Aquatic ape hypothesis

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The Aquatic Ape Hypothesis (sometimes loosely called the Aquatic Ape Theory) proposes that ancestors of modern humans went through one or more periods of time living in a semi-aquatic setting and that this accounts for many of the characteristics of modern man. This is a minority position not widely held in biology. The common and conventional view of human evolution is that the first hominids evolved in the savanna environments (the Savanna Theory).

Proponents of the Aquatic Ape Hypothesis believe that the ancestors of modern humans went through one or more periods of time living in a semi-aquatic setting, that they gathered most of their food from shallow coastal waters before their descendants returned to a more land-based existence, and that adaptations to this marine environment can be clearly identified in the modern human phenotype. There are interpretations which propose fresh-water habitats (e.g. Ellis 1993), variations in the timescale (e.g. Verhaegen et al 2002) and the proposed degree of selection arising from moving through water.

The hypothesis was originally suggested in 1942, by Max Westenhőffer in The Road to Man (Der Eigenweg des Menschen), although the author referred only briefly to it, and apparently considered that it was not a new idea. It became more well-known in 1960 when proposed in academic circles by the marine biologist Sir Alister Hardy. Hardy had had the idea privately since about 1930, independently of Westenhőffer. The early television playwright and, later, feminist writer Elaine Morgan developed and promoted it, publishing her first book on the subject, The Descent of Woman (, in 1972. Her later books on the subject are: The Aquatic Ape ( (1982), The Scars of Evolution ( (1990), The Descent of the Child ( (1994) and The Aquatic Ape Hypothesis ( (1997).



The aquatic ape hypothesis puts forward several main arguments. It should be noted that some of the facts in these arguments are in dispute and thus should be viewed with a degree of educated skepticism.

  • Bipedalism: Humans are among very few bipedal mammals, none of which adopt the fully-upright, full-time human posture. Gorillas, chimpanzees and bears are able to walk on two legs when they have a particular reason, but always revert to quadrupedalism as their basic means of locomotion. Some lemurs skip sideways on two legs when on the ground, because their adaptations to leaping through trees make ground-based quadrupedalism difficult. And of course kangaroos and similar creatures use a bipedal form of locomotion that is quite different from human walking. Even birds, with a few exceptions such as penguins, walk bipedally but with their bodies held horizontally, unlike the human upright posture. Creatures such as squirrels and meerkats often adopt an upright posture when stationary, but do not walk or run bipedally.
Although it gives us the ability to use tools whilst walking or running, bipedalism and upright posture come at a significant cost, with side effects including back problems, knee problems, varicose veins, hemorrhoids, hernias and problems with childbirth.
Since evolution works in small steps, it is hard to see how bipedalism could have evolved on the savanna: the mass of the torso makes it inherently unstable and inefficient for locomotion. Bipedalism is not observed in other savanna mammals.
Water, however, supports the body, and proboscis monkeys have been observed wading bipedally in their occasionally flooded habitats. The one other animal known to have a pelvis adapted to bipedal walking was prehistoric Oreopithecus, commonly known as the Swamp Ape owing to its flooded habitat.
  • Breathing: With the exception of humans and a few other outstanding animals (many of which had aquatic ancestry like the elephant), land mammals have no conscious control over their breathing. The voluntary control humans have over their respiratory system is similar to that of aquatic mammals which inhale as much air as they need for a dive, then return to the surface for air. Human babies also reflexively hold their breaths both when submerged, as well as swimming without training. Some young children also have the ability to completely close their nostrils at will like an aquatic mammal, which then explains the sinus cavity as a means of equalizing pressure. And humans have a descended larynx, otherwise found only in sea otters and dugongs, allowing them to quickly breathe through their mouths, a trait so useful for semi-aquatic activity that the only birds which breathe through their mouths are diving birds.
  • Fat: Humans have ten times as many fat cells in our bodies as is normal in an animal of our size. We are by far the fattest primate, and have many adipose cells even when we are considered slim by human standards. Mammals which hibernate have seasonal fat: aquatic mammals, like humans, retain fat throughout the year. Human infants are especially fat compared to apes and most other fully terrestrial mammals. The human fatty layer is also attached to the skin, like most aquatic mammals, not the muscle, like almost all land mammals. Humans also lack the layer of cutaneous muscle, present in every land mammal (including all other primates), that allows many land animals to twitch their skin, but is absent in aquatic mammals.
  • Childbirth: Dramatic increase in the size of the cranium is a prominent theme in human evolution, making childbirth difficult and dangerous. Water birthing is known to facilitate childbirth and to reduce risks to mother and infant. Human infants are born covered in vernix caseosa, a waterproof coating, and continue to draw oxygen through the umbilical cord while underwater.
  • Nutrition: Human brain tissue requires comparatively large amounts of omega-3 fatty acids, which are uncommon in the land food chain but prevalent in the marine food chain. Indeed, most animals which move to plains life tend to develop smaller brains, while aquatic animals tend to evolve larger ones, quite possibly because of access to Omega 3.
  • Tears and excessive sweating, prevalent in humans but not in other primates, are considered further evidence to support the hypothesis, insofar as they are vectors for the removal of excess water and salts from the body, as might result from the ingestion of saltwater (as in eating food from a salt marsh). Other marine animals, such as the elephant, cry saline tears, and the mechanism by which humans produce sweat from eccrine glands could have developed as a means of sheding extra salt.
  • Reproductive Traits: The most common human mating practice, missionary position, is essentially front-to-front, exactly how aquatic mammals must mate. No other land animals, save the bonobo chimpanzee, use such a position, instead mating coitus more ferarum, as with dogs is the norm. Marine animals, even non-mammals, also tend to develop a less accessible vagina to keep out water, necessitating larger male reproductive organs, a trait long noted as specific to humans and bonobos (who live partially in flooded forest) among primates.

One difficulty in evaluating this hypothesis is that the places it suggests fossils might be found are mostly below sea level at the present epoch. Furthermore, swamps and marshes are inimical to the creation of fossils.

Comparison with land-based hypotheses

  • Nakedness: The usual land-based explanation (the "thermoregulatory hypothesis") is that it was for cooling - humans sweat more per unit surface area than any other mammal, and proponents of this idea claim that it makes us particularly effective at remaining active during the heat of the day. A layer of hair would supposedly reduce the effectiveness of this (human sweat may be seen as an analogue of the water-seeking behaviors of the animals mentioned above).
  • Problems with this explanation are that body hair is needed to protect against direct sun and extreme heat as well as cold; that human sweating is highly wasteful of water and salts, which is a distinct disadvantage on the savanna; and that exposed skin is not, after all, essential for sweating to be effective (hair creates much more surface area for evaporation than skin). A prime example of this is the horse, which does sweat when hot, and yet is covered in hair. Indeed, most savanna animals have hair in part because it provides protection to skin from the heat and ultraviolet radiation of direct sunlight, not as important to semi-aquatic animals which are cooled and sheltered by water.
In addition, any such hypothesis has to explain the pattern of hair that we do have, and why women and children have less body hair than men. On the first point, why should we have retained head hair if the purpose of a naked skin is to keep cool? On the side of AAH, it may be noted that the top and the back of the head are the areas least in contact with water in the human pattern of swimming, and also the only areas covered with thick hair in both mature individuals and infants.
On the second point, it is possible to suggest an AAH scenario in which mature males spent more time near the shore, while mothers with babies stayed in deeper water out of reach of land predators. By contrast, it is hard for the temperature regulation hypothesis to accommodate a case where females and infants were more active than males, and therefore more in need of sweat-cooling, in the heat of the day.
  • Bipedalism: There are over a dozen land-based suggestions as to why the first hominids became bipedal: carrying behaviour, tool-making, and sentry behaviour, for example.
  • The difficulty with all of these is that (unlike the putative Aquatic Ape, which could have waded much more frequently) none of them apply for more than a small amount of the time; when not engaged in these behaviours, the proto-hominids would simply have reverted to quadrupedalism. In waist deep water, apes have little choice but to move bipedally and do so, very predictably. This is unusual for mammals which typically continue to wade quadrupedally, or switch to swimming.
  • Fat is very important in developing and maintaining the brain, which is a very expensive organ in terms of energy requirements.
  • However, this suggestion doesn't account for the fact that women and babies have a much higher proportion of body fat than men; while within the AAH scenario this, as with the contrast in body hair, further suggests that nursing mothers would have spent more time in water than adult males.

Objections to AAH

  • Nakedness: Human hair is drastically different from all of the aquatic species named above. The comparison to fully aquatic mammals (cetaceans, sirenians, etc.) is suspect, as these animals have evolved characteristics over a far longer period than humans. Further, many proponents of AAH claim that the putative aquatic ancestor was never that aquatic, thus presenting an internal inconsistency in their arguments when a feature of dedicated marine mammals appears without similar selective stimulus.
  • Bipedalism: No aquatic mammal is bipedal. Those animals that are temporarily bipedal (such as kangaroos and some primates) use their upright state for locomotion, feeding and sentry behaviour, which are all useful for terrestrial life. Furthermore, for standing in shallow water, it is useful to have lower limbs substantially longer than upper limbs, as is common in wading birds. Human legs do not fit this pattern (although the ratio of leg length to arm length is indeed higher in humans than in other primates).
  • Breathing: The ability to moderate breathing, to a lesser degree of control, is seen in many other animals, including other apes and dogs. The mammalian diving reflex exists in other mammals as well.


AAH provokes fierce and often acrimonious contention. Sceptics criticise the lack of direct fossil evidence; the sometimes amateurish way in which it is presented; and the occasional over-emphasis of tenuous arguments. Proponents complain about a dismissive and superior attitude; attacks on methods and personalities rather than substance; an exaggeration of the degree of aquaticism being proposed; and the failure to provide land-based alternative hypotheses that survive the very criticisms levelled at AAH.


  • Ellis, D. V., "Wetlands or Aquatic Ape? Availability of food resources." Nutrition and Health, 9, 205-217 (1993).
  • Hardy, A. C., "Was man more aquatic in the past?", New Scientist, 7,642-645 (1960).
  • Morgan, Elaine, The Aquatic Ape, 1982, Stein & Day Pub, ISBN 0-285-62509-8
  • -- The Scars of Evolution, 1990, Souvenir Press, ISBN 0-285-62996-4
  • -- The Aquatic Ape Hypothesis, 1997, Souvenir Press, ISBN 0-285-63377-5
  • Verhaegen, M., Puech, P-F, Munro, S., "Aquarboreal Ancestors?" Trends in Ecology and Evolution, 17, 212-217 (2002).

See also

External links




es:Teoría del Simio Acuático ja:アクア説


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