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Gondwana
Gondwana
( /ɡɒndˈwɑːnə/),[1] or Gondwanaland,[2] was a supercontinent that existed from the Neoproterozoic (about 550 million years ago) until the Carboniferous
Carboniferous
(about 320 million years ago). It was formed by the accretion of several cratons. Eventually, Gondwana became the largest piece of continental crust of the Paleozoic
Paleozoic
Era, covering an area of about 100,000,000 km2 (39,000,000 sq mi).[3] During the Carboniferous, it merged with Euramerica
Euramerica
to form a larger supercontinent called Pangaea. Gondwana
Gondwana
(and Pangaea) gradually broke up during the Mesozoic
Mesozoic
Era. The remnants of Gondwana
Gondwana
make up about two-thirds of today's continental area. The formation of Gondwana
Gondwana
began c. 800 to 650 Ma with the East African Orogeny – the collision of India, Madagascar, and Sri Lanka with East Africa – and was completed c. 600 to 530 Ma with the overlapping Brasiliano and Kuunga orogenies – the collision of South America
South America
with Africa
Africa
and the addition of Australia and Antarctica, respectively.[4]

Contents

1 Origin of concept 2 Formation 3 Gondwana
Gondwana
as part of Pangaea 4 Peri-Gondwana 5 Break-up

5.1 Mesozoic

5.1.1 Opening of western Indian Ocean 5.1.2 Opening of eastern Indian Ocean 5.1.3 Opening of South Atlantic Ocean 5.1.4 Early Andean orogeny

5.2 Cenozoic

5.2.1 Australia– Antarctica
Antarctica
separation

6 Biogeography

6.1 Post-Cambrian diversification 6.2 Modern diversification

7 See also 8 References

8.1 Notes 8.2 Sources

9 External links

Origin of concept[edit] The continent of Gondwana
Gondwana
was named by Austrian scientist Eduard Suess, after the Gondwana
Gondwana
region of central northern India
India
which is derived from Sanskrit
Sanskrit
for "forest of the Gonds". The name had been previously used in a geological context, first by H.B. Medlicott in 1872,[5] from which the Gondwana
Gondwana
sedimentary sequences (Permian-Triassic) are also described. The term "Gondwanaland" is preferred by some scientists in order to make a clear distinction between the region and the supercontinent.[6] Formation[edit]

Eastern Gondwana. 620 to 550 Ma post-collisional extension of the East African Orogeny
East African Orogeny
in blue and 570 to 530 Ma collisional metamorphism of the Kuunga orogeny
Kuunga orogeny
in red.[7]

The assembly of Gondwana
Gondwana
was a protracted process that occurred during the Neoproterozoic and Paleozoic
Paleozoic
which remains relatively poorly constrained because of the lack of paleo-magnetic data. Several orogenies, collectively known as the Pan-African orogeny, led to the amalgamation of most of the continental fragments of a much older supercontinent, Rodinia. One of those orogenic belts, the Mozambique Belt, formed 800 to 650 Ma and was originally interpreted as the suture between East (India, Madagascar, Antarctica, and Australia) and West Gondwana
Gondwana
( Africa
Africa
and South America). Three orogenies were recognized during the 1990s: the East African Orogeny (650 to 800 Ma) and Kuunga orogeny
Kuunga orogeny
(including the Malagasy Orogeny
Orogeny
in southern Madagascar) (550 Ma) – the collision between East Gondwana
Gondwana
and East Africa
Africa
in two steps, and the Brasiliano orogeny (660 to 530 Ma) – the collision between South American and African cratons.[8] The final stages of Gondwanan assembly overlapped with the opening of the Iapetus Ocean
Iapetus Ocean
between Laurentia
Laurentia
and western Gondwana.[9] During this interval, the Cambrian explosion
Cambrian explosion
occurred. Laurentia
Laurentia
was docked against the western shores of a united Gondwana
Gondwana
for a short period near the Precambrian/Cambrian boundary, forming the short-lived and still disputed supercontinent Pannotia.[10] The Mozambique Ocean
Mozambique Ocean
separated the Congo–Tanzania–Bangweulu Block of central Africa
Africa
from Neoproterozoic India
India
(India, the Antongil
Antongil
Block in far eastern Madagascar, the Seychelles, and the Napier and Rayner Complexes in East Antarctica). The Azania continent[11] (much of central Madagascar, the Horn of Africa
Horn of Africa
and parts of Yemen
Yemen
and Arabia) was an island in the Mozambique Ocean. The Australia/Mawson continent was still separated from India, eastern Africa, and Kalahari by c. 600 Ma when most of western Gondwana
Gondwana
had already been amalgamated. By c. 550 Ma, India
India
had reached its Gondwana
Gondwana
position which initiated the Kuunga orogeny
Kuunga orogeny
(also known as the Pinjarra orogeny). Meanwhile, on the other side of the forming Africa, Kalahari collided with Congo and Rio de la Plata which closed the Adamastor Ocean. c. 540–530 Ma the closure of the Mozambique Ocean
Mozambique Ocean
brought India
India
next to Australia– East Antarctica
East Antarctica
and both North and South China were located in proximity to Australia.[12] Other blocks which helped to form parts of the Southern Cone
Southern Cone
of South America, including a piece transferred from Laurentia
Laurentia
when the west edge of Gondwana
Gondwana
scraped against southeast Laurentia
Laurentia
in the Ordovician.[13] This is the Cuyania
Cuyania
or Precordillera terrane of the Famatinian orogeny
Famatinian orogeny
in northwest Argentina which may have continued the line of the Appalachians
Appalachians
southwards.[14]

Reconstruction showing final stages of assembly of Gondwana, 550 Mya

As the rest of Gondwana
Gondwana
formed, a complex series of orogenic events assembled the eastern parts of Gondwana
Gondwana
(eastern Africa, Arabian-Nubian Shield, Seychelles, Madagascar, India, Sri Lanka, East Antarctica, and Australia) c. 750 to 530 Ma. First the Arabian-Nubian Shield collided with eastern Africa
Africa
(in the Kenya-Tanzania region) in the East African Orogeny c.750 to 620 Ma. Then Australia and East Antarctica
East Antarctica
were merged with the remaining Gondwana
Gondwana
c. 570 to 530 Ma in the Kuunga Orogeny.[15] The later Malagasy orogeny at about 550–515 Mya affected Madagascar, eastern East Africa
Africa
and southern India. In it, Neoproterozoic India
India
collided with the already combined Azania and Congo–Tanzania–Bangweulu Block, suturing along the Mozambique Belt.[16] The 18,000 km (11,000 mi)-long Terra Australis
Terra Australis
Orogen developed along Gondwana's western, southern, and eastern margins.[17] Proto-Gondwanan Cambrian arc belts from this margin have been found in eastern Australia, Tasmania, New Zealand, and Antarctica. Though these belts formed a continuous arc chain, the direction of subduction was different between the Australian-Tasmanian and New Zealand-Antarctica arc segments.[18] Gondwana
Gondwana
as part of Pangaea[edit]

Gondwana
Gondwana
formed part of Pangaea
Pangaea
for c. 150 Ma[19]

Main article: Pangaea Gondwana
Gondwana
and Laurussia
Laurussia
formed the Pangaea
Pangaea
supercontinent during the Carboniferous. Pangaea
Pangaea
began to break up in the Mid- Jurassic
Jurassic
when the Central Atlantic opened.[20] In the western end of Pangaea, the collision between Gondwana
Gondwana
and Laurussia
Laurussia
closed the Rheic and Palaeo-Tethys oceans. The obliquity of this closure resulted in the docking of some northern terranes in the Marathon, Ouachita, Alleghanian, and Variscan orogenies, respectively. Southern terranes, such as Chortis and Oaxaca, on the other hand, remained largely unaffected by the collision along the southern shores of Laurentia. Some Peri-Gondwanan terranes, such as Yucatán and Florida, were buffered from collisions by major promontories. Other terranes, such as Carolina and Meguma, were directly involved in the collision. The final collision resulted in the Variscan-Appalachian Mountains, stretching from present-day Mexico to southern Europe. Meanwhile, Baltica
Baltica
collided with Siberia and Kazakhstania which resulted in the Uralian orogeny
Uralian orogeny
and Laurasia. Pangaea
Pangaea
was finally amalgamated in the Late Carboniferous-Early Permian
Permian
but the oblique forces continued until Pangaea
Pangaea
began to rift in the Triassic.[21] In the eastern end collisions occurred slightly later. The North China, South China, and Indochina blocks rifted from Gondwana
Gondwana
during the middle Paleozoic
Paleozoic
and opened the Proto-Tethys Ocean. North China docked with Mongolia and Siberia during the Carboniferous–Permian followed by South China. The Cimmerian blocks then rifted from Gondwana
Gondwana
to form the Palaeo-Thethys and Neo-Tethys oceans in the Late Carboniferous
Carboniferous
and docked with Asia
Asia
during the Triassic
Triassic
and Jurassic. Western Pangaea
Pangaea
began to rift while the eastern end was still being assembled.[22] The formation of Pangaea
Pangaea
and its mountains had a tremendous impact on global climate and sea levels, which resulted in glaciations and continent-wide sedimentation. In North America, the base of the Absaroka sequence coincides with the Alleghanian and Ouachita orogenies and are indicative of a large-scale change in the mode of deposition far away from the Pangaean orogenies. Ultimately, these changes contributed to the Permian– Triassic
Triassic
extinction event and left large deposits of hydrocarbons, coal, evaporite, and metals.[23] The break-up of Pangaea
Pangaea
began with the Central Atlantic magmatic province (CAMP) between South America, Africa, North America, and Europe. CAMP covered more than seven million square kilometres over a few million years, reached its peak at c. 200 Ma, and coincided with the Triassic– Jurassic
Jurassic
extinction event.[24] The reformed Gondwanan continent was not precisely the same as that which had existed before Pangaea
Pangaea
formed; for example, most of Florida
Florida
and southern Georgia and Alabama
Alabama
is underlain by rocks that were originally part of Gondwana, but this region stayed attached to North America
North America
when the Central Atlantic opened.[25] Peri-Gondwana[edit] A large number of terranes were accreted to Eurasia
Eurasia
during Gondwana's existence but the Cambrian or Precambrian origin of many of these terranes remains uncertain. For example, some Palaeozoic terranes and microcontinents that now make up Central Asia, often called the "Kazakh" and "Mongolian terranes", were progressively amalgamated into the continent Kazakhstania in the Late Silurian. Whether these blocks originated on the shores of Gondwana
Gondwana
is not known.[26] In the Early Palaeozoic the Armorican terrane, which today form large parts of France, was part of either Peri- Gondwana
Gondwana
or core Gondwana; the Rheic Ocean
Rheic Ocean
closed in front of it and the Palaeo-Tethys Ocean opened behind it. Precambrian rocks from the Iberian Peninsula
Iberian Peninsula
suggest it too probably formed part of core Gondwana
Gondwana
before its detachment as an orocline in the Variscan orogeny
Variscan orogeny
close to the Carboniferous– Permian
Permian
boundary.[27] South-east Asia
Asia
is made of Gondwanan and Cathaysian continental fragments that were assembled during the Mid-Palaeozoic and Cenozoic. This process can be divided into three phases of rifting along Gondwana's northern margin: firstly, in the Devonian, North and South China together with Tarim and Quidam (north-western China) rifted opening the Palaeo-Tethys behind them. These terranes accreted to Asia during Late Devonian
Devonian
and Permian. Secondly, in the Late Carboniferous to Early Permian, Cimmerian terranes opened Meso-Tethys Ocean; Sibumasu and Qiantang were added to south-east Asia
Asia
during Late Permian
Permian
and Early Jurassic. Thirdly, in the Late Triassic
Triassic
to Late Jurassic, Lhasa, West Burma, Woyla terranes opened the Neo-Tethys Ocean; Lhasa collided with Asia
Asia
during the Early Cretaceous and West Burma and Woyla during the Late Cretaceous.[28] During the Neoproterozoic to Palaeozoic phase of the Terra Australis orogen a series of terranes were rafted from the Andean margin when the Iapteus Ocean opened, to be added back to Gondwana
Gondwana
during the closure of that ocean.[29] Gondwana's long, northern margin had remained a mostly passive margin throughout the Palaeozoic. The Early Permian
Permian
opening of the Neo-Tethys Ocean along this margin produced a long series of terranes many of which were and still are being deformed in the Himalaya Orogeny. From Turkey to north-eastern India: the Taurides in southern Turkey; the Lesser Caucasus Terrane in Georgia; the Sanand, Alborz, and Lut terranes in Iran; the Mangysglak or Kopetdag Terrane in the Caspian Sea; the Afghan Terrane; the Karakorum Terrane in northern Pakistan; and the Lhasa and Qiangtang terranes in Tibet. The Permian–Triassic widening of the Neo-Tethys pushed all these terranes across the Equator and over to Eurasia.[30] Break-up[edit] Mesozoic[edit] Antarctica, the centre of the supercontinent, shared boundaries with all other Gondwana
Gondwana
continents and the fragmentation of Gondwana propagated clockwise around it. The break-up was the result of one of the Earth's most extensive large igneous provinces c. 200 to 170 Ma, but the oldest magnetic anomalies between South America, Africa, and Antarctica
Antarctica
are found in what is now the southern Weddell Sea
Weddell Sea
where initial break-up occurred during the Jurassic
Jurassic
c. 160 to 180 Ma.[31] Opening of western Indian Ocean[edit]

The first ocean floor formed between Madagascar
Madagascar
and Africa
Africa
c. 150 Ma (left) and between India
India
and Madagscar c. 70 Ma (right).

Gondwana
Gondwana
began to break up in the early Jurassic
Jurassic
following the extensive and fast emplacement of the Karoo-Ferrar flood basalts c. 184 Ma. Before the Karoo plume initiated rifting between Africa
Africa
and Antarctica, it separated a series of smaller continental blocks from Gondwana's southern, Proto-Pacific margin (along what is now the Transantarctic Mountains): the Antarctic Peninsula, Marie Byrd Land, Zealandia, and Thurston Island; the Falkland Islands
Falkland Islands
and Ellsworth–Whitmore Mountains (in Antarctica) were rotated 90° in opposite directions; and South America
South America
south of the Gastre Fault (often referred to as Patagonia) was pushed westward.[32] The history of the Africa- Antarctica
Antarctica
break-up can be studied in great detail in the fracture zones and magnetic anomalies flanking the Southwest Indian Ridge.[33] The Madagascar
Madagascar
block and the Mascarene Plateau, stretching from the Seychelles
Seychelles
to Réunion, were broken off India; elements of this breakup nearly coincide with the Cretaceous–Paleogene extinction event. The India–Madagascar– Seychelles
Seychelles
separations appear to coincide with the eruption of the Deccan basalts, whose eruption site may survive as the Réunion
Réunion
hotspot. The Seychelles
Seychelles
and the Maldives are now separated by the Central Indian Ridge. During the initial break-up in the Early Jurassic
Jurassic
a marine transgression swept over the Horn of Africa
Horn of Africa
covering Triassic planation surfaces with sandstone, limestone, shale, marls and evaporites.[34][35]

Opening of eastern Indian Ocean[edit]

The first ocean floor formed between India
India
and Antarctica
Antarctica
c. 120 Ma (left). The Kerguelen LIP began to form the Ninety East ridge c. 80 Ma (centre). The Indian and Australian plates merged c. 40 Ma (right).

East Gondwana, comprising Antarctica, Madagascar, India, and Australia, began to separate from Africa. East Gondwana
Gondwana
then began to break up c. 132.5 to 96 Ma when India
India
moved northwest from Australia-Antarctica.[36] The Indian Plate
Indian Plate
and the Australian Plate are now separated by the Capricorn Plate
Capricorn Plate
and its diffuse boundaries.[37] During the opening of the Indian Ocean, the Kerguelen hotspot first formed the Kerguelen Plateau
Kerguelen Plateau
on the Antarctic Plate
Antarctic Plate
c. 118 to 95 Ma and then the Ninety East Ridge
Ninety East Ridge
on the Indian Plate at c. 100 Ma.[38] The Kerguelen Plateau
Kerguelen Plateau
and the Broken Ridge, the southern end of the Ninety East Ridge, are now separated by the Southeast Indian Ridge. Separation between Australia and East Antarctica
East Antarctica
began c. 132 Ma with sea-floor spreading occurring c. 96 Ma. A shallow seaway developed over the South Tasman Rise during the Early Cenozoic
Cenozoic
and as oceanic crust started to separate the continents during the Eocene
Eocene
c. 35.5 Ma global ocean temperature dropped significantly.[39] A dramatic shift from arc- to rift magmatism c. 100 Ma separated Zealandia, including New Zealand, the Campbell Plateau, Chatham Rise, Lord Howe Rise, Norfolk Ridge, and New Caledonia, from West Antarctica
West Antarctica
c. 84 Ma.[40]

Opening of South Atlantic Ocean[edit]

At c. 126 Ma (left) the Falkland Plateau began to slide past southern Africa
Africa
and the Paraná-Etendeka LIP had opened the Mid-Atlantic Ridge. At c. 83 Ma (right) the South Atlantic was fully opened and the Romanche Fracture Zone was forming near the Equator.

The opening of the South Atlantic Ocean
South Atlantic Ocean
divided West Gondwana
Gondwana
(South America and Africa), but there is a considerable debate over the exact timing of this break-up. Rifting propagated from south to north along Triassic–Early Jurassic
Jurassic
lineaments, but intra-continental rifts also began to develop within both continents in Jurassic–Cretaceous sedimentary basins; subdividing each continent into three sub-plates. Rifting began c. 190 Ma at Falkland latitudes, forcing Patagonia to move relative to the still static remainder of South America
South America
and Africa, and this westward movement lasted until the Early Cretaceous 126.7 Ma. From there rifting propagated northward during the Late Jurassic
Jurassic
c. 150 Ma or Early Cretaceous c. 140 Ma most likely forcing dextral movements between sub-plates on either side. South of the Walvis Ridge
Walvis Ridge
and Rio Grande Rise
Rio Grande Rise
the Paraná and Etendeka magmatics resulted in further ocean-floor spreading c. 130 to 135 Ma and the development of rifts systems on both continents, including the Central African Rift
Rift
System and the Central African Shear Zone
Central African Shear Zone
which lasted until c. 85 Ma. At Brazilian latitudes spreading is more difficult to assess because of the lack of palaeo-magnetic data, but rifting occurred in Nigeria at the Benue Trough
Benue Trough
c. 118 Ma. North of the Equator the rifting began after 120.4 Ma and continued until c. 100 to 96 Ma.[41]

Early Andean orogeny[edit] The first phases of Andean orogeny
Andean orogeny
in the Jurassic
Jurassic
and Early Cretaceous were characterized by extensional tectonics, rifting, the development of back-arc basins and the emplacement of large batholiths.[42][43] This development is presumed to have been linked to the subduction of cold oceanic lithosphere.[43] During the mid to Late Cretaceous (ca. 90 million years ago) the Andean orogeny
Andean orogeny
changed significantly in character.[42][43] Warmer and younger oceanic lithosphere is believed to have started to be subducted beneath South America around this time. Such kind of subduction is held responsible not only for the intense contractional deformation that different lithologies were subject to, but also the uplift and erosion known to have occurred from the Late Cretaceous onward.[43] Plate tectonic reorganization since the mid-Cretaceous might also have been linked to the opening of the South Atlantic Ocean.[42] Another change related to mid-Cretaceous plate tectonic changes was the change of subduction direction of the oceanic lithosphere that went from having south-east motion to having a north-east motion at about 90 million years ago.[44] While subduction direction changed it remained oblique (and not perpendicular) to the coast of South America, and the direction change affected several subduction zone-parallel faults including Atacama, Domeyko and Liquiñe-Ofqui.[43][44] Cenozoic[edit] The Indian subcontinent
Indian subcontinent
began to collide with Asia
Asia
c. 70 Ma since when more than 1,400 km (870 mi) of crust has been absorbed by the Himalayan-Tibetan orogen. During the Cenozoic
Cenozoic
the orogen resulted in the construction of the Tibetan Plateau
Tibetan Plateau
between the Tethyan Himalayas
Himalayas
in the south and the Kunlun and Qilian mountains in the north.[45] Later, South America
South America
was connected to North America
North America
via the Isthmus of Panama, cutting off a circulation of warm water and thereby making the Arctic
Arctic
colder,[46] as well as allowing the Great American Interchange. The breakup of Gondwana
Gondwana
can be said to continue in eastern Africa
Africa
at the Afar Triple Junction, which separates the Arabian, Nubian, and Somali plates, resulting in rifting in the Red Sea
Red Sea
and East African Rift.[47] Australia– Antarctica
Antarctica
separation[edit] In the Early Cenozoic
Cenozoic
Australia was still connected to Antarctica
Antarctica
c. 35–40° south of its current location and both continents were largely unglaciated. A rift between the two developed but remained an embayment until the Eocene-Oligocene boundary when the Circumpolar Current developed and the glaciation of Antarctica
Antarctica
began.[48] Australia was warm and wet during the Palaeocene and dominated by rainforest. The opening of the Tasman Gateway at the Eocene-Oligocene boundary (33 Ma) resulted in abrupt cooling but the Oligocene became a period of high rainfall with swamps in southeast Australia. During the Miocene a warm and humid climate developed with pockets of rainforests in central Australia but before the end of the period colder and drier climate severely reduced this rainforest. A brief period of increased rainfall in the Pliocene was followed by drier climate which favoured grassland. Since then the fluctuation between wet interglacial periods and dry glacial periods has developed into the present arid regime. Australia has thus experienced various climate changes over a 15 million year period with a gradual decrease in precipitation.[49] The Tasman Gateway between Australia and Antarctica
Antarctica
began to open c. 40 to 30 Ma. Palaeontological evidences indicate the Antarctic Circumpolar Current
Antarctic Circumpolar Current
(ACC) was established in the Late Oligocene c. 23 Ma with the full opening of the Drake Passage
Drake Passage
and the deepening of the Tasman Gateway. The oldest oceanic crust in the Drake Passage, however, is 34 to 29 Ma-old which indicates spreading between the Antarctic and South American plates began near the Eocene/Oligocene boundary.[50] Deep sea environments in Tierra del Fuego and the North Scotia Ridge
North Scotia Ridge
during the Eocene
Eocene
and Oligocene indicate a "Proto-ACC" opened opened during this period. Later, 26 to 14 Ma, a series of events severally restricted the Proto-ACC: change to shallow marine conditions along the North Scotia Ridge; closure of the Fuegan Seaway, the deep sea that existed in Tierra del Fuego; and uplift of the Patagonian Cordillera. This, together with the reactivated Iceland plume, contributed to global warming. During the Miocene, the Drake Passage
Drake Passage
began to widen and as water flow between South America
South America
and the Antarctic Peninsula increased, the renewed ACC resulted in cooler global climate.[51] Since the Eocene
Eocene
the northward movement of the Australian Plate
Australian Plate
has resulted in an arc-continent collision with the Philippine and Caroline plates and the uplift of the New Guinea Highlands.[52] From the Oligocene to the late Miocene, the climate in Australia, dominated by warm and humid rainforests before this collision, began to alternate between open forest and rainforest before the continent became the arid or semiarid landscape it is today.[53] Biogeography[edit] See also: Evolutionary history of plants

Banksia, a grevilleoid Proteaceae, is an example a plant with a Gondwanan distribution

The adjective "Gondwanan" is in common use in biogeography when referring to patterns of distribution of living organisms, typically when the organisms are restricted to two or more of the now-discontinuous regions that were once part of Gondwana, including the Antarctic flora.[6] For example, the plant family Proteaceae, known from all continent in the Southern Hemisphere, has a "Gondwanan distribution" and is often described as an archaic, or relict, lineage. The distributions in Proteaceae
Proteaceae
is, nevertheless, the result of both Gondwanan rafting and later oceanic dispersal.[54] Post-Cambrian diversification[edit] During the Silurian
Silurian
Gondwana
Gondwana
extended from the Equator (Australia) to the South Pole (North Africa
Africa
and South America) whilst Laurasia
Laurasia
was located on the Equator opposite to Australia. A short-lived Late Ordovician
Ordovician
glaciation was followed by a Silurian
Silurian
Hot House period.[55] The End- Ordovician
Ordovician
extinction, which resulted in 27% of marine invertebrate families and 57% of genera going extinct, occurred during this shift from Ice House to Hot House.[56]

Reconstructions of (left) a Late Silurian
Silurian
Cooksonia, the first land plant, and (right) a Late Devonian
Devonian
Archaeopteris, the first large tree.

By the end of the Ordovician
Ordovician
Cooksonia, a slender, ground-covering plant, became the first vascular plant to establish itself on land. This first colonisation occurred exclusively around the Equator on landmasses then limited to Laurasia
Laurasia
and, in Gondwana, to Australia. In the Late Silurian
Silurian
two distinctive linages, zosterophylls and rhyniophytes, had colonised the tropics. The former evolved into the lycopods, that were to dominate the Gondwanan vegetation over a long period, whilst the latter evolved into horsetails and gymnosperms. Most of Gondwana
Gondwana
was located far from the Equator during this period and remained a lifeless and barren landscape.[57] West Gondwana
Gondwana
drifted north during the Devonian
Devonian
which brought Gondwana and Laurasia
Laurasia
close together. Global cooling contributed to the Late Devonian
Devonian
extinction (19% of marine families and 50% of genera went extinct) and glaciation occurred in South America. Before Pangaea
Pangaea
had formed terrestrial plants, such as pteridophytes, began to diversify rapidly resulting in the colonisation of Gondwana. The Baragwanathia Flora, found only in the Yea Beds of Victoria, Australia, occurs in two strata separated by 1,700 m (5,600 ft) or 30 Ma; the upper assemblage is more diverse and includes Baragwanathia, the first primitive herbaceous lycopod to evolve from the zosterophylls. During the Devonian
Devonian
giant club mosses replaced the Baragwanathia Flora, introducing the first trees, and by the Late Devonian
Devonian
this first forest was accompanied by the progymnosperms, including the first large trees Archaeopteris.[58] The Late Devonian
Devonian
extinction probably also resulted in osteolepiform fishes evolving into the amphibian tetrapods, the earliest land vertebrates, in Greenland and Russia. The only traces of this evolution in Gondwana
Gondwana
are amphibian footprints and a single jaw from Australia.[59] The closure of the Rheic Ocean
Rheic Ocean
and the formation of Pangaea
Pangaea
in the Carboniferous
Carboniferous
resulted in the rerouting of ocean currents which initiated an Ice House period. As Gondwana
Gondwana
began to rotate clockwise, Australia shifted south to more temperate latitudes. An ice cap initially covered most of southern Africa
Africa
and South America
South America
but began to spread to eventually cover most of the supercontinent, save for northern-most Africa- South America
South America
and eastern Australia. Giant lycopod and horsetail forests continued to evolve in tropical Laurasia together with a diversified assemblage of true insects. In Gondwana, in contrast, ice and, in Australia, volcanism decimated the Devonian flora to a low-diversity seed fern flora – the pteridophytes were increasingly replaced by the gymnosperms which were to dominate until the Mid-Cretaceous. Australia, however, was still located near the Equator during the Early Carboniferous
Carboniferous
and during this period temnospondyl and lepospondyl amphibians and the first amniote reptileans evolved, all closely related to the Laurasian fauna, but spreading ice eventually drove these animals away from Gondwana entirely.[60]

Fossilised Walchia
Walchia
and Utrechtia, two voltzialean pines from which modern conifers evolved

Still extant Triassic
Triassic
conifers (Agathis, Wollemia, Araucaria, and Podocarpus) that once dominated Gondwana

The Gondwana
Gondwana
ice sheet melted and sea levels dropped during the Permian
Permian
and Triassic
Triassic
global warming. During this period, the extinct glossopterids colonised Gondwana
Gondwana
and reached peak diversity in the Late Permian
Permian
when coal-forming forests covered much of Gondwana. The period also saw the evolution of Voltziales; one of the few plant orders to survive the end- Permian
Permian
extinction (57% of marine families and 83% of genera went extinct) which came to dominate in the Late Permian
Permian
and from whom true conifers evolved. Tall lycopods and horsetails dominated the wetlands of Gondwana
Gondwana
in the Early Permian. Insects co-evolved with glossopterids across Gondwana
Gondwana
and diversified with more than 200 species in 21 orders by the Late Permian, many known from South Africa
Africa
and Australia. Beetles and cockroaches remained minor elements in this fauna. Tetrapod
Tetrapod
fossils from the Early Permian
Permian
have only been found in Laurasia
Laurasia
but they became common in Gondwana
Gondwana
later during the Permian. The arrival of the therapsids resulted in the first plant-vertebrate-insect ecosystem.[61] Modern diversification[edit] During the Mid- to Late Triassic
Triassic
Hot House condition coincided with a peak in biodiversity — the end- Permian
Permian
extinction was huge and so was the radiation that followed. Two families of conifers, Podocarpaceae
Podocarpaceae
and Araucariaceae, dominated Gondwana
Gondwana
in the Early Triassic, but Dicroidium, an extinct genus of fork-leaved seed ferns, dominated woodlands and forests of Gondwana
Gondwana
during most of the Triassic. Conifers evolved and radiated during the period with six of eight extant families already present before the end of it. Bennettitales
Bennettitales
and Pentoxylales, two now extinct orders of gymnospermous plants, evolved in the Late Triassic
Triassic
and became important in the Jurassic
Jurassic
and Cretaceous. It is possible that gymnosperm biodiversity surpassed later angiosperm biodiversity and that the evolution of angiosperms began during the Triassic
Triassic
but, if so, in Laurasia
Laurasia
rather than in Gondwana. Two Gondwanan classes, lycophytes and sphenophytes, saw a gradual decline during the Triassic while ferns, though never dominant, managed to diversify.[62] The brief ice house of the Triassic–Jurassic extinction event
Triassic–Jurassic extinction event
had a dramatic impact on dinosaurs but left plants largely unaffected. The Jurassic
Jurassic
was mostly a hot house and while vertebrates managed to diversify in this environment plants has left little evidence of such development, with the exception of Cheiroleidiacean conifers and Caytoniales
Caytoniales
and other groups of seed ferns. In terms of biomass the Jurassic
Jurassic
flora was dominated by conifers families and other gymnosperms that had evolved during the Triassic. The Pteridophytes, that had dominated during the Palaeozoic, were now marginalised, except for ferns. In contrast to Laurentia, very few insect fossils have been found in Gondwana
Gondwana
to a large extent because of widespread deserts and volcanism. While plants had a cosmopolitan distribution, dinosaur evolved and diversified in a pattern that reflects the Jurassic
Jurassic
break-up of Panagaea.[63] The Cretaceous saw the arrival of the angiosperms, or flowering plants, a group that probably evolved in western Gondwana
Gondwana
(South America-Africa). From there the angiosperms diversified in two stages: the monocots and magnoliids evolved in the Early Cretaceous followed by the hammamelid dicots. By the Mid-Cretaceous angiosperms constituted half of the flora in northeastern Australia. There is, however, no obvious connection between this spectacular angiosperm radiation and an extinction event nor vertebrate/insect evolution. Insect orders associated with pollination, such as beetles, flies, butterflies and moths, and wasps, bees, and ants, radiated continuously from the Permian- Triassic
Triassic
long before the arrival of the angiosperms. Well-preserved insect fossils have been found in the lake deposits of the Santana Formation
Santana Formation
in Brazil, the Koonwarra Lake fauna in Australia, and the Orapa diamond mine
Orapa diamond mine
in Botswana.[64] Dinosaurs continued to prosper but, as the angiosperm diversified, conifers, bennettitaleans and pentoxylaleans disappeared from Gondwana
Gondwana
c. 115 Ma together with the specialised herbivorous ornithischians whilst generalist browsers such as several families of sauropodomorph Saurischia
Saurischia
prevailed. The Cretaceous–Paleogene extinction event killed off all dinosaurs except birds, but plant evolution in Gondwana was hardly affected by the event.[64] Gondwanatheria
Gondwanatheria
is an extinct group of non-therian mammals with a Gondwanan distribution (South America, Africa, Madagascar, India, and Antarctica) during the Late Cretaceous and Palaeogene.[65] Xenarthra
Xenarthra
and Afrotheria, two placental clades, are of Gondwanan origin and probably began to evolve separately c. 105 Ma when Africa
Africa
and South America
South America
separated.[66]

The Nothofagus
Nothofagus
plant genus illustrates Gondwanan distribution, having descended from the supercontinent and existing in present-day Australia, New Zealand, New Caledonia, and the Southern Cone. Fossils have also recently been found in Antarctica.[67]

The laurel forest of Australia, New Caledonia, and New Zealand
New Zealand
have a number of other related species of the laurissilva de Valdivia, through the connection of the Antarctic flora
Antarctic flora
as gymnosperms and deciduous angiosperm Nothofagus. Corynocarpus laevigatus
Corynocarpus laevigatus
is called the bay of New Zealand, Laurelia novae-zelandiae
Laurelia novae-zelandiae
belongs to the same genus Laurelia. The sempervirens tree niaouli grows in Australia, New Caledonia, and New Zealand. New Caledonia
New Caledonia
and New Zealand
New Zealand
ecoregions became separated from Australia by continental drift 85 million years ago. The islands still retain plants that originated in Gondwana
Gondwana
and spread to the Southern Hemisphere continents later. However, strong evidence exists of glaciation during the Carboniferous
Carboniferous
to Permian time, especially in South Africa.[citation needed] See also[edit]

Continental drift, the movement of the Earth's continents relative to each other Australasian realm Gondwana
Gondwana
Rainforests of Australia The Great Escarpment of Southern Africa Plate tectonics, a theory which describes the large-scale motions of Earth's lithosphere South Polar dinosaurs, which proliferated during the Early Cretaceous (145–100 Mya) while Australia was still linked to Antarctica
Antarctica
to form East Gondwana Tarkine wilderness

References[edit] Notes[edit]

^ "Gondwana". Dictionary.com. Lexico Publishing Group. Retrieved 18 January 2010.  ^ "Gondwanaland". Merriam-Webster Online Dictionary. Retrieved 18 January 2010.  ^ Torsvik & Cocks 2013, Abstract ^ Meert & Van Der Voo 1997, Abstract ^ Suess 1885, p. 768: "Wir nennen es Gondwána-Land, nach der gemeinsamen alten Gondwána-Flora, … " (We name it Gondwána-Land, after the common ancient flora of Gondwána …) ^ a b McLoughlin 2001, Gondwana
Gondwana
or Gondwanaland?, pp. 272–273 ^ Meert 2003, Fig. 10, p. 19 ^ Meert & Van Der Voo 1997, Introduction, pp. 223–226 ^ Miashita & Yamamoto 1996 ^ Meert & Van Der Voo 1997, p. 229 ^ Defined but not named in Collins & Pisarevsky 2005: "Azania" was a Greek name for the East African coast ^ Li et al. 2008, The birth of Gondwanaland (600–530 Ma), p. 201 ^ Rapalini 2001; Rapalini 1998, pp. 105–106 ^ Dalla Salda et al. 1998, Abstract; Vujovich, van Staal & Davis 2004, Conclusions, p. 1053 ^ Meert 2003, Abstract ^ Grantham, Maboko & Eglington 2003 ^ Cawood 2005, Definition and Tectonic Framework, pp. 4–6 ^ Münker & Crawford 2000, Abstract ^ Li et al. 2008, Abstract ^ Torsvik & Van Der Voo 2002, Data selection and reconstruction fits, p. 772 ^ Blakey 2003, Assembly of Western Pangaea: Carboniferous–Permian, pp. 453–454 ^ Blakey 2003, Assembly of Eastern Pangaea: Late Permian–Jurassic, p. 454 ^ Blakey 2003, Summary: significance of Pangaean events, pp. 454–455 ^ Marzoli et al. 1999, Abstract ^ " Gondwana
Gondwana
Remnants in Alabama
Alabama
And Georgia: Uchee Is An 'Exotic' Peri-Gondwanan Arc Terrane, Not Part Of Laurentia". ScienceDaily. February 4, 2008. Retrieved 2011-10-22.  ^ Torsvik & Cocks 2013, Marginal microcontinents and terranes, p. 1008 ^ Torsvik & Cocks 2013, Southern Europe, pp. 1008–1009 ^ McLoughlin 2001, Cimmerian terranes, p. 278 ^ Cawood 2005, Peri-Gondwanan continental basement assemblages, pp. 15–16 ^ Torsvik & Cocks 2013, South-central and eastern Asia ^ Jokat et al. 2003, Introduction, pp. 1–2 ^ Encarnación et al. 1996, Early rifting and Gondwana
Gondwana
breakup, pp. 537–538 ^ Royer et al. 1988, Figg. 7 a–j, pp. 248–257 ^ Abbate, Ernesto; Bruni, Piero; Sagri, Mario (2015). "Geology of Ethiopia: A Review and Geomorphological Perspectives". In Billi, Paolo. Landscapes and Landforms of Ethiopia. World Geomorphological Landscapes. pp. 33–64. ISBN 978-94-017-8026-1.  ^ Coltorti, M.; Dramis, F.; Ollier, C.D. (2007). "Planation surfaces in Northern Ethiopia". Geomorphology. 89: 287–296.  access-date= requires url= (help) ^ Powell, Roots & Veevers 1988, Abstract ^ DeMets, Gordon & Royer 2005, Introduction; Fig. 1, p. 446 ^ Müller, Royer & Lawver 1993, Model results, pp. 277–278 ^ McLoughlin 2001, East Antarctica–Australia, p. 280 ^ McLoughlin 2001, West Antarctica–Tasmantia, p. 280 ^ Seton et al. 2012, South Atlantic, pp. 217–218 ^ a b c Ramos 2009, Abstract ^ a b c d e Charrier, Pinto & Rodríguez 2006, pp. 45–46 ^ a b Hoffmann-Rothe et al. 2006 ^ Yin & Harrison 2000, Abstract ^ Luyendyk, Forsyth & Phillips 1972, Abstract ^ Jestin, Huchon & Gaulier 1994, Abstract ^ Martin 2006, Palaeogeography, pp. 538–539 ^ Martin 2006, Conclusions, pp. 557–558 ^ Lagabrielle et al. 2009, Timing of opening of the Drake Passage region, pp. 198–199 ^ Lagabrielle et al. 2009, Conclusions, p. 210 ^ Hill & Hall 2003, Abstract ^ Travouillon et al. 2009, Abstract ^ Barker et al. 2007, Abstract ^ Anderson et al. 1999, SILURIAN: terrestrial life appears in the tropics, p. 148 ^ Anderson et al. 1999, The First Extinction, p. 151 ^ Anderson et al. 1999, The Silurian
Silurian
revolution, p. 151 ^ Anderson et al. 1999, DEVONIAN: colonising Gondwana; The Second Extinction; Global colonisation of plants, pp. 151, 153 ^ Anderson et al. 1999, Amphibian prelude, p. 153 ^ Anderson et al. 1999, CARBONIFEROUS: competing with ice, pp. 153–154 ^ Anderson et al. 1999, PERMIAN: the glossopterid empire, pp. 153–154 ^ Anderson et al. 1999, TRIASSIC: the gymnosperm heyday, pp. 155–156 ^ Anderson et al. 1999, JURASSIC: volcanism, conifers and bennettitaleans, pp. 156, 158 ^ a b Anderson et al. 1999, CRETACEOUS: of flowers and pollination, pp. 158–159 ^ Gurovich & Beck 2009, Introduction, pp. 25–26 ^ Woodburne, Rich & Springer 2003, Gondwana
Gondwana
and early mammal evolution, p. 375 ^ HaoMin & ZheKun 2007

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External links[edit]

Houseman, Greg. "Animation of the dispersal of Gondwanaland". University of Leeds. Retrieved 21 October 2008.  Barend Köbben; Colin Reeves; Maarten de Wit. "Interactive animation of the breakup of Gondwana". ITC, University of Twente. Retrieved 16 October 2017.  Graphical subjects dealing with Tectonics and Paleontology Gondwana
Gondwana
Reconstruction and Dispersion The Gondwana
Gondwana
Map Project

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