1996). Araucarian conifers show increased
importance compared with podocarps in northern areas of
Gondwana (India, Argentina, northern Australia: Dettmann
1994) at this time. The conifer pollen Hoegisporis is
restricted to northern and western areas of Australia and
cheirolepidacean pollen is more common in the Indo-
Eromanga region than in the Tasman subprovince (Dettmann
1994). Many palynomorph taxa occurring in both
subprovinces appear slightly earlier in the stratigraphic
record in the former region (Helby et al. 1987) suggesting
poleward advance with climatic amelioration in the mid-
Cretaceous. The Indo-Eromanga palynofloras generally have
a higher diversity than those of the Tasman region (Dettmann
1973; Burger 1990). Both subprovinces share
Phyllopteroides ferns although this group reaches its greatest
abundance and diversity in south-eastern Australia. Indian
floras incorporate reticulate-veined bennettitalean species.
These taxa have not yet been recorded from the Tasman
subprovince.
Southern African Early Cretaceous floras are
characterised by bennettitaleans with large leaves and, in
several cases, reticulate venation. Pentoxylalean leaves are
also larger than the eastern Gondwanan forms. Although
several fern genera are shared with other Gondwanan
regions, the South African forms are distinct at species level
(Anderson and Anderson 1985).
South American and Antarctic Peninsula floras
(Patagonia–Palmer subprovince) are also characterised by
bennettitaleans with large leaves. The South American
assemblages in particular have a greater diversity than most
other Gondwanan floras and incorporate several endemic
genera (e.g. Ticoa, Ruflorinia, Mesosingeria and Ktalenia:
Archangelsky 1963). Several species are shared between
southern South America and the Antarctic Peninsula,
including large (presently undescribed) permineralised
cycadophyte trunks. These similarities suggest that the
regions were closely affiliated in the Early Cretaceous
although the presence of sparse Phyllopteroides (a genus
otherwise restricted to eastern Gondwana) in the Antarctic
Peninsula suggests that this region was at somewhat higher
latitudes than southern South America and retained some
floristic links with the Tasman subprovince.
Beyond the mid-Cretaceous floristic transition
Late Cretaceous and Cenozoic fossil assemblages generally
reflect increasing divergence in the floras of the segregated
parts of Gondwana. Some angiosperm elements (e.g.
Nothofagaceae, Winteraceae and Proteaceae) had clearly
evolved before the separation of at least some Gondwanan
landmasses, and to a significant extent the distribution of
extant representatives of these groups probably reflects
vicariance events. However, in some cases the available
geological and palaeontological evidence suggests that
continental breakup occurred before the differentiation of
plant groups that are traditionally cited as having a
‘Gondwanan’ heritage based on their wide Southern
Hemisphere distribution. The presence of Proteaceae in
southern Africa, including some elements with close
relationships (taxa associated by distal nodes on molecular
phylogenies) to south-western Australian clades (Hoot and
Douglas 1998), suggests a close biogeographic relationship
between these areas. The earliest putative Proteaceae pollen
occur in Cenomanian–Turonian (99–89 million years ago)
strata of northern Gondwana (Muller 1981), but apparent
diversification of the Proteaceae (based on pollen records)
was centred in southern high latitudes in the latest
Cretaceous (Dettmann and Jarzen 1991). However, evidence
from seafloor spreading suggests that Africa separated from
other Gondwanan continents early in the breakup history of
the supercontinent (in Late Jurassic–Early Cretaceous:
Fig. 4). For vicariance to be a plausible mechanism to
explain these distributions, a large number of extinctions
would need to be invoked and/or extension of the
stratigraphic record of closely related (derived) taxa, now
separated by ocean barriers, back to the Early Cretaceous.
This would further necessitate the origin of more ‘basal’ taxa
to times even earlier in the Cretaceous where the fossil
record for angiosperms as a whole is relatively tenuous.
Long-distance dispersal may, therefore, have been an
important process accounting for the extant distribution of
such taxa. Its role may have been more important during the
Cretaceous and Paleogene when oceanic barriers in the
Southern Hemisphere were less extensive than at present.
The processes of regional extinction and recent range
alteration through climate change or terrane modification
292 S. McLoughlin
may also detract from simple vicariance models to explain
the distribution of some taxa. The relationship of the extant
representatives of the Nothofagus subgenus Brassospora in
New Caledonia and New Guinea might initially suggest a
close geological relationship between these landmasses.
However, the fossil record indicates an extensive distribution
of the subgenus through much of southern Gondwana in the
Paleogene (Dettmann et al. 1990). The modern
representatives in New Caledonia and New Guinea may have
only recently colonised those regions from separate source
areas and parent stocks following Neogene climatic and
terrain changes (Swenson et al. 2001). Similarly, on the basis
of the extant distribution of taxa, the Casuarinaceae could be
considered to have an Indo-Pacific palaeoequatorial origin
(Fig. 8). However, an extensive and long fossil pollen record
in Gondwana (MacPhail et al. 1994) and limited macrofossil
evidence (Frenguelli 1943; Campbell and Holden 1984)
indicate a much wider, but predominantly southern
Gondwanan, distribution of this group during the Paleogene.
This early distribution included regions as far away from the
modern area of endemism as New Zealand, northern
Patagonia, southern Africa and the Oligocene oceanic
islands of the Ninetyeast Ridge (Fig. 8). Again, a vicariance
scenario would necessitate derivation of this group prior to
the mid-Albian, yet a pre-Cenozoic fossil record for
Casuarinaceae is lacking. Perhaps the Late Cretaceous
fossils of this group have not yet been found. However, given
the presence of a range of other angiosperm pollen in Upper
Cretaceous sediments, the preponderance of anemophily
within the Casuarinaceae and the common occurrence of
casuarinacean pollen in post-Paleocene sediments, its
absence in the Cretaceous seems unlikely to be attributable
to taphonomic or ecological biases. Whether vicariance or
long-distance dispersal was the primary factor involved in
the distribution of the Casuarinaceae on Gondwanan
continental fragments, its modern presence on the oceanic
islands of the central and western Pacific (Fig. 8) must have
involved dispersal over extensive marine hurdles as these
geologically young islands have never had land connections
to the continents. It is perhaps the past and prevailing
distribution of appropriate mesothermal–megathermal
climates that have dictated the extinction of this group in
southern Gondwana, its survival in Australia (accompanying
continental movement into lower latitudes) and its probable
recent expansion into the South-east Asia–Pacific region.
India provides another Gondwanan case where a purely
vicariant scenario would appear not to provide a complete
solution to Cenozoic plant distributions. India became
isolated early in the breakup history of Gondwana and its
rapid northward movement through warmer climatic belts
was accompanied by radical floristic changes. By the Early
Paleocene India had reached equatorial latitudes but
according to most reconstructions it remained isolated from
Asia (Barron 1987; Lee and Lawver 1995; Scotese 1997). At
this time, some similarities remained between the flora of
India and its Gondwanan sister landmasses (Bande 1992),
but it also hosted an extensive range of plant groups
(including palms, bamboo, Bignoniaceae, Musaceae and
Abietinae) suggestive of Laurasian or palaeotropical links
across a significant marine barrier (Bande and Prakash 1986;
Extant
Pre-Quaternary fossils
Fig. 8. Distribution of principal pre-Quaternary fossil records (grey circles; located on mostly southern
high latitude ‘Gondwanan’ landmasses that were isolated between 105 and 28 million years ago) and extant
representatives (shaded black; on mostly Australian–Malesian–Melanesian landmasses) of Casuarinaceae
(adapted from Johnson and Wilson 1989). The northern and eastern limit of extant representatives is
chiefly defined by the distribution of the littoral taxon Casuarina equisetifolia Forst. and For st .f.
Gondwana break-up and pre-Cenozoic floristic provincialism 293
Bande 1992). Substantially greater affinities with Asian
floras, evidenced by an influx of dipterocarp and legume
taxa, were established from the beginning of the Neogene
following India–Asia collision (Guleria 1992).
Conclusions
‘Gondwana’ refers to the unified, mostly Southern
Hemisphere, continental entity that amalgamated at the
beginning of the Palaeozoic and progressively fragmented
from the Devonian to present (with a major phase of
continental breakup in the Cretaceous). The question of
whether to retain the term Gondwanan for terranes and their
biota after breakup is somewhat academic as they essentially
chart different tectonic and palaeoclimatic courses after
separation. Similarly, if biotic dispersal has occured between
two post-breakup Gondwanan terranes (e.g. Madagascar to
India) then it may be more appropriate to describe the
dispersed taxon in terms of its source area (e.g. Madagascan)
rather than applying the broader term ‘Gondwanan’ or
‘dispersed-Gondwanan’.
High southern latitude floras have retained a distinct
floristic composition through most of the late Palaeozoic and
Mesozoic, even when Gondwana was united to Laurasia as
part of Pangea and through global extinction events when the
dominant plant groups were replaced. The phytogeographic
zonation of Gondwana through this time is strongly
correlative to palaeolatitudinal belts and/or palaeoclimatic
zones that can be independently documented through
palaeomagnetic and sedimentary facies analysis. The
distribution of many modern plants may be influenced as
much by current and past climatic regimes as their historical
geographic derivation. The robustness of a geographic
barrier to dispersal or range-expansion decreases with the
length of time it is operative. Extensive marine domains have
not proved to be absolute barriers to the colonisation of
geologically young oceanic islands. It is, therefore, likely
that floristic interchange between isolated landmasses was
even more common during the Cretaceous and Paleogene
when more equable global climates prevailed and
intercontinental distances in the Southern Hemisphere were
less than at present.
A detailed biogeographic understanding of the Southern
Hemisphere floras must ultimately depend on a thorough
understanding of the timing of geological and biological
events that led to modern terrane and floristic distributions.
For most parts of Gondwana, the amalgamation and breakup
histories and palaeoclimatic regimes are now well-
documented via palaeomagnetic studies, radiometric dating,
tectonostratigraphic analyses, palaeontological investiga-
tions and comparison of sedimentary facies. On the
biological side of the problem, detailed morphological and
molecular analyses together with cladistic methodologies are
now producing robust phylogenies for some plant groups.
Furthermore, biogeographic analyses of widespread groups
with well-constrained phylogenies can, with the elimination
of geographic paralogy, provide a useful measure of the
historical similarities or ‘relatedness’ of the biotas between
areas. How one interprets the historical processes that led to
those patterns of relatedness remains a difficulty and, as
noted by Pole (1994), the same data can be interpreted by
different biogeographers to support radically different
hypotheses of historical processes.
Progress in this field will require better techniques and
methodologies for obtaining age constraints on biological
differentiation events (cladogenesis). It should then be
possible to compare the age of biological differentiation with
the ages of terrane separation in order to test biogeographic
hypotheses or at least to derive degrees of probability to
assess various hypotheses. Palaeontological data and
mutation rates derived from DNA sequencing currently
provide the best information on the age of plant clades. Both
techniques have their limitations and frequently yield
conflicting results, yet the former can at least provide
minimum ages for cladogenesis. Increased focus on this
weakest link in biogeographic analysis is likely to yield the
most informative results in the future.
Acknowledgments
This research is supported by an Australian Research
Fellowship and Large Grant funding from the Australian
Research Council. A.-M. Tosolini is thanked for providing
constructive criticism on geological aspects of the
manuscript.
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