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Intermediate disturbance hypothesis
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The intermediate disturbance hypothesis (IDH) suggests that local species diversity is maximized when ecological disturbance is neither too rare nor too frequent. At high levels of disturbance, due to frequent forest fires or human impacts like deforestation, all species are at risk of going extinct. According to IDH theory, at intermediate levels of disturbance, diversity is thus maximized because species that thrive at both early and late successional stages can coexist. IDH is a nonequilibrium model used to describe the relationship between disturbance and species diversity. IDH is based on the following premises: First, ecological disturbances have major effects on species richness within the area of disturbance.[1][2][3] Second, interspecific competition results from one species driving a competitor to extinction and becoming dominant in the ecosystem.[1][2][3] Third, moderate ecological scale disturbances prevent interspecific competition.[1][2][3]
Graph shows principles of intermediate disturbance hypothesis: I. at low levels of ecological disturbance species richness decreases as competitive exclusion increases, II. at intermediate levels of disturbance, diversity is maximized because species that thrive at both early and late successional stages can coexist, III. at high levels of disturbance species richness is decreased due an increase in species movement.
Disturbances act to disrupt stable ecosystems and clear species' habitat. As a result, disturbances lead to species movement into the newly cleared area.[1] Once an area is cleared there is a progressive increase in species richness and competition takes place again. Once disturbance is removed, species richness decreases as competitive exclusion increases.[4] "Gause's Law", also known as competitive exclusion, explains how species that compete for the same resources cannot coexist in the same niche.[2] Each species handles change from a disturbance differently; therefore, IDH can be described as both "broad in description and rich in detail".[1] The broad IDH model can be broken down into smaller divisions which include spatial within-patch scales, spatial between-patch scales, and purely temporal models.[4] Each subdivision within this theory generates similar explanations for the coexistence of species with habitat disturbance. Joseph H. Connell[5] proposed that relatively low disturbance leads to decreased diversity and high disturbance causes an increase in species movement. These proposed relationships lead to the hypothesis that intermediate disturbance levels would be the optimal amount of disorder within an ecosystem. Once K-selected and r-selected species can live in the same region, species richness can reach its maximum. The main difference between both types of species is their growth and reproduction rate. These characteristics attribute to the species that thrive in habitats with higher and lower amounts of disturbance. K-selected species generally demonstrate more competitive traits. Their primary investment of resources is directed towards growth, causing them to dominate stable ecosystems over a long period of time; an example of K-selected species the African elephant, which is prone to extinction because of their long generation times and low reproductive rates. In contrast, r-selected species colonize open areas quickly and can dominate landscapes that have been recently cleared by disturbance.[3] An ideal examples of r-selected groups are algae. Based on the contradictory characteristics of both of these examples, areas of occasional disturbance allow both r and K species to benefit by residing in the same area. The ecological effect on species relationships is therefore supported by the intermediate disturbance hypothesis.
Disturbed vegetation due to milpa farming. Cayo District, Belize. [Macrae 2008].
Contents
[hide]
1 History
2 Support and critiques
3 See also
4 References
History[edit]
David Wilkinson gives a thorough history of the hypothesis in his paper titled, "The disturbing history of the intermediate disturbance".[1] In this paper, he explains that the idea of disturbance relating to species richness can be traced back to the 1940s in Eggeling 1947,[6] Watt 1947,[7] and Tansley 1949.[8] Though studies supporting the hypothesis began in the 1960s, the first concrete statements of the intermediate disturbance hypothesis didn't occur until the 1970s.[1] The hypothesis was initially illustrated using what has been referred to as a "hump-backed model", which graphed the proposed relationship between diversity and disturbance.[1] This graph appeared first in Grime's 'Competitive exclusion in herbaceous vegetation'[9] where it was used to show the relationship between species density and both environmental stress and intensity of management. The graph appears again in Horn's 'Markovian properties of forest succession'[10] and Connell's 'The influence of interspecific competition and other factors on the distribution of the barnacle'.[5] Though Grime was the first to provide a model for the relationship and Horn was the first to explicitly state the hypothesis, Connell is generally cited in text books and journals as the founder of the hypothesis.[1]
The hypothesis caused concern among the marine science community because of the discrepancy with the 1976 Competition/Predation/Disturbance model proposed by Menge and Sutherland[11] In this model, low disturbance influences high predation and high disturbance creates low predation, causing competitive exclusion to take place. Menge & Sutherland formulated a new model, one that incorporated Connell's ideas in a two part graph published in The American Naturalist (1987).[11] This model proposes that predation, competition, and disturbance are all responsible for shaping the diversity of a community under certain circumstances.
Research regarding the effects of intermediate disturbance is ongoing. In one study, dry and tropical forest regions were compared to determine how the effects of IDH change due to varying climate.[12] More recently, the intermediate disturbance hypothesis has been examined in marine and freshwater ecosystems[13][14] and protist microcosms.[15]
Support and critiques[edit]
Debates over the validity of the IDH are ongoing within the discipline of tropical ecology as the theory is tested in various ecological communities. Other evidence exists for[15][16] and against[17][18] the hypothesis. The intermediate disturbance hypothesis has been supported by several studies involving marine habitats such as coral reefs and macroalgal communities. In shallow coastal waters off of south-west Western Australia, a study was conducted to determine whether or not the extremely high diversity observed in macroalgal communities was due to disturbance from waves.[13] Using a numerical wave model to estimate the forces caused by waves, researchers were able to determine that there was a significant relationship between species diversity and disturbance index; this is consistent with the intermediate disturbance hypothesis.[13] Furthermore, diversity was lower at exposed offshore sites where disturbance from waves was highest, and at extremely sheltered site where disturbance from waves was minimized.[13] The study provided evidence that biodiversity in microalgal reef communities possess some relationship with their proximity to the outer edge of lagoon systems typical of the Western Australian coast.[13] While this study may have been localized to the Western Australian coast, it still provides some evidence to support the validity of the IDH.
Additionally, a study done in the Virgin Islands National Park found that species diversity, in some locations, of shallow coral reefs increased after infrequent hurricane disturbance.[14] In 1982, reefs in Kona, Hawaii were reported to have an increase in diversity after a moderate storm, although the effects of the storm varied with the reef zones.[14] In 1980, Hurricane Allen increased species diversity in shallow zones of the Discovery Bay Reef in Jamaica because the more dominant corals were reduced; giving the other types a chance to propagate following the disturbance.[14] Similar findings have been reported in shallow reefs in which dominant species of coral have suffered more damage than the less common species.[14] While more long-term studies are required to completely support the Intermediate Disturbance Hypothesis, the studies that have been conducted thus far have proven that IDH does have some validity while attempting to describe the relationship between diversity and the rate of occurrences of disturbance in an area.
Even though the IDH was designed for species-rich environments, like tropical rainforests, "most studies that evaluate the IDH are based on limited data with: few species, a limited range of disturbance and/or only a small geographic area, compared with the scale of interest".[19] In this experiment, Rogers, Poorter, Hawthorne, and Sheil evaluate the IDH on a larger scale and compare different tropical forest types in Ghana. Their dataset consisted of 2504 one-hectare plots with a total of 331,567 trees. These plots were divided classified into three forest types: wet (446 plots), moist (1322 plots), and dry forest (736 plots).[19] They found that diversity does peak at intermediate level of disturbance but little variation is explained outside dry forests. Therefore, disturbance is less important for species diversity patterns in wet tropical rain forests than previously thought. The number of species was about the same for each forest type, and wet forests had only slightly fewer pioneer species, slightly more shade-tolerant and an equal number of pioneer light-demanders compared with the moist and dry forests.[19] Their results generally supported the IDH as an explanation of why diversity varies across sites, but concluded that disturbance is less important for species richness patterns in wet tropical rain forests than previously thought.[19]
IDH has been subject to criticism since its inception but not to the degree that other species density hypotheses have been. Recently there has been a call for a critical reassessment of IDH.[20] Criticisms have focused on the increasing amount of empirical data that disagrees with IDH. This can be found within approximately 80% of over 100 reviewed studies that are examining the predicted peak of diversity in intermediate disturbance levels.[15][20][21][22] The rationales behind these discrepancies have been leveled at the simplicity of IDH and its inability to grasp the complexity found within the spatial and intensity aspects of disturbance relationships.[23] In addition, many IDH proven circumstances have been suggested to be a reflection of skewed research methods based on researchers looking for humped diversity-disturbance relations only in systems where they believed it has occurred.[20] Other criticisms are suggesting several subtle theoretical issues with IDH. First, while disturbances weaken competition by reducing species densities and per-capita growth rates, it also reduces the strength of competition needed to push per capita growth into a negative territory and reduce density to zero.[20][23] Second, intermediate disturbances slow competitive exclusion by increasing the long-term average mortality rate, and thereby reducing the differences in the average growth rates of competing species. The difference in the growth rates between competitively superior and inferior species determines the rates of competitive exclusion; therefore intermediate disturbances are affecting species' abundance but not coexistence.[20] Third, intermediate disturbances temporarily affect relative species fitness. However, no matter what the rate of disturbance is, the species with favored fitness will out-compete the rest of the species.[24]
Several alternatives hypotheses have been proposed. One example is by Denslow,[25] he states that the species diversity in a disturbance-mediated coexistence between species is maximized by the presence of a disturbance regime resembling the historic processes. This is because species generally adapt to the level of disturbance in their ecosystem through evolution (whether disturbance is of high, intermediate or low level). Many species (e.g. ruderal plants, fire-adapted species) even depend on disturbance in ecosystems where it often occurs.
(Disturbed vegetation due to milpa farming, Contreras Valley, Cayo District, Belize [Macrae 2010].
Disturbance due to tree fall, Gainesville, Florida [Daniel 2012].
See also[edit]
Competition-colonization trade-off
Ecological succession
Habitat fragmentation
Patch dynamics
References[edit]
^ Jump up to: a b c d e f g h i Wilkinson, David M. (1999). "The Disturbing History of Intermediate Disturbance". Oikos. 84 (1): 145–7. doi:10.2307/3546874. JSTOR 3546874.
^ Jump up to: a b c d Kricher, John C. (2011). Tropical Ecology. New Jersey, Princeton: Princeton University Press. [page needed]
^ Jump up to: a b c d Catford, Jane A.; Daehler, Curtis C.; Murphy, Helen T.; Sheppard, Andy W.; Hardesty, Britta D.; Westcott, David A.; Rejmánek, Marcel; Bellingham, Peter J.; et al. (2012). "The intermediate disturbance hypothesis and plant invasions: Implications for species richness and management". Perspectives in Plant Ecology, Evolution and Systematics. 14 (3): 231–41. doi:10.1016/j.ppees.2011.12.002.
^ Jump up to: a b Vandermeer, John; Boucher, Douglas; Perfecto, Ivette; de la Cerda, Inigo Granzow (1996). "A Theory of Disturbance and Species Diversity: Evidence from Nicaragua After Hurricane Joan". Biotropica. 28 (4): 600–13. doi:10.2307/2389100. JSTOR 2389100.
^ Jump up to: a b Connell, J. H. (1978). "Diversity in Tropical Rain Forests and Coral Reefs". Science. 199 (4335): 1302–10. Bibcode:1978Sci...199.1302C. doi:10.1126/science.199.4335.1302. PMID 17840770.
Jump up ^ Eggeling, W. J. (1947). "Observations on the Ecology of the Budongo Rain Forest, Uganda". Journal of Ecology. 34 (1): 20–87. doi:10.2307/2256760. JSTOR 2256760.
Jump up ^ Watt, Alex S. (1947). "Pattern and Process in the Plant Community". Journal of Ecology. 35 (1/2): 1–22. doi:10.2307/2256497. JSTOR 2256497.
Jump up ^ Tansley, A. G. (1949). Britain's Green Mantle. London: George Allen and Unwin. [page needed]
Jump up ^ Grime, J. P. (1973). "Competitive Exclusion in Herbaceous Vegetation". Nature. 242 (5396): 344–7. Bibcode:1973Natur.242..344G. doi:10.1038/242344a0.
Jump up ^ Horn, H. S. (1975). "Markovian Properties of Forest Succession". In Cody, M. L.; Diamond, J. M. Ecology and evolution of communities. Massachusetts: Belknap Press. pp. 196–211. ISBN 0-674-22444-2.
^ Jump up to: a b Menge, Bruce A.; Sutherland, John P. (1987). "Community regulation: variation in disturbance, competition, and predation in relation to environmental stress and recruitment". The American Naturalist. 130 (5): 730–57. doi:10.1086/284741. JSTOR 2461716.
Jump up ^ Sousa, Wayne P. (1979). "Disturbance in Marine Intertidal Boulder Fields: The Nonequilibrium Maintenance of Species Diversity". Ecology. 60 (6): 1225–39. doi:10.2307/1936969. JSTOR 1936969.
^ Jump up to: a b c d e England, Phillip R.; Phillips, Julia; Waring, Jason R.; Symonds, Graham; Babcock, Russell (2008). "Modelling wave-induced disturbance in highly biodiverse marine macroalgal communities: Support for the intermediate disturbance hypothesis". Marine and Freshwater Research. 59 (6): 515. doi:10.1071/MF07224.
^ Jump up to: a b c d e Rogers, C. S. (1993). "Hurricanes and coral reefs: The intermediate disturbance hypothesis revisited". Coral Reefs. 12 (3–4): 127–37. Bibcode:1993CorRe..12..127R. doi:10.1007/BF00334471.
^ Jump up to: a b c Mackey, Robin L.; Currie, David J. (2001). "The Diversity-Disturbance Relationship: Is It Generally Strong and Peaked?". Ecology. 82 (12): 3479–92. doi:10.2307/2680166. JSTOR 2680166.
Jump up ^ Randall Hughes, A.; Byrnes, Jarrett E.; Kimbro, David L.; Stachowicz, John J. (2007). "Reciprocal relationships and potential feedbacks between biodiversity and disturbance". Ecology Letters. 10 (9): 849–64. doi:10.1111/j.1461-0248.2007.01075.x. PMID 17663718.
Jump up ^ Collins, Scott L.; Glenn, Susan M.; Gibson, David J. (1995). "Experimental Analysis of Intermediate Disturbance and Initial Floristic Composition: Decoupling Cause and Effect". Ecology. 76 (2): 486–92. doi:10.2307/1941207. JSTOR 1941207.
Jump up ^ Warren, Philip H. (1996). "Dispersal and Destruction in a Multiple Habitat System: An Experimental Approach Using Protist Communities". Oikos. 77 (2): 317–25. doi:10.2307/3546071. JSTOR 3546071.
^ Jump up to: a b c d Bongers, Frans; Poorter, Lourens; Hawthorne, William D.; Sheil, Douglas (2009). "The intermediate disturbance hypothesis applies to tropical forests, but disturbance contributes little to tree diversity". Ecology Letters. 12 (8): 798–805. doi:10.1111/j.1461-0248.2009.01329.x. PMID 19473218.
^ Jump up to: a b c d e Fox, Jeremy W. (2013). "The intermediate disturbance hypothesis should be abandoned". Trends in Ecology & Evolution. 28 (2): 86–92. doi:10.1016/j.tree.2012.08.014. PMID 22981468.
Jump up ^ Scholes, Lianna; Warren, Philip H.; Beckerman, Andrew P. (2005). "The combined effects of energy and disturbance on species richness in protist microcosms". Ecology Letters. 8 (7): 730–8. doi:10.1111/j.1461-0248.2005.00777.x.
Jump up ^ Lubchenco, Jane (1978). "Plant Species Diversity in a Marine Intertidal Community: Importance of Herbivore Food Preference and Algal Competitive Abilities". American Naturalist. 112 (983): 23–39. doi:10.1086/283250. JSTOR 2460135.
^ Jump up to: a b Chesson, Peter; Huntly, Nancy (1997). "The Roles of Harsh and Fluctuating Conditions in the Dynamics of Ecological Communities". The American Naturalist. 150 (5): 519–53. doi:10.1086/286080. JSTOR 286080. PMID 18811299.
Jump up ^ Violle, Cyrille Violle, Zhichao Pu, Lin Jiang; Pu, Zhichao; Jiang, Lin; Schoener, Thomas W. (2010). "Experimental demonstration of the importance of competition under disturbance". Proceedings of the National Academy of Sciences. 107 (29): 12925–9. Bibcode:2010PNAS..10712925V. doi:10.1073/pnas.1000699107. JSTOR 25708640. PMC 2919955  . PMID 20616069.
Jump up ^ Hall, A. R.; Miller, A. D.; Leggett, H. C.; Roxburgh, S. H.; Buckling, A.; Shea, K. (2012). "Diversity-disturbance relationships: Frequency and intensity interact". Biology Letters. 8 (5): 768–71. doi:10.1098/rsbl.2012.0282. PMC 3440969  . PMID 22628097.
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Gondwana
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This article is about the supercontinent. For other uses, see Gondwana (disambiguation).
Gondwana 420 million years ago. View centered on South Pole.
Gondwana ( /ɡɒndˈwɑːnə/),[1] or Gondwanaland,[2] was a supercontinent that existed from the Neoproterozoic (about 550 million years ago) until the 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 Era, covering an area of about 100,000,000 km2 (39,000,000 sq mi).[3] During the Carboniferous, it merged with Euramerica to form a larger supercontinent called Pangaea. Gondwana (and Pangaea) gradually broke up during the Mesozoic Era. The remnants of Gondwana make up about two-thirds of today's continental area.
The formation of 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 with Africa and the addition of Australia and Antarctica, respectively.[4]
Contents
[hide]
1 Origin of concept
2 Formation
3 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 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 was named by Austrian scientist Eduard Suess, after the Gondwana region of central northern India which is derived from 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 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 in blue and 570 to 530 Ma collisional metamorphism of the Kuunga orogeny in red.[7]
The assembly of Gondwana was a protracted process during the Neoproterozoic and Paleozoic, which however remains incompletely understood 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 (Africa and South America). Three orogenies were recognized during the 1990s: the East African Orogeny (650 to 800 Ma) and Kuunga orogeny (including the Malagasy Orogeny in southern Madagascar) (550 Ma) – the collision between East Gondwana and East 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 between Laurentia and western Gondwana.[9] During this interval, the Cambrian explosion occurred. Laurentia was docked against the western shores of a united Gondwana for a short period near the Precambrian/Cambrian boundary, forming the short-lived and still disputed supercontinent Pannotia.[10]
The Mozambique Ocean separated the Congo–Tanzania–Bangweulu Block of central Africa from Neoproterozoic India (India, the 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 and parts of 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 had already been amalgamated. By c. 550 Ma, India had reached its Gondwana position which initiated the 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 brought India next to Australia–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 of South America, including a piece transferred from Laurentia when the west edge of Gondwana scraped against southeast Laurentia in the Ordovician.[13] This is the Cuyania or Precordillera terrane of the Famatinian orogeny in northwest Argentina which may have continued the line of the Appalachians southwards.[14]
Reconstruction showing final stages of assembly of Gondwana, 550 Mya
As the rest of Gondwana formed, a complex series of orogenic events assembled the eastern parts of 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 (in the Kenya-Tanzania region) in the East African Orogeny c.750 to 620 Ma. Then Australia and East Antarctica were merged with the remaining 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 and southern India. In it, Neoproterozoic 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 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 as part of Pangaea[edit]
Gondwana formed part of Pangaea for c. 150 Ma[19]
Main article: Pangaea
Gondwana and Laurussia formed the Pangaea supercontinent during the Carboniferous. Pangaea began to break up in the Mid-Jurassic when the Central Atlantic opened.[20]
In the western end of Pangaea, the collision between Gondwana and 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 collided with Siberia and Kazakhstania which resulted in the Uralian orogeny and Laurasia. Pangaea was finally amalgamated in the Late Carboniferous-Early Permian but the oblique forces continued until 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 during the middle 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 to form the Palaeo-Thethys and Neo-Tethys oceans in the Late Carboniferous and docked with Asia during the Triassic and Jurassic. Western Pangaea began to rift while the eastern end was still being assembled.[22]
The formation of 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 extinction event and left large deposits of hydrocarbons, coal, evaporite, and metals.[23]
The break-up of 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 extinction event.[24] The reformed Gondwanan continent was not precisely the same as that which had existed before Pangaea formed; for example, most of Florida and southern Georgia and Alabama is underlain by rocks that were originally part of Gondwana, but this region stayed attached to North America when the Central Atlantic opened.[25]
Peri-Gondwana[edit]
A large number of terranes were accreted to 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 is not known.[26]
In the Early Palaeozoic the Armorican terrane, which today form large parts of France, was part of either Peri-Gondwana or core Gondwana; the Rheic Ocean closed in front of it and the Palaeo-Tethys Ocean opened behind it. Precambrian rocks from the Iberian Peninsula suggest it too probably formed part of core Gondwana before its detachment as an orocline in the Variscan orogeny close to the Carboniferous–Permian boundary.[27]
South-east 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 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 during Late Permian and Early Jurassic. Thirdly, in the Late Triassic to Late Jurassic, Lhasa, West Burma, Woyla terranes opened the Neo-Tethys Ocean; Lhasa collided with 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 during the closure of that ocean.[29]
Gondwana's long, northern margin had remained a mostly passive margin throughout the Palaeozoic. The Early 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 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 are found in what is now the southern Weddell Sea where initial break-up occurred during the Jurassic c. 160 to 180 Ma.[31]
Opening of western Indian Ocean[edit]
The first ocean floor formed between Madagascar and Africa c. 150 Ma (left) and between India and Madagscar c. 70 Ma (right).
Gondwana began to break up in the early Jurassic following the extensive and fast emplacement of the Karoo-Ferrar flood basalts c. 184 Ma. Before the Karoo plume initiated rifting between 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 and Ellsworth–Whitmore Mountains (in Antarctica) were rotated 90° in opposite directions; and South America south of the Gastre Fault (often referred to as Patagonia) was pushed westward.[32] The history of the Africa-Antarctica break-up can be studied in great detail in the fracture zones and magnetic anomalies flanking the Southwest Indian Ridge.[33]
The Madagascar block and the Mascarene Plateau, stretching from the Seychelles to Réunion, were broken off India; elements of this breakup nearly coincide with the Cretaceous–Paleogene extinction event. The India–Madagascar–Seychelles separations appear to coincide with the eruption of the Deccan basalts, whose eruption site may survive as the Réunion hotspot. The Seychelles and the Maldives are now separated by the Central Indian Ridge.
During the initial break-up in the Early Jurassic a marine transgression swept over the 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 and 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 then began to break up c. 132.5 to 96 Ma when India moved northwest from Australia-Antarctica.[36] The Indian Plate and the Australian Plate are now separated by the Capricorn Plate and its diffuse boundaries.[37] During the opening of the Indian Ocean, the Kerguelen hotspot first formed the Kerguelen Plateau on the Antarctic Plate c. 118 to 95 Ma and then the Ninety East Ridge on the Indian Plate at c. 100 Ma.[38] The 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 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 and as oceanic crust started to separate the continents during the 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 c. 84 Ma.[40]
Opening of South Atlantic Ocean[edit]
At c. 126 Ma (left) the Falkland Plateau began to slide past southern 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 divided West 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 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 and Africa, and this westward movement lasted until the Early Cretaceous 126.7 Ma. From there rifting propagated northward during the Late 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 and 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 System and the 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 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 in the 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 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 began to collide with 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 the orogen resulted in the construction of the Tibetan Plateau between the Tethyan Himalayas in the south and the Kunlun and Qilian mountains in the north.[45]
Later, South America was connected to North America via the Isthmus of Panama, cutting off a circulation of warm water and thereby making the Arctic colder,[46] as well as allowing the Great American Interchange.
The breakup of Gondwana can be said to continue in eastern Africa at the Afar Triple Junction, which separates the Arabian, Nubian, and Somali plates, resulting in rifting in the Red Sea and East African Rift.[47]
Australia–Antarctica separation[edit]
In the Early Cenozoic Australia was still connected to 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 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 began to open c. 40 to 30 Ma. Palaeontological evidences indicate the Antarctic Circumpolar Current (ACC) was established in the Late Oligocene c. 23 Ma with the full opening of the 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 during the 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 began to widen and as water flow between South America and the Antarctic Peninsula increased, the renewed ACC resulted in cooler global climate.[51]
Since the Eocene the northward movement of the 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 is, nevertheless, the result of both Gondwanan rafting and later oceanic dispersal.[54]
Post-Cambrian diversification[edit]
During the Silurian Gondwana extended from the Equator (Australia) to the South Pole (North Africa and South America) whilst Laurasia was located on the Equator opposite to Australia. A short-lived Late Ordovician glaciation was followed by a Silurian Hot House period.[55] The End-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 Cooksonia, the first land plant, and (right) a Late Devonian Archaeopteris, the first large tree.
By the end of the 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 and, in Gondwana, to Australia. In the Late 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 was located far from the Equator during this period and remained a lifeless and barren landscape.[57]
West Gondwana drifted north during the Devonian which brought Gondwana and Laurasia close together. Global cooling contributed to the Late Devonian extinction (19% of marine families and 50% of genera went extinct) and glaciation occurred in South America. Before 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 giant club mosses replaced the Baragwanathia Flora, introducing the first trees, and by the Late Devonian this first forest was accompanied by the progymnosperms, including the first large trees Archaeopteris.[58] The Late 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 are amphibian footprints and a single jaw from Australia.[59]
The closure of the Rheic Ocean and the formation of Pangaea in the Carboniferous resulted in the rerouting of ocean currents which initiated an Ice House period. As Gondwana began to rotate clockwise, Australia shifted south to more temperate latitudes. An ice cap initially covered most of southern Africa and South America but began to spread to eventually cover most of the supercontinent, save for northern-most Africa-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 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 and Utrechtia, two voltzialean pines from which modern conifers evolved
Still extant Triassic conifers (Agathis, Wollemia, Araucaria, and Podocarpus) that once dominated Gondwana
The Gondwana ice sheet melted and sea levels dropped during the Permian and Triassic global warming. During this period, the extinct glossopterids colonised Gondwana and reached peak diversity in the Late 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 extinction (57% of marine families and 83% of genera went extinct) which came to dominate in the Late Permian and from whom true conifers evolved. Tall lycopods and horsetails dominated the wetlands of Gondwana in the Early Permian. Insects co-evolved with glossopterids across Gondwana and diversified with more than 200 species in 21 orders by the Late Permian, many known from South Africa and Australia. Beetles and cockroaches remained minor elements in this fauna. Tetrapod fossils from the Early Permian have only been found in Laurasia but they became common in 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 Hot House condition coincided with a peak in biodiversity — the end-Permian extinction was huge and so was the radiation that followed. Two families of conifers, Podocarpaceae and Araucariaceae, dominated Gondwana in the Early Triassic, but Dicroidium, an extinct genus of fork-leaved seed ferns, dominated woodlands and forests of 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 and Pentoxylales, two now extinct orders of gymnospermous plants, evolved in the Late Triassic and became important in the Jurassic and Cretaceous. It is possible that gymnosperm biodiversity surpassed later angiosperm biodiversity and that the evolution of angiosperms began during the Triassic but, if so, in 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 had a dramatic impact on dinosaurs but left plants largely unaffected. The 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 and other groups of seed ferns. In terms of biomass the 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 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 break-up of Panagaea.[63]
The Cretaceous saw the arrival of the angiosperms, or flowering plants, a group that probably evolved in western 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 long before the arrival of the angiosperms. Well-preserved insect fossils have been found in the lake deposits of the Santana Formation in Brazil, the Koonwarra Lake fauna in Australia, and the Orapa diamond mine in Botswana.[64] Dinosaurs continued to prosper but, as the angiosperm diversified, conifers, bennettitaleans and pentoxylaleans disappeared from Gondwana c. 115 Ma together with the specialised herbivorous ornithischians whilst generalist browsers such as several families of sauropodomorph 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 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 and Afrotheria, two placental clades, are of Gondwanan origin and probably began to evolve separately c. 105 Ma when Africa and South America separated.[66]
The 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 have a number of other related species of the laurissilva de Valdivia, through the connection of the Antarctic flora as gymnosperms and deciduous angiosperm Nothofagus. Corynocarpus laevigatus is called the bay of New Zealand, Laurelia novae-zelandiae belongs to the same genus Laurelia. The sempervirens tree niaouli grows in Australia, New Caledonia, and New Zealand. New Caledonia and New Zealand ecoregions became separated from Australia by continental drift 85 million years ago. The islands still retain plants that originated in Gondwana and spread to the Southern Hemisphere continents later. However, strong evidence exists of glaciation during the 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 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 to form East Gondwana
Tarkine wilderness
References[edit]
Notes[edit]
Jump up ^ "Gondwana". Dictionary.com. Lexico Publishing Group. Retrieved 18 January 2010.
Jump up ^ "Gondwanaland". Merriam-Webster Online Dictionary. Retrieved 18 January 2010.
Jump up ^ Torsvik & Cocks 2013, Abstract
Jump up ^ Meert & Van Der Voo 1997, Abstract
Jump up ^ 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 …)
^ Jump up to: a b McLoughlin 2001, Gondwana or Gondwanaland?, pp. 272–273
Jump up ^ Meert 2003, Fig. 10, p. 19
Jump up ^ Meert & Van Der Voo 1997, Introduction, pp. 223–226
Jump up ^ Miashita & Yamamoto 1996
Jump up ^ Meert & Van Der Voo 1997, p. 229
Jump up ^ Defined but not named in Collins & Pisarevsky 2005: "Azania" was a Greek name for the East African coast
Jump up ^ Li et al. 2008, The birth of Gondwanaland (600–530 Ma), p. 201
Jump up ^ Rapalini 2001; Rapalini 1998, pp. 105–106
Jump up ^ Dalla Salda et al. 1998, Abstract; Vujovich, van Staal & Davis 2004, Conclusions, p. 1053
Jump up ^ Meert 2003, Abstract
Jump up ^ Grantham, Maboko & Eglington 2003
Jump up ^ Cawood 2005, Definition and Tectonic Framework, pp. 4–6
Jump up ^ Münker & Crawford 2000, Abstract
Jump up ^ Li et al. 2008, Abstract
Jump up ^ Torsvik & Van Der Voo 2002, Data selection and reconstruction fits, p. 772
Jump up ^ Blakey 2003, Assembly of Western Pangaea: Carboniferous–Permian, pp. 453–454
Jump up ^ Blakey 2003, Assembly of Eastern Pangaea: Late Permian–Jurassic, p. 454
Jump up ^ Blakey 2003, Summary: significance of Pangaean events, pp. 454–455
Jump up ^ Marzoli et al. 1999, Abstract
Jump up ^ "Gondwana Remnants in Alabama And Georgia: Uchee Is An 'Exotic' Peri-Gondwanan Arc Terrane, Not Part Of Laurentia". ScienceDaily. February 4, 2008. Retrieved 2011-10-22.
Jump up ^ Torsvik & Cocks 2013, Marginal microcontinents and terranes, p. 1008
Jump up ^ Torsvik & Cocks 2013, Southern Europe, pp. 1008–1009
Jump up ^ McLoughlin 2001, Cimmerian terranes, p. 278
Jump up ^ Cawood 2005, Peri-Gondwanan continental basement assemblages, pp. 15–16
Jump up ^ Torsvik & Cocks 2013, South-central and eastern Asia
Jump up ^ Jokat et al. 2003, Introduction, pp. 1–2
Jump up ^ Encarnación et al. 1996, Early rifting and Gondwana breakup, pp. 537–538
Jump up ^ Royer et al. 1988, Figg. 7 a–j, pp. 248–257
Jump up ^ 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.
Jump up ^ Coltorti, M.; Dramis, F.; Ollier, C.D. (2007). "Planation surfaces in Northern Ethiopia". Geomorphology. 89: 287–296.
Jump up ^ Powell, Roots & Veevers 1988, Abstract
Jump up ^ DeMets, Gordon & Royer 2005, Introduction; Fig. 1, p. 446
Jump up ^ Müller, Royer & Lawver 1993, Model results, pp. 277–278
Jump up ^ McLoughlin 2001, East Antarctica–Australia, p. 280
Jump up ^ McLoughlin 2001, West Antarctica–Tasmantia, p. 280
Jump up ^ Seton et al. 2012, South Atlantic, pp. 217–218
^ Jump up to: a b c Ramos 2009, Abstract
^ Jump up to: a b c d e Charrier, Pinto & Rodríguez 2006, pp. 45–46
^ Jump up to: a b Hoffmann-Rothe et al. 2006
Jump up ^ Yin & Harrison 2000, Abstract
Jump up ^ Luyendyk, Forsyth & Phillips 1972, Abstract
Jump up ^ Jestin, Huchon & Gaulier 1994, Abstract
Jump up ^ Martin 2006, Palaeogeography, pp. 538–539
Jump up ^ Martin 2006, Conclusions, pp. 557–558
Jump up ^ Lagabrielle et al. 2009, Timing of opening of the Drake Passage region, pp. 198–199
Jump up ^ Lagabrielle et al. 2009, Conclusions, p. 210
Jump up ^ Hill & Hall 2003, Abstract
Jump up ^ Travouillon et al. 2009, Abstract
Jump up ^ Barker et al. 2007, Abstract
Jump up ^ Anderson et al. 1999, SILURIAN: terrestrial life appears in the tropics, p. 148
Jump up ^ Anderson et al. 1999, The First Extinction, p. 151
Jump up ^ Anderson et al. 1999, The Silurian revolution, p. 151
Jump up ^ Anderson et al. 1999, DEVONIAN: colonising Gondwana; The Second Extinction; Global colonisation of plants, pp. 151, 153
Jump up ^ Anderson et al. 1999, Amphibian prelude, p. 153
Jump up ^ Anderson et al. 1999, CARBONIFEROUS: competing with ice, pp. 153–154
Jump up ^ Anderson et al. 1999, PERMIAN: the glossopterid empire, pp. 153–154
Jump up ^ Anderson et al. 1999, TRIASSIC: the gymnosperm heyday, pp. 155–156
Jump up ^ Anderson et al. 1999, JURASSIC: volcanism, conifers and bennettitaleans, pp. 156, 158
^ Jump up to: a b Anderson et al. 1999, CRETACEOUS: of flowers and pollination, pp. 158–159
Jump up ^ Gurovich & Beck 2009, Introduction, pp. 25–26
Jump up ^ Woodburne, Rich & Springer 2003, Gondwana and early mammal evolution, p. 375
Jump up ^ 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 Reconstruction and Dispersion
The Gondwana Map Project
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Gondwana – Afrikaans
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Gondwana – Asturian
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https://az.wikipedia.org/wiki/Qondvana
Qondvana – Azerbaijani
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https://bar.wikipedia.org/wiki/Gondwana
Gondwana – Bavarian
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https://bs.wikipedia.org/wiki/Gondvana
Gondvana – Bosnian
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Gondwana – Catalan
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Gondwana – Czech
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Gondwana – Welsh
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Gondwana – German
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Gondwana – Estonian
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Gondwana – Spanish
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Gondvano – Esperanto
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Gondwana – Basque
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Gondwana – French
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Gondwana – Scottish Gaelic
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Gondwana – Galician
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Gondvana – Croatian
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Gondwana – Indonesian
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Gondvana – Icelandic
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Gondwana – Italian
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Gondwana – Malagasy
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Gondwana – Norwegian
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Gondwana – Norwegian Nynorsk
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Gondwana – Occitan
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Gondvana – Uzbek
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Gondwana – Piedmontese
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Gondwana – Polish
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Gondwana – Portuguese
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Gondwana – Romanian
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Gondwana – Scots
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Gondwana – Slovak
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Gondvana – Slovenian
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Gondvana – Serbo-Croatian
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Gondwana – Finnish
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Gondwana – Swedish
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Gondwana – Tagalog
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Gondvana – Turkish
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Gondwana – Vietnamese
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Gondwana – West Flemish
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Gondwana – Zazaki
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