AAH Refuted, or just misunderstood?
The So-called “Aquatic Ape Hypothesis”:
Refuted, or just misunderstood?
Algis Kuliukas (UWA) akuliukas@anhb.uwa.edu.au
Talk given at ANU, at 10:30 am, 9th December 2004
Abstract
Since Hardy (1960) published a request for comments about his controversial idea
that ‘man was more aquatic in the past’, the total response has remained rather muted,
if generally dismissive, from paleoanthropologists. Students of the discipline today
could be forgiven for concluding that the idea had been decisively refuted by the
scientific method, that perhaps some unequivocal evidence had been published in the
literature which dismissed the predictions made by the hypothesis.
It is argued here, that no such decisive refutation has, in fact, been forthcoming. The
two works in the literature which have set out to do so, Langdon (1997) and parts of
Roede et al (1991), both seem to have critiqued rather exaggerated versions of what
Hardy appears to have originally postulated.
In this talk the case is made that, rather than exaggerating its claims in this way, it is
possible to scale them back, in several ways, and that if one does so, the hypothesis
only gains in credibility.
It is proposed, in fact, that this hypothesis has yet to even be clearly defined (thus
removing the large scope for personal interpretation), let alone refuted. A starting
point – a simple and unambiguous definition - therefore, is suggested from which the
hypothesis can be tested and scientific rigour applied in the future.
Paper Presentation – Full Transcript of the talk follows:
Algis Kuliukas
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AAH Refuted, or just misunderstood?
The soso-called “Aquatic Ape Hypothesis”
Hypothesis”:
Refuted... or just misunderstood?
Algis Kuliukas MSc
University of Western Australia
Supervisors:
Dr Nick Milne (Anatomy & Human Biology) &
Dr Paul Fournier (Human Movement Science)
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Images: from left to right, top to bottom: Alister Hardy FRS, Elaine Morgan; Image of infant floating, unaided, face-up from
Desmond Morris’ book: Manwatching; An image of ‘paradise’; Thin strip of gallery forest – dry climates cause forests to shrink
close to water sources; chimp wading up to shoulders in Conkuoati, Congo; Paleogeography of Miocene Europe – an
archipelago of islands subject to changing coastlines.
Introduction
Good afternoon. My name is Algis Kuliukas and I’m a PhD student at UWA looking
at hominid bipedal origins and, specifically, the hypothesis that wading through
shallow water was a significant factor in its evolution.
This subject, some may have noticed, is closely associated with a certain, apparently
controversial, idea on human evolution: the so-called ‘aquatic ape theory.’
I should say from the start that I have never understood this controversy. In fact it is
through trying to understand it that has driven me to the point where I turned my back
on a lucrative IT career to return to academia. Since doing so, whilst being a Masters
student at UCL, in London and a PhD student here in Australia, several eminent
people have advised me not to mention the ‘aquatic ape hypothesis’ by name when
writing theses, PhD proposals and whilst giving talks such as this. “It might be
misunderstood”, I was told. I have carefully followed this advice until today. But
when David Cameron asked me to come here today to give a talk specifically about
the “Aquatic Ape Hypothesis” at this year’s ASHB, I realised that it would be
impossible to heed that advice any longer. I decided that this would, in fact, provide
the ideal opportunity to finally put this taboo to the sword, to lift up the carpet and
pull out from under it this thing that seems to have been brushed there for over forty
years, to pick it up off the floor, dust it down, rather like one might an old fossil. To
put it on the table and get people to gather round and examine it, perhaps more
carefully than they have done before. That is what I intend to do today.
Before I start I’d like to get a feel for the level of awareness and support for this idea
in the audience.
How many people have heard of the aquatic ape theory?
[Almost everyone puts their hand up]
Has anyone not heard of it?
[Just a small number raise their hands]
Ok.
Algis Kuliukas
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AAH Refuted, or just misunderstood?
Are you a proponent of the AAH?
NO
YES
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Now, who would regard themselves as a proponent of the hypothesis? On a scale
from Blue to Green (I didn’t want to use a numeric scale for fear of being accused of
bias), where ‘Blue’ means you think Elaine Morgan is wonderful and that all the main
differences between humans and chimps can be explained by moving through water;
‘Green’ means you thinks it’s just rubbish, pseudoscience and we really shouldn’t be
discussing it at a forum such as this; and ‘Turquoise’ means you’re rather agnostic –
who would rate themselves a blue shade of turquoise?
[about two hands out of about 60 go up]
No surprise there!
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AAH Refuted, or just misunderstood?
But can you cite a modern and
unambiguous definition in the literature?
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I have a final, rhetorical question for you: ... So, can you tell me what it is?
Now, by this, I don’t mean what you think it is, or what your impression of it is. I
mean: Can you cite a piece from the scientific literature where you remember reading
a paragraph, or even a couple of sentences, where the idea was clearly,
unambiguously defined in a context that still makes sense according to the fossil
record we know today?
I say it’s a rhetorical question, because I really don’t think such a definition exists.
[one person puts hand up, as if to dispute what I just said]
Now, I expect some people are maybe thinking of Alister Hardy’s 1960 paper or
perhaps one of Elaine Morgan’s five books but I would suggest that even those do not
really contain a definition that is clear, unambiguous and relatively up to date in terms
of recent fossil evidence and other findings.
[Hand goes down]
Hardy’s original article certainly included a description of what he was thinking
about. But it was only a request for comments, not a carefully worked out hypothesis.
And it certainly contained ambiguities: On the one hand, he alluded to fully aquatic
mammals, like whales and dolphins as analogues for human subcutaneous fat, but on
the other hand said that he never envisaged out ancestors being even as aquatic as an
otter or spending more than a few hours in water every day. And, of course, since his
paper there have been many fossil discoveries which would surely force him to alter
his position, if he weer alive today.
Elaine Morgan wrote five books on this subject. Surely, she must have defined it
carefully somewhere in one of those. Apparently not. I wrote to her about this and she
told me that she deliberately avoided tying any particular flag to the AAH mast for
fear of it being attacked. Her aim was always, only to promote Hardy’s idea and ask:
Why has no-one looked at this thing? You might not consider this to be very scientific
but the fact is, today we remain in a situation where everyone has heard of the
Aquatic Ape Hypothesis, very few support it but no-one can tell you what it is!
Now I don’t know about you, but I don’t think that’s a very sound basis for a
refutation.
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AAH Refuted, or just misunderstood?
Agenda
1. Which ‘AAH’?
2. What has been the response to it in academia?
Sparse.
Majority open.
Minority rejecting – but also exaggerating.
3. How could it have been interpreted differently?
4. It’s time for an unambiguous definition.
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Now, I’ve already taken up about five minutes just on the intro, so I need to accelerate
a little. It’s a very big subject and I have very little time, so I’m just going to make a
few basic points.
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AAH Refuted, or just misunderstood?
1. Which ‘Aquatic Ape Hypothesis?’
Hypothesis?’








Westenhöfer (1942)
Hardy (1960)
Morgan (1972 - 1997)
Ellis (1984)
Williams (1997)
Verhaegen, Munro et al (2000)
Crawford et al (2000)
Others.
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Images: Max Westenhöfer; Alister Hardy FRS; Elaine Morgan; Marc Verhaegen.
The first point is this: There is not one aquatic ape hypothesis. Just as there is not one
‘orthodox’ idea, there are a variety of different models which favour some kind of
aquatic theme. There is not time to go through all of these in detail here but suffice it
to say that they vary in their proposed timescales, the type of habitat and the kind of
traits suggested as evidence for it.
I want to mention one name on this slide specifically, however, and that is Stephen
Munro, who is a student here at ANU. He has co-authored several papers with Marc
Verhaegen which promote, in my view, a very plausible model of human evolution.
The point is that the idea is not a simple monolith which, of course, makes it all the
more difficult to refute.
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AAH Refuted, or just misunderstood?
2. What has been the academic
response?



Sparse

A trickle of responses in 1960.

Few (if any) Ph.D. students assigned to it.
Majority, open: (Excepting that of proponents ~ at least 40 pieces)

Sauer (1960) – Coastal foraging.

Knight (1991) – Menstruation and ‘sex strike theory’.

Evans (1992) – Adaptive explanation for the paranasal sinuses.

Groves (1993) – Roede et al. Book Review.

Tobias (1995) – Water and Human Evolution – 3 papers.

Wescott (1995) – Aquaticism and Quantalism.

Ashford (2000) – Water-borne parasites indicators of human evolution.

Stringer (2000) - “What about wading?”

Attenborough (2002) – Wading, in ‘Life of Mammals.

Niemitz (2002) – “Amphibisce Generalistheorie”

Cameron & Groves (2004) – ‘...we insist that the AAH take its place...’
Minority, rejecting:

Lowenstein & Zihlman (1980)

Langdon (1997)

Roede et al (1991) – Half in favour, half against.
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Now, what has been the academic response to this idea? It can be summed up in just
five words: ‘Sparse, majority open, minority rejecting.’
First of all, ‘sparse’. In the weeks that followed Hardy’s original New Scientist article
there were a smattering of replies (mostly supportive) but then nothing happenned for
twelve years until Elaine Morgan wrote her first book. (Actually that itself was
inspired by Desmond Morris’ book ‘The Naked Ape’ specifically referring to the idea
in 1967.) I do not know of a single PhD student who has been encouraged to study
this thing. Clearly it would not have been a good career move. On the contrary most
students, I suspect, have been gently persuaded not to waste their time on it.
Now, considering how the hypothesis would appear to have been rejected by the field,
there is a surprising amount in the literature which is supportive or merely open to it.
Even if we take away all the papers and books written by AAH proponents, people
such as Hardy, Morgan, Verhaegen and Crawford – and that’s about forty pieces of
literature – and only take those written by people who would be regarded as
somewhat neutral, one finds that there are still over a dozen pieces that are open to the
idea. People like Phillip Tobias has openly called on his peers to look at this thing.
Chris Stringer, David Attenborough and the German Anthropologist Carsten Nimietz
have all argued in favour of the wading hypothesis for bipedal origins. And here in
Australia, Colin Groves and David Cameron have, on at least two occasions to my
knowledge, published statements which have urged people not to be so dismissive of
the hypothesis.
The literature against the hypothesis, on the other hand, only numbers at most three
pieces, and yet it is the ‘rejection’ which somehow has won primacy in this debate.
The three pieces of literature against are ‘Lowenstein and Zihlman’, Langdon and
Roede et al. Let’s take a close look at those now:
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The ‘Refutations’:



Lowenstein & Zihlman (1980)
 Ridiculing. (e.g. compares a’pith limb size to flippers.)
Langdon’s (JHE 1997) Critique
 See my paper for a more thorough critique, but…
 Comparisons with fully aquatic mammals seem to exaggerate
Hardy’s claims.
 Parades 26 aquatic ‘traits’ equally and superficially.
 “Savannah hypothesis... a straw man” invented by Morgan.
 Doesn’t draw upon Roede et al (1991)
Roede et al (1991)
 Proceedings of the Valkenburg Symposium 1987.
 Balanced but polarised (11 for 11 against) debate.
 On balance, the verdict was against.
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Images of Langdon’s Paper and Roede et al ‘Aquatic Ape: Fact or Fiction?’
Lowenstein and Adrienne Zihlman published a critique of the aquatic ape hypothesis
in the journal ‘Oceans’ in 1980. I only just came across this paper. Basically, it is a
mickey-take of the idea, exaggerating it to breaking point and then ridiculing the
remains. For example they compare australopithecines limb sizes with those of seals
and dolphins and conclude that, as they show no sign of reduction this, somehow,
shows the AAH wanting. Now clearly that is an absurd misrepresentation of the
hypothesis. Nobody has ever suggested that humans were on the way to becoming as
aquatic as seals and dolphins. Or, if they were, on a scale from one to a hundred,
they’d made, say, notch number three.
The most respected critique of the aquatic ape hypothesis, or at least, the one
published in the most respected journal, was Langdon’s 1997 paper published in the
Journal of Human Evolution.
Now I’ve written a full critique of this paper outlining my detailed criticisms and I’ve
put it in a PDF on the server here if anyone’s interested. But, just to summarise the
main points: Firstly, he makes the same kind of exaggerations that Lowenstein and
Zihlman make, comparing human ancestors to true aquatics. Secondly he parades 26
traits taken from one of Elaine Morgan’s books with equal superficiality. I put it to
you that this was just not fair. For example, in her latest book, Elaine Morgan wrote
four chapters on the wading idea, a whole chapter on nakedness and another on
subcutaneous fat. The remaining chapters discussed the other twenty three in ever
lighter and more speculative manner. To line them all up, as Langdon did, and imply
that they are all as important parts of the argument as each other is simply a straw
man. He did not do any justice to the most important argument, in my view: the
wading hypothesis. There, he simply took one (of many) of Elaine Morgan’s
arguments, misrepresented it and then argued for his own view of a brachiationist
model in just two paragraphs. This was typical of his treatment.
He used a straw man argument against Elaine Morgan but then had the fantastic
audacity to accuse her of doing so when he accused her of inventing the ‘savannah
theory’ merely as a straw man to be knocked down in favour of the aquatic theory.
Now I do not know if anyone has ever read a book on human evolution in the past 50
years where the savannah was not explicitly referred to, or at least the more open
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AAH Refuted, or just misunderstood?
habitats that it implies, but I certainly have not. It is surely just a grotesque distortion
to imply that this was all an invention of Elaine Morgan’s. Perhaps she was the first to
coin the phrase ‘savannah theory’ but, as she so eloquently puts it herself: “I would no
more have expected them to use that phrase that I would expect a Creationist to refer
to 'the God theory' - their faith in it was too strong for that.” Morgan (1997:14).
But perhaps the greatest indictment on Langdon’s critique is that it all but ignores the
one serious scientific study that has been published about it in the literature, Roede et
al. He does mention it, making his omission all the more alarming, but no more. I put
it to you that if an undergraduate student of yours handed you an essay on the aquatic
ape hypothesis that did not significantly draw upon the findings of Roede et al you
would put a red line through it and tell the student to do it again. And yet Langdon’s
paper was peer reviewed, was published in the hallowed pages of the Journal of
Human Evolution and remains, to this day, the great refutation, still without reply,
seven years on.
So, what about Roede et al? Well, as I said, it remains the single most serious piece of
scientific work on the subject, being the proceedings of a three-day symposium in
Valkenberg, Holland in 1987.
It was fair and balanced, but rather polarised, having eleven proponents and eleven
opponents, gathered to contest the issue, rather like two football teams.
It has to be said that the overall findings of the symposium were against...
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Team Against won, but only
The
Final? decision’
after aRoede
‘dodgy refereeing
Team Against
Team For
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... but, in football parlance, it was hardly a thrashing. In fact, some might say it was
more like a 1-0 win with a goal in the last minute of injury time following a dodgy
refereeing decision!
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The Roede et al (1991) Findings

Conclusion were overall against the hypothesis, but,
Vernon Reynolds, the chief editor, had this to say...
"Overall, it will be clear that I do not think it
would be correct to designate our early hominid
ancestors as ‘aquatic’. But at the same time
there does seem to be evidence that not only
did they take to the water from time to time but
that the water (and by this I mean inland lakes
and rivers) was a habitat that provided enough
extra food to count as an agency for selection."
Reynolds (1991:340)
Groves’ book review in 1993 came to the
same conclusion: If only taken this far, as
with Cliff Jolly’s seed eating hypothesis “How
© Algis Kuliukas 2004
stupid of one not to have thought of that oneself!"
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Let me explain what I mean. Vernon Reynolds was one of the four editors with the
unenviable task of trying to arrive at some kind of ‘verdict’ and a form of words to
summarise the whole symposium. He was, like Jan Wind, was one of the opponents
but was the more open of the two. The other two editors, Machteld Roede herself
(perhaps the book was named after her as some kind of concession), and John Patrick
were proponents. I would suggest that if you were looking to find out the thinking
behind the rejection of the aquatic ape hypothesis at this symposium, the best place to
look for it would be in Vernon Reynold’s chapters.
These are the words he used to do so...
"Overall, it will be clear that I do not think it would be correct to designate our early
hominid ancestors as ‘aquatic’. But at the same time there does seem to be evidence
that not only did they take to the water from time to time but that the water (and by
this I mean inland lakes and rivers) was a habitat that provided enough extra food to
count as an agency for selection." Reynolds (1991:340)
Note those words: “water ... was a habitat that provided enough extra food to count as
an agency for selection.”
Now when I read this, I must admit, I was confused. You see, I thought that was the
aquatic ape hypothesis! I never thought for a second that Hardy was actually
proposing a truly aquatic ape, in any sense of the phrase.
And it’s not just me. Colin Groves, in his book review of Roede et al makes
practically the same point. “If only Hardy and Morgan had gone this far” he wrote,
perhaps there would have been no problem. Perhaps we’d all have been kicking
ourselves for not thinking of it ourselves.
I must say that I agree with Colin exactly on this point.
Clearly, one’s impression of this hypothesis is a function of one’s interpretation of it.
The more aquatic you think it is proposing our ancestors to be, the more sceptical you
are likely to be, and vice versa. And, there is a lot of ground for interpretation....
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3. How could it have been
interpreted differently?


How ‘aquatic’ is ‘More aquatic?’ – All Hardy asked.
 Isn’t a generalist also, by definition, more aquatic?
Which wetland habitat?
 Coastal, islands, mangroves, marsh, swamp, gallery forests?
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Most obviously, there is the question of how aquatic is “more aquatic”? More aquatic
is a comparative term so it heavily implies that we need to compare our aquaticism
with something else. Clearly, if you choose to make comparisons with dolphins, or
even otters, humans are poor in water. However, if you compare us to chimpanzees,
you get a very different impression.
Most anthropologists hold the view that a big part of human evolution is a trend
towards being more generalist, as opposed to being adapted to any particular niche.
But if this is true, then isn’t a generalist, by definition, more aquatic than an
arborealist?
Then, there is the matter of habitat. Langdon and Roede et al rejected the AAH
largely on the idea of a distinct, early, coastal phase, but there are many habitats
where similar aquatic pressures could work.
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Orthodox ‘Linear’
Linear’ View of
Human Evolution
Savannah
Colder, Drier
Rain forest
Warmer, wetter
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Most orthodox texts on human evolution seem to espouse a rather simple, linear trend
explaining the process of hominization. Our ancestors lived in tropical rainforests
when the climate was wet and warm. Then, as the climate changed, and the habitat
became more sparsely populated with trees, our ancestors were forced to try to exploit
the rather more meagre resources of more open, savannah-like, habitats. They were
the products of a distinct trend towards greater aridity.
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Another dimension to consider:
Ma Savannah
Gallery Forest
River
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Adapted from Potts (1998)
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And yet there is another factor to be considered. Forests do not simply get more
sparse as the climate gets drier, they tend to cluster around lakes and rivers in gallery
forests. Therefore any fauna which have evolved in such forests are likely to cling
onto such refugias as long as they can, rather than venture out into new habitats to
which they are not yet adapted.
Also the climatic trend is clearly not so simple. There was a general shift to greater
aridity, but it was associated with ever increasing phases and frequencies from wet to
dry. This must have resulted in phases when forests shrunk back closer to the water’s
edge followed by other phases where those same gallery forests were prone to
flooding. This, paradoxically, must have placed our hominid ancestors living in the
Pleistocene ever closer to the water’s edge.
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3. How could it have been
interpreted differently?




How ‘aquatic’ is ‘More aquatic?’ – All Hardy asked.
 Isn’t a generalist also, by definition, more aquatic?
Which wetland habitat?
 Coastal, islands, mangroves, marsh, swamp, gallery forests?
Chronology:
 Pre- or post- Homo, Pre- or post– LCA?
 A distinct ‘aquatic’ phase, many cycles, or a constant trend?
 Can’t a ‘mosaic’ include ‘more aquatic’ pieces?
A hybridisation factor?
 Could Homo sapiens be the result of a introgression of two
hominid sub-species, one being more aquatic than the other?
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Another area open to interpretation is the chronology. When did this happen? The
Hardy-Morgan model posited a distinct ‘full-on’ aquatic phase somewhere between
ten million years ago and three million years ago. The time between the, as then seen,
fossil gap between Proconsul and Australopithecus. But clearly, this is not necessarily
the case. The ‘phase’ could have started at any time, even before the last common
ancestor of humans and chimps, and it could have ended only recently, perhaps as the
‘Out of Africa’ coastal and riverside migrations gave way to more in-land huntergathering.
Perhaps there was no ‘phase’, but rather a general adaptive trend towards living by the
water’s edge. And, as one of the most fashionable words to use in human evolution
today is ‘mosaic’, can’t this mosaic include ‘more aquatic’ bits?
Finally, my own pet theory is that Homo sapiens is a hybrid of two distinct hominid
taxa. This might best explain how we, uniquely among the Hominoidae, have just 46
chromosomes. If so, then we may have not one but two ancestral representatives in
the Plio-Pleistocene. If one of these were more aquatic and one less, then this might
result in a Homo sapiens species that betrayed an unusual mix of features which
appeared anomalous and hence more difficult to diagnose.
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4. A Proposed Definition for the AAH:
“The hypothesis that water has acted as an
agent of selection in the evolution of humans
more than it has in the evolution of our ape
cousins...”
“...And that, as a result, many of the major physical
differences between humans and the other apes may be
explained, to a large extent, as adaptations to moving
(wading, swimming and/or diving) better through various
aquatic media and from greater feeding on resources that
might be procured from such habitats.”
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I put it to you that there are lots of ways in which this hypothesis can be interpreted
differently. Therefore it is most important that we try to define it so that, at least, we
can agree about what it is we are discussing.
I have tried to do this. I’ve taken the lowest common denominator from all the
different aquatic ape theories and merged them into this overarching, simple
definition....
The hypothesis that water has acted as an agent of selection in the evolution of
humans more than it has in the evolution of our ape cousins...”
And just to add some detail...
“...And that, as a result, many of the major physical differences between humans
and the other apes may be explained, to a large extent, as adaptations to moving
(wading, swimming and/or diving) better through various aquatic media and from
greater feeding on resources that might be procured from such habitats.”
I believe that if we all agree to use this form of words, which was endorsed by both
Elaine Morgan and Marc Verhaegen, we might finally be able to apply some scientific
rigour to the idea to see if it might actually be useful.
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AAH Refuted, or just misunderstood?
Summary
1.
2.
3.
4.
5.
6.
AAH has been largely misunderstood, perhaps due
to ambiguities in Hardy’s original paper.
The academic response, although inadequate, has
been largely open to it.
Despite this, two or three refutations have won
primacy.
Two of the attempted refutations are very weak.
A mild and moderate definition has been defined
and endorsed by proponents.
Can we now mention the dreaded ‘a’ word, please?
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To summarise:
AAH has been largely misunderstood, perhaps due to ambiguities in Hardy’s
original paper.
The academic response, although inadequate, has been largely open to it.
Despite this, two or three refutations have somehow won primacy.
Two of the attempted refutations are very weak and one was very close and even
this rejected an exaggerated version of the hypothesis.
A mild and moderate definition has been defined and endorsed by proponents.
So.... can we please now mention the dreaded ‘a’ word?
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Question:
[Paraphrased] ‘This isn’t so much a question but a comment. I’m not an
paleoanthropologist but I’m a nutritionist. You mentioned Michael Crawford in your
talk. His work, along with his collaborators, has argued that human brain growth
requires a significant amount of essential fatty acids which are best taken from the
marine food chain. I was part of a team that wrote a paper which argued against this.
We found that eating brains and bone marrow would also provide enough specific
nutrients and energy to fuel the process of encephalisation.”
Answer:
I agree with you. I must say I have never been wholly convinced that we need to eat a
great deal of food from the marine food chain in order to have large brains, as many
people who live a long way from the coast, or who eat little or not fish, seem to
continue to produce infants with very large and healthy brains.
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References:
Ashford, R. W. (1991). The human parasite fauna: towards an analysis and
interpretation. An Trop Med Parasit. 85(1), 189-198.
Attenborough, D. (2002). Life of Mammals. St Helier: Domino.
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