Biology:Savannah hypothesis

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The savannah hypothesis (or savanna hypothesis) is a hypothesis that human bipedalism evolved as a direct result of human ancestors' transition from an arboreal lifestyle to one on the savannas. According to the hypothesis, hominins left the woodlands that had previously been their natural habitat millions of years ago and adapted to their new habitat by walking upright.

The idea that a climate-driven retraction of tropical forests forced early hominini into bipedalism has been around for a long time, often implicitly. Some early authors saw savannahs as open grasslands, while others saw a mosaic of environments from woodlands to grasslands. The hypothesis has seen rising criticism since at least the late 1960s.[1]:98 The open grasslands version is mostly dismissed; in contrast, the mosaic version still has relatively wide support. However, the transition from forest to savanna probably was more gradual than previously thought.

History

The fundamental ideas behind it date back to Lamarck, Darwin and Wallace.[2][3][4] Also Gustav Steinmann saw reducing rain forest due to climate change as an important driver for bipedalism.[5] Osborn thought man probably originated from the forests and flood-plains of southern Asia.[6] Hilzheimer stated it was open landscapes that stimulated development.[7]

The hypothesis first came to prominence however with the discovery of Australopithecus africanus by Raymond Dart in 1924. In an article on the discovery, published in the journal Nature, Dart wrote:

"For the production of man a different apprenticeship was needed to sharpen the wits and quicken the higher manifestations of intellect – a more open veldt country where competition was keener between swiftness and stealth, and where adroitness of thinking and movement played a preponderating role in the preservation of the species. Darwin has said, "no country in the world abounds in a greater degree with dangerous beasts than Southern Africa." and, in my opinion, Southern Africa, by providing a vast open country with occasional wooded belts and a relative scarcity of water, together with a fierce and bitter mammalian competition, furnished a laboratory such as was essential to this penultimate phase of human evolution."
—Raymond Dart, Australopithecus africanus: The Man-Ape of South Africa [8]

Weinert stated apes are very reluctant to leave the safety of the trees, and the ancestors of modern man did not leave the trees, but the trees left them.[9] Grabau echoed this by saying Instead of the apes leaving the trees, the trees left the apes.[10]

Not everyone agreed with this hypothesis, such as Weidenreich, but he did conclude it was a widely spread belief.[11]

The work of Robert Ardrey helped popularize the ideas that Dart had developed with a wide audience.

In the decades following Dart's discovery, more hominid fossils were found in Eastern and Southern Africa, leading researches to conclude that these were savanna dwellers as well. Much of the academic discussion at the time took for granted that the transition to the savannas was responsible for the emergence of bipedalism, and focused instead on determining particular mechanisms by which this happened.[12]

One of the proposed mechanisms was the knuckle-walking hypothesis; the claim that early human ancestors walked on all fours when they first emerged into the savannas. This was based on observations of morphological characteristics found in Australopithecus anamensis and Australopithecus afarensis, and posited that knuckle-walking was an example of convergent evolution in chimpanzees and gorillas, which was then lost by the genus Homo.[13] Paleoanthropologists also posited that the upright posture would have been advantageous to savanna-dwelling hominids, as it allowed them to peer over tall grasses for predators, or in search of prey.[14] P. E. Wheeler suggested that another advantage lay in reducing the amount of skin exposed to the sun, which helped regulate body temperatures.[15] The turnover-pulse hypothesis, first described by Elizabeth Vrba seemed to support the savanna hypothesis by suggesting that climate change events resulting in the shrinking of forested areas forced animals out into the open grasslands.[12]

Robinson investigated adaptive radiation for Australopithecus and saw grass savanna and other more arid environments expanding at this time, thus providing increased opportunity for animals capable of adapting to such conditions.[16] Monod investigated the role in human evolution of the Sahara during wet periods as a place that was covered with steppes, savannas, and lakes. He saw advantages for the process of hominization in a wooded savanna.[17]

In analogy with gelada Jolly proposed that [i]n the basal hominid, therefore, the 'gelada' specialisations would be superimposed upon a behavioural repertoire and post-cranial structure already attuned to some degree of truncal erectness. The transition to bipedality would have been instigated by seed-eating and would probably took place in a dambo-like environment, later shifting to wider floodplains.[18]

An early critic of the savanna hypothesis was Lovejoy in 1981. He stated [i]t is more likely that hominids venturing into open habitats were already bipedal and that their regular occupation of savannahs was not possible until intensified social behavior was well developed.[19]

Kortlandt sought the barrier required for geographic speciation to take place. According to him, the Great Rift Valley, the Nile and the Zambezi acted as a double barrier when a period of desiccation set in East Africa. This must have converted the last-surviving dryopithecine (Proconsul) ape there into an upright-walking, drought-adapted, and "humanoid" type of bush and grassland ape, i.e., in all probability the Homininae, strictly speaking.[20] This corresponded with the location of some important fossils that had been found until then, such as in 1939 the Australopithecus afarensis in Laetoli by Ludwig Kohl-Larsen and the Paranthropus boisei in the Olduvai Gorge in 1959 by Mary Leakey. This Rift Valley theory became known as the East Side Story by Yves Coppens.[21]

Shifting consensus

In the latter parts of the 20th century, new fossil evidence began to emerge which called the savanna hypothesis into question. These newly-discovered remains showed indications that they were still well adapted to climbing trees, even after they had begun to walk upright.[22] Both humans and chimpanzees tend to walk upright when moving along long branches of trees, increasing their reach.[23]

In 1993, 4.4 million year old fossil teeth were found in Aramis, Ethiopia, by a group led by Tim D. White attributed to a new species, Australopithecus ramidus, later called Ardipithecus ramidus. The age was thus half a million years older than previously known A. afarensis and had a more monkey-like appearance.[24] After extensive research, in 2009 in a series of eleven articles in Science, more was published about Ardi. It concluded that Ar. ramidus preferred more wooded areas instead of open grassland, which would not support the climate-driven savannah hypothesis.[25]

A year later, these conclusions were questioned: In contrast, we find the environmental context of Ar. ramidus at Aramis to be represented by what is commonly referred to as "tree or bush savanna" with 25% or less woody canopy cover. The habitats involved probably ranged from riparian forest to grassland.[26]

For Phillip Tobias, the 1994 find of Little Foot, the collection of Australopithecus africanus foot bones demonstrating features consistent with tree-climbing as well as an upright gait, contributed to calling the savannah hypothesis obsolete, stating Open the window and throw out the savannah hypothesis; it's dead and we need a new paradigm.[27]

In 2000 Brigitte Senut and Martin Pickford found the 6 million year old Orrorin tugenensis in Kenya. The skeleton seems to indicate both bipedalism and good climbing skills. The latter indicates a wooded environment, as does the discovery of black-and-white colobuses. The discovery of impalas points more towards a more open landscape.[28] It later led Senut to the conclusion that the savannah hypothesis was no longer tenable.[29] If these fossils are indeed early ancestors of modern man, then the environment of the later Australopithecus is less relevant.

In 2001 the 7 million year old Sahelanthropus tchadensis was discovered in Chad. Based on animal finds in the vicinity, this suggests a mosaic of environments from gallery forest at the edge of a lake area to a dominance of large savannah and grassland, although more research was needed to determine this precisely.[30] The 5.6 million year old Ardipithecus kadabba discovered in 1997 was found in a similar terrain.[31]

Definition of savannah

Not everyone was willing to write off the savannah hypothesis. A poor definition of what a savannah actually is contributed to this. Critics of the hypothesis often saw the savannah as open grasslands with sporadic tree growth. However, savannas can have a high tree density and can also be humid. The big difference between savannas and forests is the lack of grasses in the latter. Thure E. Cerling developed a method to determine the forest cover of ancient landscapes, thus no longer requiring a definition of what a savannah is. By distinguishing between the C3 plants of the tropical forests and the mix of trees and C4 grasses of the savannah, they investigated the stable carbon isotope of paleosols from some sites in East Africa. They described landscapes varying from forest, woodland/bushland/shrubland, wooded grasslands to grasslands. They concluded that the early hominini lived in a more open environment than Australopithecus, rendering the savannah hypothesis still a plausible possibility.[32]

Following on from Cerling, Manuel Domínguez-Rodrigo stated that the usual division of landscapes into grassy, wooded and wooded is of little use, because it tells nothing about the evolutionary pressure on mammals. For example, the selection pressure of grass fields in tropical forests is incomparable to the grasslands of savannas. Tropical forests also have many different species of trees, while savannas only have a few species, which hardly carry any fruit. Another factor is that of scale. Paleontologists often only investigate the site itself, an area of several hundred to thousands of meters. These habitats are referred to as biomes, yet, this latter term includes many hundreds of kilometres. According to Domínguez-Rodrigo, the savannah hypothesis can still give a good explanation, although the transition of environment has probably been less abrupt than some earlier authors thought.[33]

See also

References

  1. ↑ Richmond, G. B., Begun, R. D., and Strait, D. S. (2001): 'Origin of Human Bipedalism: The Knuckle-Walking Hypothesis Revisited' in Yearbook of Physical Anthropology
  2. ↑ Lamarck, J.B. de (1809): Philosophie zoologique, ou Exposition des considérations relative à l'histoire naturelle des animaux, Dentu
  3. ↑ Darwin, C.R. (1871): The Descent of Man, and Selection in Relation to Sex, John Murray
  4. ↑ Wallace, A.R. (1889): Darwinism. An Exposition of the Theory of Natural Selection with Some of Its Applications, Macmillan and Co.
  5. ↑ Steinmann, G. (1908): Die geologischen Grundlagen der Abstammungslehre, W. Engelmann
  6. ↑ Osborn, H.F. (1915): Men of the Old Stone Age. Their Environment, Life and Art, Charles Scribner's Sons
  7. ↑ Hilzheimer, O.J.M. (1921): 'Aphoristische Gedanken über einen Zusammenhang zwischen Erdgeschichte, Biologie, Menschheitsgeschichte und Kulturgeschichte' in Zeitschrift für Morphologie und Anthropologie, 21, p. 185-208
  8. ↑ Dart, Raymond (7 February 1925). "Australopithecus africanus: The Man-Ape of South Africa". Nature 115 (2884): 195–199. doi:10.1038/115195a0. Bibcode: 1925Natur.115..195D. http://www.nature.com/nature/ancestor/pdf/115195.pdf. Retrieved 26 September 2017. 
  9. ↑ Weinert, H. (1932): Ursprung der Menschheit. Über den engeren Anschluss des Menschengeschlechts an die Menschenaffen, Ferdinand Enke
  10. ↑ Grabau, A.W. (1961): The World We Live in. A New Interpretation of Earth History, Geological Society of China
  11. ↑ Weidenreich, F. (1939): 'Six Lectures on Sinanthropus Pekinensis and Related Problems' in Bulletin of the Geological Society of China, Volume 19, p. 1-92
  12. ↑ 12.0 12.1 Shreeve, James (1 July 1996). "Sunset on the Savanna". Discover 17 (7). http://www.cabrillo.edu/~crsmith/Sunset.Savanna.doc. Retrieved 26 September 2017. 
  13. ↑ Richmond, Brian; Strait, David (23 March 2000). "Evidence that humans evolved from a knuckle-walking ancestor". Nature 404 (6776): 382–385. doi:10.1038/35006045. PMID 10746723. Bibcode: 2000Natur.404..382R. 
  14. ↑ Falk, Dean (22 February 2000). Primate Diversity (2 ed.). New York: W. W. Norton & Company. ISBN 978-0393974287. http://books.wwnorton.com/books/webad.aspx?id=10762. Retrieved 26 September 2017. 
  15. ↑ Wheeler, P. E. (1 January 1984). "The evolution of bipedality and loss of functional body hair in hominids". Journal of Human Evolution 13 (1): 91–98. doi:10.1016/S0047-2484(84)80079-2. 
  16. ↑ Robinson J.T. (1963): 'Adaptive radiation in the Australopithecines and the origin of man' in Howell, F.C.; Bourlière, F. African Ecology and Human Evolution, Aldine, p. 385-416
  17. ↑ Monod, T. (1963): 'Late Tertiary and Pleistocene in the Sahara' in Howell; Bourlière, p. 117–229
  18. ↑ Jolly, C.J. (1970): 'The Seed-Eaters: A New Model of Hominid Differentiation Based on a Baboon Analogy' in Man, Volume 5, p. 5–26
  19. ↑ Lovejoy, C.O. (1981): 'The Origin of Man' in Science, Volume 211, Number 4480, p. 341-350
  20. ↑ Kortlandt, A. (1972): New Perspectives on Ape and Human Evolution, Stichting voor Psychobiologie
  21. ↑ Coppens, Y. (1994): 'East Side Story: The Origin of Humankind' in Scientific American, Volume 270, no. 5, p. 88-95
  22. ↑ Green, Alemseged, David, Zeresenay (2017). "Australopithecus afarensis Scapular Ontogeny, Function, and the Role of Climbing in Human Evolution". Science 338 (6106): 514–517. doi:10.1126/science.1227123. PMID 23112331. 
  23. ↑ Thorpe, S. K.; Holder, R.L; Crompton, R. H. (2007). "Origin of human bipedalism as an adaptation for locomotion on flexible branches". Science 316 (5829): 1328–31. doi:10.1126/science.1140799. PMID 17540902. Bibcode: 2007Sci...316.1328T. 
  24. ↑ White, T.D.; Suwa, G.; Asfaw, B. (1994): 'Australopithecus ramidus, a new species of early hominid from Aramis, Ethiopia' in Nature, Volume 371, p. 306–312
  25. ↑ White, T.D.; Asfaw, B.; Beyene, Y.; Haile-Selassie, Y.; Lovejoy, C.O.; Suwa, G.; WoldeGabriel, G. (2009): 'Ardipithecus ramidus and the Paleobiology of Early Hominids' in Science, Volume 326, p. 75-86
  26. ↑ Cerling, T.E.; Levin, N.E.; Quade, J.; Wynn, J.G.; Fox, D.L.; Kingston, J.D.; Klein, R.G.; Brown, F.H. (2010): 'Comment on the Paleoenvironment of Ardipithecus ramidus' in Science, Volume 328, 1105
  27. ↑ Vaneechoutte, M.; Kuliukas, A.; Verhaegen, M. (2011): Was Man More Aquatic in the Past? Fifty Years After Alister Hardy – Waterside Hypotheses of Human Evolution, Bentham Science Publishers
  28. ↑ Pickford, M.; Senut, B. (2001): 'The geological and faunal context of Late Miocene hominid remains from Lukeino, Kenya' in Comptes Rendus de l’Academie des Sciences, Series IIA, Earth and Planetary Science, Volume 332, No. 2, p. 145-152
  29. ↑ Senut, B. (2015): 'Morphology and environment in some fossil Hominoids and Pedetids (Mammalia)' in Journal of Anatomy, Volume 228, Issue 4
  30. ↑ Vignaud, P. et al. (2002): 'Geology and palaeontology of the Upper Miocene Toros-Menalla hominid locality, Chad' in Nature, Volume 418, p. 152-155
  31. ↑ Su, D.F.; Ambrose, S.H.; DeGusta, D.; Haile-Selassie, Y. (2009): 'Paleoenvironment' in Haile-Selassie, Y.; WoldeGabriel, G. Ardipithecus Kadabba. Late Miocene Evidence from the Middle Awash, Ethiopia, University of California Press
  32. ↑ Cerling, T.E.; Wynn, J.G.; Andanje, S.A.; Bird, M.I.; Korir, D.K.; Levin, N.E.; Mace, W.; Macharia, A.N.; Quade, J.; Remien, C.H. (2011): 'Woody cover and hominin environments in the past 6 million years' in Nature, Volume 476, p. 51-56
  33. ↑ Domínguez-Rodrigo, M. (2014): 'Is the “Savanna Hypothesis” a Dead Concept for Explaining the Emergence of the Earliest Hominins?' in Current Anthropology, Volume 55, Number 1, p. 59-81



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