After being shown proudly around the campus of a prestigious American university built in gothic style, Bertrand Russell is said to have exclaimed, “Remarkable. As near Oxford as monkeys can make.” Much earlier, Immanuel Kant had expressed a less ironic amazement, “Two things fill the mind with ever new and increasing admiration and awe … : the starry heavens above and the moral law within.” Today many who look at morality through a Darwinian lens can’t help but find a charming naïveté in Kant’s thought. “Yes, remarkable. As near morality as monkeys can make.”

So the question is, just how near is that? Optimistic Darwinians believe, near enough to be morality. But skeptical Darwinians won’t buy it. The great show we humans make of respect for moral principle they see as a civilized camouflage for an underlying, evolved psychology of a quite different kind.

Read the rest of the article on the New York Times website, then feel free to return here for a conversation with Peter Railton.

http://opinionator.blogs.nytimes.com/2010/07/18/moral-camouflage-or-moral-monkeys/?hp

Darwin’s radical new history of the world did not give a central place to the human. It challenged human exceptionalism and emphasised what was shared, across all organisms extant and extinct. He thought of himself initially as a geologist, so was constantly alert to the ghosting presence of past life forms, visible now only as vestiges, impacted, fossilised, fired, crumbling but discernable, and capable of being re-imagined. In The Voyage of the Beagle encounters with human beings from other tribes and cultures became important and helped his thinking to thrive, but in the years up to and including the publication of the Origin, and well beyond it, his main concerns and researches were with forms of life other than the human, including barnacles and plants. As is well known, he withheld discussion of the human in the Origin, for what he called ‘diplomatic’ reasons. In fact, though, this refusal was profoundly disruptive since it had the effect of simply including us in the general class of primates without a special space or reach reserved. From the start Darwin made it clear that there is no simple opposition between organism and environment since environment is itself composed of the interpenetrating needs, desires, and claims of all the other organisms that surround and include any single being. But it is striking that in trying to describe primordial life he figures the ancestor as single almost as often as he describes it as a pair: the ‘ single progenitor’, ‘one primordial form’.

Such an imagined being is asexual or ‘hors-sexe’, outside sex, and much of Darwin’s research life was spent studying life-forms in which the methods of reproduction are through parthenogenesis (virgin birth, as in some reptiles, fishes and plants), splitting (as with amoebas) or hermaphroditism (as with some barnacles and slugs who fertilizes themselves). Sex is a mechanism of reproduction that speeds up the possibilities of change. In sex two streams of unlike material from the parents enter the progeny, and the spectrum of outcome is much greater. Darwin did not have the language or the knowledge of genetics to work with. He saw the outcome of hyperproductivity and difference but he struggled to explain how such changes were carried between generations. It’s worth emphasizing at the outset that sex difference is not a universal condition since, looking at things from our human point of view, we tend to see it as normative.

Until the 1870s Darwin did not publish extensively about human beings and their descent or liaisons. Then in quick succession he published The Descent of Man, and Selection in Relation to Sex (1871) and The Expression of the Emotions in Man and Animals (1872). That second work was originally to have formed part of the Descent but it grew too large. The Descent itself is on an enormous scale and explores the issue of sexual selection in ways that demanded quite new thinking from Darwin, though it had been touched on in the Origin. In the Descent he brings the human to the foreground of his argument and that produces new tensions in his relation to his readers and in his own mind.

Darwin’s later theory of ‘sexual selection’ placed sex at the centre of explanation, supplementing the emphasis already established in the Origin through natural selection on the resilience of family ties across generations. In the Origin Darwin expanded the idea of family, away from the exclusiveness of what he called “pedigrees and armorial bearings” (Origin 486), to embrace all “the past and present inhabitants of the world” (488) – and by ‘inhabitants’ he did not mean merely the human.

In this brief essay I shall concentrate on a single effect of the newly emphasised presence of the human: what happens when he is writing about man and woman rather than simply male and female. The terms ‘male’ and ‘female’ are fundamental to his argument across species, and to his insistence on the kinship of all species (even those that reproduce through methods other than sexual difference). As soon as his vocabulary enlarges to include man and woman particular difficulties arise – and these difficulties are exacerbated because he does not have a vocabulary that would allow him to discriminate between sex and gender.

The sexual behaviour of different human groups is studied in the Descent alongside that of other kinds, as also are the physical differences between sexes in a range of creatures. And here we begin to see the problem that Darwin has not so much introduced as illuminated by setting the human among other kinds. In his descriptions of species behaviour it is often difficult to discriminate human values from structures. Where he finds physical likeness between the sexes he comments, using the observations of colleagues, on contrasted behaviour:

In one of the sand-wasps (Ammophila) the jaws in the two sexes are closely alike, but are used for widely different purposes: the males, as Professor Westwood observes, ‘are exceedingly ardent, seizing their partners round the neck with their sickle-shaped jaws’; whilst the females use these organs for burrowing in sand-banks and making their nests. (p317)

- a striking example of separate spheres among sand-wasps. Darwin clearly felt some little scepticism himself since he adds a footnote stating that ‘Mr. Walsh, who called my attention to the double use of the jaws, says that he has repeatedly observed this fact.’ And fact it may be, since we cannot just wish away such structural and performative differences between sexes within species, even as we note the gendered interpretation being offered.

Darwin’s later years were spent seeking a system implicit in the inordinate, the decorative, the ornamental, in the drive of sexual desire. Sexual selection demanded flaunting, extravagance, smells and song. The males of most species, his researches showed, were driven to display, the females were the choosers (though ‘choice’ might sometimes be a false word to describe the process of accepting the successful male’s advances). Beauty re-emerged as a key element in his enquiry, and he argued that humans were not the sole possessors of aesthetics and of delight in art.

We have evidence of this capacity even low down in the animal scale thus Crustaceans are provided with auditory hairs of different lengths, which have been seen to vibrate when the proper musical notes are struck. (634-5)

Bird-song was prior to language; it expressed territorial and erotic claims by means of all the pleasures of skilled elaboration. The primordial skills are singing, dancing, and poetry, he argued, skills shared across many species (636). Music becomes his key example of the powers of selection. And he compared the capacities of gnats, dogs, and seals alongside humans. The key point to observe is that once again the whole ground of his argument is the uninterrupted continuum between human experience and that of other life forms, here predominantly animals and birds, but often also including plants.

This discussion of aesthetic life across species and its importance in sexual selection follows one of his most controversial arguments (at least so far as his fellow-humans are concerned), which again relies on analogy (here claimed as homology) with other animals: ‘Differences in the Mental Powers of the Two Sexes’. That is, the two sexes of human beings. He approaches this topic through lengthy discussions of differences between the sexes in a variety of animals, insects, beetles, and birds: in size, in strength, in colouration, in smell, in voice. Once that difference is established he turns in Part III to ‘Sexual Selection in Relation to Man, and Conclusion’. The first paragraph describes the greater muscular development of the male. The second opens boldly, and flatly:

Man is more courageous, pugnacious and energetic than woman, and has a more inventive genius. (622)

The third paragraph opens ‘As with animals of all classes, so with man’.

This equalising between the human and other animals is the argumentative gesture that recurs throughout the discussion, and is in line with all that Darwin has written earlier. It comes as a surprise only because it is here focussed specifically on human beings in a transhistorical and generalising manner, whereas the other crucial aspect of his argument until now has been the emphasis on variability. He acknowledges that ‘some writers doubt whether there is any such inherent difference’ between the human sexes. He has in mind John Stuart Mill, as becomes evident on the next page where he joins in open dispute with Mill. In the text Darwin writes:

Now, when two men are put into competition, or a man with a woman, both possessed of every mental quality in equal perfection, save that one has higher energy, perseverance, and courage, the latter will generally become more eminent in every pursuit, and will gain the ascendancy.

The footnote runs:

J. Stuart Mill remarks ( ‘The subjection of Women’, 1869, p.122), ‘The things in which man most excels woman are those which require most plodding, and long hammering at single thoughts.’ What is this but energy and perseverance? (630)

The tone of exasperation as the qualities slide downhill in Mill’s description from energy and perseverance to plodding and one-track mind sounds as if Darwin has felt Mill’s comments as a personal affront. He has earlier, with a certain ethical self-abnegation spoken of competition, ambition, and selfishness as the ‘natural and unfortunate birthright’ of men. (629) Here, ‘natural’ seems a cover-word for social. Darwin is struggling, and the effect is to make him much more emphatic than is his wont.

Once he substitutes the term ’Man’ for ‘male’, his descriptor for all other species, a rush of social assumptions gathers behind his statements. One is that ‘Man’ (capitalized) in human generalising discourse is to cover both sexes whereas in his descriptions of all other sexed species he painstakingly discriminates between male and female. In a mordant aside he suggests that, unlike most of his argument for sexual selection in which the male displays and the female selects, women may have ‘first acquired musical powers in order to attract the other sex.’ ‘But if so,[he asserts] this must have occurred long ago, before our ancestors had become sufficiently human to treat and value their women merely as useful slaves.’ (639) He sees human behaviour as an aberration in the processes of sexual selection since men seek wealth and beauty in their women, and having social dominance can require that. They do the choosing. Even when women choose:

their choice is largely influenced by the social position and wealth of the men; and the success of the latter in life depends upon their intellectual powers and energy, or the fruits of these same powers in their forefathers.(653)

That women might bring intellectual powers and energy into the marriage does not enter his argument here.

Occasionally, Darwin reaches a different form of inclusivity, ‘human beings’ rather than ‘man’ become his aim:

But we should bear in mind that the activity of the mind in vividly recalling past impressions is one of the fundamental though secondary bases of conscience. This affords the strongest argument for educating and stimulating in all possible ways the intellectual faculties of every human being. (681)

And he argues earlier that :

In order that woman should reach the same standard as man, she ought, when nearly adult, to be trained to energy and perseverance, and to have her reason and imagination exercised to the highest point; and then she would probably transmit these qualities chiefly to her adult daughters. (631)

But he has little hope of equality since men are by him assumed to be destined to be the breadwinners:

they generally undergo a severe struggle in order to maintain themselves and their families: and this will tend to keep up or even increase their mental powers, and, as a consequence, the present inequality between the sexes. (631)

It turns out that it is not quite possible to transpose the physical traits and behaviour of other species to the human and use that to delimit human potential, even in the terms of his own argument. The discussion of men and women’s powers is a matter of about seven pages in a text of seven hundred pages plus, but inevitably it attracts our attention, disconcertingly so. Again, the linking of the two words ‘new and improved’, which, conjoined, lurk at the heart of natural selection haunt this argument too.

Thus man has ultimately become superior to woman. It is, indeed, fortunate that the law of equal transmission of characters to both sexes prevails with mammals; otherwise it is probable that man would have become as superior in mental endowment to woman, as the peacock is in ornamental plumage to the peahen. (631)

But – and this is an important but – Darwin is here describing what has not happened: the law of equal transmission of characters to both sexes does prevail, he acknowledges, against this possible vast mental superiority of man over woman. And that word ‘ultimately’, (‘man has ultimately become superior to woman’) must, if it is to be in key with the rest of his evolutionary thinking even if reluctantly granted here, signify ‘at the moment’ rather than ‘for ever’.

Were I to have the chance to ask him one question I would want to ask him how he sees the changed achievements of women and the altered relations between the sexes in some (dare one say ‘many’?) societies now. Sexual selection as a process is poised between the ‘natural’ and the ‘artificial’ – that is to say, it includes ‘choice’ and discrimination in the drive of desire between individuals and, as he acknowledges, social assumptions and pressures exercise their power in the selection. Darwin’s founding argument that we are ‘all netted together’ across species and that all forms of life are kin, is a wholesome and enfranchising belief. But in the Descent it often seems to have congealed into the assertion that analogies between species debar social change.

The Descent is the work in which Darwin must face the further implications of his insistence on kinship between all organic life, and the place of ‘improvement’ in his argument. The foregrounding of the human forces these issues: male and female become man and woman – and these two are gathered into the title ‘Man’ . He is torn by the difficulty of descrying what is temporary and what eternal in the evolutionary process, what social and what physical in the relations of creatures to each other and in the human sexes too. His intelligence often drives him past the position that his argument can reach.

Darwin rejected Wallace’s belief that the human was a special case, distinguished from other creatures by the possession of a soul, yet he struggles with the question of how far male and female can translate directly into man and woman. Indeed, his daughter Henrietta, who acted as his much-valued commentator and critic while he was writing the Descent, teases him that ‘ you think an apology is wanting for writing abt[sic] anything so unimportant as the mind of man!’(Correspondence, 18, 25) She does not capitalise Man. She knows that for Darwin the human is not the measure of all things and she here pinpoints the difficulty he faced when he wrote a book that placed the human at the centre of his discussion.

References: page references in the text refer to the editions below

• The Correspondence of Charles Darwin, Volume 18, 1870, edited Frederick Burkhardt and James A. Secord (Cambridge: Cambridge University Press, 2010)
• The Descent of Man, and Selection in Relation to Sex, second edition , 1879, introduction James Moore and Adrian Desmond (London: Penguin, 2004)
• On the Origin of Species, A Facsimile of the first Edition, introduction Ernst Mayr (Cambridge, Mass.: Harvard University Press, 1964)

This is a précis of an argument that I developed in an article called “Did Darwin Write the Origin Backwards?” The article was published in 2009 and may be found on my web set at http://philosophy.wisc.edu/sober/recent.html. An expanded version of the argument is the first chapter of a book that I’m publishing at the end of 2010 with Prometheus Books. The book has the same title as the 2009 article.

Although Darwin’s theory is often described as the theory of evolution by natural selection, most commentators recognize that common ancestry (the idea that all organisms now alive on earth and all present day fossils trace back to one or a few “original progenitors”) is an important part of the Darwinian picture. What has been less explored in Darwin studies is how these two parts of Darwin’s theory – common ancestry and natural selection — are related to each other. Ernst Mayr and others have noted that they are logically independent. But this leaves open how the two ideas are evidentially related. How does common ancestry affect the way in which evidence concerning natural selection should be evaluated? And how does natural selection affect the way in which evidence concerning common ancestry should be evaluated?

Darwin addresses one of these two questions very succinctly in a passage from the Origin:

… adaptive characters, although of the utmost importance to the welfare of the being, are almost valueless to the systematist. For animals belonging to two most distinct lines of descent, may readily become adapted to similar conditions, and thus assume a close external resemblance; but such resemblances will not reveal – will rather tend to conceal their blood-relationship to their proper lines of descent.

The fact that human beings and monkeys have tailbones is evidence for common ancestry precisely because tailbones are useless in humans. Contrast this with the torpedo shape that sharks and dolphins share; this similarity is useful in both groups. One might expect natural selection to cause the torpedo shape to evolve in large aquatic predators whether or not they have a common ancestor. This is why the adaptive similarity is almost valueless to the systematist who is trying to reconstruct patterns of common ancestry.

In this passage, Darwin is saying that to determine whether a trait shared by two species is strong evidence that they have a common ancestor, one must be able to judge whether there was selection for the trait in the lineages leading to each. In this sense, knowledge of natural selection is a prerequisite for interpreting evidence concerning common ancestry. However, there is a subtly different question that has a very different answer. Must natural selection have been an important influence on trait evolution for there to be strong evidence for common ancestry? Darwin’s answer to this question is no. A world in which organisms are saturated with neutral and deleterious similarities, while adaptive similarities are rare or non-existent, would be an epistemological paradise so far as the hypothesis of common ancestry is concerned. That’s the point that Darwin is making in the passage I just quoted. Inferring common ancestry does not require that natural selection has occurred.

What about the converse question – how does the fact of common ancestry affect the interpretation of evidence for natural selection? One of Darwin’s most famous arguments concerning natural selection does not depend one whit on common ancestry. This is Darwin’s Malthusian argument. If reproduction in a population outstrips the supply of food, the population will be cut back by starvation. If the organisms in the population vary with respect to characteristics that affect their ability to survive, and if offspring inherit these fitness-affecting traits from their parents, the population will evolve. The process of natural selection is a consequence of these conditions and it can and will occur even if no two species have a common ancestor.

All this is correct, but there is more to the Darwinian picture of natural selection than this. The Malthusian argument establishes that selection has occurred – that some traits changed frequency because of their influence on the viability of organisms. But which traits evolved by natural selection? Darwin doesn’t think that every trait we observe evolved because there was selection for it; recall his comment in the Origin that selection is “the main but not the exclusive cause” of evolution. And if a trait did evolve under the influence of natural selection, why was it favored by natural selection? It is these questions, which concern the detailed application of the hypothesis of natural selection to examples, that common ancestry helps to answer.

An interesting illustration of how Darwin uses the assumption of common ancestry to think about natural selection may be found in his discussion of why mammals in utero have skull sutures that allow them to pass through the birth canal:

The sutures in the skulls of young mammals have been advanced as a beautiful adaptation for aiding parturition [live birth], and no doubt they facilitate, or may be indispensable for this act; but as sutures occur in the skulls of young birds and reptiles, which have only to escape from a broken egg, we may infer that this structure has arisen from the laws of growth, and has been taken advantage of in the parturition of the higher animals.

On the face of it, Darwin’s reasoning here is odd. If he wants to evaluate the hypothesis that mammals have skull sutures because this facilitates live birth, why does he consider the fact that nonmammals have the sutures but not the live birth? Let us hope that he isn’t thinking that if a trait T evolved because the trait facilitated X in one lineage, that T cannot be present without X in any organisms on earth. Penguins do not refute the hypothesis that wings evolved to facilitate flight in birds. And the hypothesis that a species of lizard evolved its green coloration because this color provided camouflage does not require that every green organism on earth gains protective coloration from its being green.

What Darwin is doing in this and in other similar passages is exploiting the fact of common ancestry to test hypotheses about natural selection. The reason that birds and reptiles are relevant to the question of why mammals have skull sutures is that all these organisms share a common ancestor. Common ancestry allows Darwin to infer what happened in the lineage leading to modern mammals. The fact that present day birds and reptiles have sutures but no live birth is evidence that sutures were present in the lineage leading to modern mammals before live birth evolved. If so, the sutures did not evolve because they facilitated live birth. On the contrary, live birth evolved after the sutures were already in place.

Darwin does not spell out the details of this inference, but modern evolutionary biologists will recognize it as an application of the principle of parsimony. Consider the phylogenetic tree shown in the accompanying figure. The tips of the tree represent modern mammals, reptiles, and birds. This is not the tree that a modern biologist would draw, but it may well have been the one that Darwin thought is true. As you move down the page, you are moving from present to past. The lines represent lineages; when two of them coalesce, you have reached a common ancestor. The tree says that mammals and birds are more closely related to each other than either is to reptiles; A2 is an ancestor of the first two, but not of the third. If you go sufficiently far into the past, you will find a common ancestor (A1) that unites all three of these present day groups.

The figure also indicates the traits (±skull sutures; ± live birth) that contemporary mammals, birds and reptiles exhibit. Given this tree, and the features exhibited by its tips,

what is the most reasonable inference concerning the characteristics of the ancestors A1 and A2? The most parsimonious inference is that A1 and A2 both have skull sutures but no live birth. This is the most parsimonious reconstruction in the sense that it requires fewer changes in character states in the lineages leading to the present than any other reconstruction. If this most parsimonious reconstruction is correct, we can deduce that skull sutures evolved before live birth made its appearance in the lineage leading to modern mammals; the mammalian lineage is represented in the figure by a broken line. This parsimony argument justifies Darwin’s statement that sutures now facilitate, or may even be indispensible for, live birth in mammals, but this is not why the sutures evolved.

The argument just described raises an interesting philosophical question: why should we think that the principle of parsimony is a good inferential rule? Why should we think that the most parsimonious hypothesis is true? I won’t pursue this enticing question here. Rather, the point of relevance is that in Darwin’s theory, and in the evolutionary biology of the present, common ancestry is not an unrelated add-on that supplements the hypothesis of natural selection. Instead, common ancestry provides a framework within which hypotheses about natural selection can be tested. In Darwinian biology, a lineage is like a mineshaft that extends from the surface of the earth to deep below, with multiple portholes connecting surface to shaft at varying depths. By peering into a porthole, we gain evidence about what is happening in the shaft. The more portholes there are, the more evidence we can obtain. Thanks to common ancestry, facts about the history of natural selection become knowable.

There is an asymmetry in how common ancestry and natural selection are related to each other in Darwin’s theory. To get evidence for common ancestry, natural selection need not have caused any of the traits we now observe. But to get evidence for natural selection, Darwin needs to be able to think of present day organisms as tracing back to common ancestors. Selection doesn’t make adaptations out of nothing; adaptations are modifications of the traits of ancestors. To know what those ancestors were like, we need to be able to infer their characteristics from what we now observe. It is common ancestry that makes those inferences possible.

If this is the right picture of how common ancestry and natural selection are related in Darwin’s theory, a puzzle presents itself: why did Darwin write the Origin by front-loading natural selection? Darwin does mention some ideas about common ancestry early in the book, but the big picture of there being one tree of life for the whole living world emerges only gradually, and later. On the whole, it is natural selection that comes first. Why is the book structured like this? Why didn’t Darwin begin by defending the idea of common ancestry and then gradually introduce natural selection as a secondary theme?

Introduction

Since Darwin’s time, the human language capacity has been a perennially cited paragon of extreme complexity that defies the explanatory powers of natural selection. And it is not just critics of Darwinism who have argued that this most distinctive human capacity is problematic. Alfred Russel Wallace—the co-discoverer of natural selection theory and in many ways more of an ultra-Darwinian than Darwin himself—famously argued that the human intellectual capacity which makes language possible, is developed to a level of complexity that far exceeds what is achievable through natural selection alone. While fiercely defending natural selection theory with respect to the traits of other species, he argued that in the case of humans, “… natural selection could only have endowed the savage with a brain a little superior to that of an ape.” (p. 392) And Charles Lyell—who personally promoted Darwin’s work and generally supported the evolutionary perspective—also worried that language was just too complex to have evolved by natural means. Not only are the vast vocabulary and baroquely structured grammar and syntax of even the most simple of natural languages orders of magnitude more complex than any other species’ communication system, but the capacity this all provides for expressing esoteric concepts and conveying aesthetic experiences seems far removed from anything with direct adaptive consequence.

Darwin himself fretted over the possibility that natural selection alone might be incapable of accounting for exaggerated functional complexity in nature. In a letter he wrote to Asa Gray shortly after the publication of On the Origin of Species, he admits that “The sight of a feather in a peacock’s tail, whenever I gaze at it, makes me feel sick!” Despite the spectacular and elaborately formed details of this adornment, it was a burden that negatively impacted health and survival and so could not have been the subject to natural selection with respect to the environment. But it was the extravagance of traits such as this, despite their lack of utility, that suggested to Darwin an approach to the challenge of explaining human mental capacities.

In the case of the peacock tail, and other similar traits, Darwin realized that, indeed, something other than natural selection with respect to environmental conditions was responsible. Recognizing that reproduction rather than individual survival was the critical factor in evolution, he argued that competition with respect to reproductive access (sexual selection) could result in runaway selection on certain traits, independent of their environmental suitability. Darwin argued that a display feature or fighting ability that led an individual to out-compete others in gaining access to mates would also favor proliferation and evolutionary exaggeration of these traits, even at some cost to individual health and survival. Analogously, he postulated that selection with respect to sex might also explain such extravagant and highly divergent traits as human language. In his book The Descent of Man and Selection in Relation to Sex—which is typically referred to by only the first half of its title—he argues that language and other human traits that appear exaggerated beyond survival value, can be explained as consequences of sexual selection. So, for example, he imagines that language might have evolved from something akin to bird song, used as a means to attract mates, and that the ability to produce highly elaborate vocal behaviors was progressively exaggerated by a kind of arms-race competition for the most complex vocal display.

Unfortunately, there are strong reasons for doubting the relevance of sexual selection to this most distinctive of human traits. This is because sexual selection inevitably produces complementary divergence of male and female traits, as is exemplified by peacock tails and moose antlers, which are exhibited only by males. While there are indeed a few highly divergent traits distinguishing women from men (e.g. patterns of fat deposition in breasts and hips, etc.), the sexes differ only very subtly in their intellectual and language abilities. Thus accounting for the extravagant complexity of language in terms of sexual selection requires explaining why it lacks this otherwise ubiquitous mark of extreme sexual dimorphism. To explain the origin of the highly structured human-unique adaptation inevitably requires addressing Wallace’s challenge concerning the complexity and apparent non-adaptive aspects of these features.

Long evolution in an artificial niche

In my work I use the phrase, symbolic species, quite literally, to argue that symbols have literally changed the kind of biological organism we are. I believe that we think and behave in many ways that are quite odd compared to other species because of the way that language has changed us. In many respects symbolic language has become a major part of the environment to which we have had to adapt in order to flourish. In the same way that our ancestors’ bodies evolved in the context of the demands posed by bipedal foraging with stone tools and incorporating meat into the diet, their brains evolved in the context of a rich fabric of symbolic cultural communication. As it became increasingly important to be able to enter into the social web of protolinguistic and other early forms of symbolic social communication in order to survive and reproduce, the demands imposed by this artificial niche would have selectively favored mental capacities that guaranteed successful access to this essential resource. So rather than merely intelligent or wise (sapient) creatures, we are creatures whose social and mental capacities have been quite literally shaped by the special demands of communicating with symbols. And this doesn’t just mean that we are adapted for language use, but also for all the many ancillary mental biases that support reliable access and use of this social resource.

But this claim depends on language-like communication being a long-time feature of hominid evolution. Theories suggesting that human language is a very recent and suddenly evolved phenomenon would not make this prediction. To them language is almost epiphenomenal. This is particularly true if the claim is that language appeared suddenly due to some marvelous accidental mutation that transformed dumb (but large brained) brutes into articulate speakers. This sort of scenario has become commonplace in recent years, though the evidence supporting it is mostly very indirect (e.g. archeological evidence of representational forms and objects for adornment, appearing in the Upper Paleolithic). I think that it is mostly a reflection of a caricatured view of the human/animal distinction and a sort of hero metaphor imposed upon the fossil evidence. The way that modern human brains accommodate language can be used as a clue to how old language is.

If language is a comparatively recent feature of human social interaction, that is if it is only, say, a hundred thousand years old or so, then we should expect that it had little effect on human brains. Any structural tweaks of brain architecture that evolved to support it would have had to be either minimal or else major but dependent on comparatively few genetic changes. A recent origin of language would give it little opportunity to impose selection pressure on human brains, so language function would not be supported by any widespread and well integrated neurological changes. This would predict that language abilities are essentially an evolutionary after-thought, inserted unsystematically into an otherwise typical (if enlarged) ape brain. With little time for the genetic fixation of many supportive traits to occur, this adaptation would likely depend on only a few key genetic and neurological changes. As a consequence, language function should be poorly integrated with other cognitive functions, relatively fragile if faced with impoverished learning contexts, susceptible to catastrophic breakdown as a result of certain small but critical genetic defects, and severely affected by congenital mental impairment.

None of these seems to be the case.

On the other hand, if language has been around for a good deal of our evolutionary past, say a million years or so, that amount of time would have been adequate for the demands of language to have affected brain evolution more broadly. A large network of subtle gene changes and neurological adjustments would be involved, and as a result it should be a remarkably well integrated and robust neurological function. Indeed, there is ample evidence to suggest that language is both well-integrated into almost every aspect of our cognitive and social lives, that it utilizes a significant fraction of the forebrain, and is acquired robustly under even quite difficult social circumstances and neurological impairment. It is far from fragile.

The co-evolutionary interaction goes both ways. Languages also have to adapt to brains. Since the language one learns has to be passed from generation to generation, the more learnable its structures, and fitted to human limitations, the more effective its reproduction in each generation. Languages and brains will evolve in tandem, converging towards each other, though not symmetrically. But brain evolution is a ponderously slow and unyielding process in comparison to the more facile evolution of languages. So we should expect that languages are more modified for brains than brains are for language. Nevertheless, if we have been evolving in a symbolic niche for a million years or more, we should expect that human brains will have been tweaked in many different ways to aid life in this virtual world.

The world of symbols is an artificial niche. Its ecology is radically different than the biological niche we also find ourselves in (or at least our ancestors found themselves in). In the same way that beaver dam building has created an aquatic niche to which beaver bodies have adapted over their evolutionary history, our cognitive capacities have adapted to our self-constructed niche: a symbolic niche. This is not a new idea. Indeed the anthropologist Clifford Geertz suggested something like this many decades ago. I think that today we may be at a point in our evolutionary theorizing and our understanding of brains to begin to explore exactly what this might mean.

The most intense and unusual demands of this niche should be reflected in the ways that human cognition diverges from patterns more typical of other species. Although it has long been popular to think of the human difference in terms of general intelligence, I think this bias may have misled us into ignoring what may be a more important constellation of more subtle differences. These likely included differences in social cognition (e.g. joint attention, empathy, the ability to anticipate another’s intended actions), differences in how we learn (e.g. superior transfer learning, a predisposition to assume that associations are bidirectional—known as stimulus equivalence, a comparative ease at mimicking) or even just unusual motor capacities (e.g. unprecedented articulatory and vocal control). These are members of a widely distributed and diverse set of adaptations that fractionally and collectively contribute to our language abilities.

With respect to the brain, we need to confront another mystery. How could these many diverse brain traits have become so functionally intertwined and interdependent as to provide such a novel means of communication? This is particularly challenging to explain because language is in effect an emergent function, not some prior function just requiring fine-tuning. Our various inherited vocalizations, such as laughter, shrieks of fright, and cries of anguish, are comparatively localized in their neurological control (mostly subcortical) as are other modes of communication in animals. In comparison, language depends on a widely dispersed constellation of cortical systems, each of which can be found in other primate brains, but evolved for very different functions. These brain systems have become collectively recruited for language only because their previously evolved functions overlapped significantly with some processing demand necessitated by language, though evolved for quite different functions altogether. Indeed, the neural structures and circuits involved in the production and comprehension of language are homologous to structures found ubiquitously in most monkey and ape brains: old structures performing unprecedented new tricks.

A related mystery concerns the extent to which this dominant form of communication depends on information maintained by social transmission. Even for theories postulating an innate universal grammar, the vast quantity and high fidelity of the information constituting even a typical vocabulary stands out as exceedingly anomalous from a biological point of view. How did such a large fraction of our communicative capacity wind up offloaded onto social transmission? And what explains the remarkable reliability of this process?

Relaxed selection and complexity

Perhaps the most surprising and controversial point to be made follows from the realization of the importance of relaxed selection. The higher-order synergy of systems that contribute to language requires the cooperative functioning of component brain systems. But it appears to paradoxically require that this synergy among diverse systems must already be in place in order for selection to have honed it for language.

The co-evolutionary niche construction scenario sketched above still does not account for the generation of the novel functional synergy between neural systems that language processing requires. The discontinuities between call control systems and speech and language control systems of the brain suggest that a co-evolutionary logic alone is insufficient to explain the shift in substrate. Recent investigation of a parallel shift in both complexity and neural substrate in birdsong may be able to shed some light on this.

In a comparative study of a long-domesticated bird, the Bengalese Finch, and its feral cousin, the White-Rump Munia, it was discovered that the domesticated lineage was a far more facile song-learner with a much more complex and flexible song than its wild cousin. This was despite the fact that the Bengalese Finch was bred in captivity for coloration, not singing (Okanoya, 2004). The domestic/feral difference of song complexity and song learning in these close finch breeds parallels what is found in comparisons between species that are song-learners and non-learners. This difference also correlates with a much more extensive neural control of song in birds that learn a complex and variable song.

The fact that this behavioral and neural complexity can arise spontaneously without specific breeding for singing is a surprising finding since it is generally assumed that song complexity evolves under the influence of intense sexual selection. This was, however, blocked by domestication. One intriguing interpretation is that the relaxation of natural and sexual selection on singing paradoxically was responsible for its elaboration in this example. In brief, with song becoming irrelevant to species identification, territorial defense, mate attraction, predator avoidance, and so on, degrading mutations and existing deleterious alleles affecting the specification of the stereotypic song would not have been weeded out. The result appears to have been the reduction of innate biases controlling song production. The domestic song could thus be described as both less constrained and more variable because it is subject to more kinds of perturbations. But with the specification of song structure no longer strictly controlled by the primary forebrain motor center (called nucleus RA), other linked brain systems can begin to play a biasing role. With innate motor biases weakened, auditory experience, social context, learning biases, and attentional factors could all begin to influence singing. The result is that the domestic song became more variable, more complicated, and more influenced by social experience. The usual consequence of relaxed selection is genetic drift—increasing the genetic and phenotypic variety of a population by allowing random reassortment of alleles—but neurologically, drift in the genetic control of neural functions should cause constraints to become less specific, generating increased behavioral flexibility and greater conditional sensitivity to other neurological and contextual factors.

This is relevant to the human case, because a number of features of the human language adaptation also appear to involve a relaxation of innate constraints allowing multiple other influences besides fixed links to emotion and immediate context to affect vocalization. Probably the clearest evidence for this is infant babbling. This unprecedented tendency to freely play with vocal sound production occurs with minimal innate constraint on what sound can follow what (except for physical constraints on vocal sound generation). Babbling occurs also in contexts of comparatively low arousal state, whereas laughter, crying, or shrieking are each produced in comparatively specific high arousal states and with specific contextual associations. This reduction of innate arousal and contextual constraint on sound production, opens the door for numerous other influences to begin to play a role. Like the domesticated bird, this allows many more brain systems to influence vocal behavior, including socially acquired auditory experience. In fact, this freedom from constraint is an essential precondition for being able to correlate learned vocal behaviors with the wide diversity of objects, events, properties, and relationships language is capable of referring to. It is also a plausible answer to the combinatorial synergy problem (above) because it demonstrates an evolutionary mechanism that would spontaneously result in the emergence of multi-system coordination of neural control over vocal behavior.

But although an evolutionary de-differentiation process may be a part of the story for human language adaptation, it is clearly not the whole story. This increased flexibility and conditionality likely exposed many previously irrelevant interrelationships between brain systems to selection for the new functional associations that have emerged. Most of these adaptations remain to be identified. However, if such a dedifferentiation effect has been involved in our evolution, then scenarios hypothesizing selection for increased innateness or extrapolation from innate referential calls to words become less plausible.

Some concluding speculations

In closing, I would like to reflect on some of the more esoteric features of humanness that may be illuminated by the paired processes of symbolic niche construction effects and relaxed selection.

For example, I think it makes sense to think of ourselves as symbolic savants, unable to suppress the many predispositions evolved to aid in symbol acquisition, use, and transmission. In order to be so accomplished at this strange cognitive task, we almost certainly have evolved a predisposition to see things as symbols, whether they are or not. This is probably manifest in the make-believe of young children, the way we find meaning in coincidental events, see faces in clouds, are fascinated by art, charmed by music, and run our lives with respect to dictates presumed to originate from an invisible spirit world. Like the flight play of birds, the manipulation of objects by monkeys, the attraction of cats to small feathered toys, our special adaptation is the lens through which we see the world. With it comes an irrepressible predisposition to seek for a cryptic meaning hiding beneath the surface of appearances. Almost certainly many of our most distinctive social capacities and biases—e.g. tendencies to conformity and interest in copying the speech we hear as infants—are also reflections of this adaptation to an ecosystem of symbolic relationships. And of course there is literature and theater. How effortlessly we project ourselves into the experiences of someone else, feeling the joys and sorrows almost as intensely as our own.

Relaxation of selection, on the other hand, may have contributed to another suite of distinctively human traits. Widely distributed dedifferentiation at the genetic and epigenetic level would have increased flexibility of a variety of once phylogenetically constrained cognitive and motivational systems. Perhaps the most striking feature of humans is their flexibility and cultural variety. Consider the incredible diversity of marital and kinship organizations. Most species have fairly predictable patterns of sexual association, kin association, and offspring care, and although they are somewhat flexible, this variety is mediated almost entirely by individual motivational systems. In contrast, despite the evolutionary importance of reproduction, human mating and reproduction are largely controlled by symbolically mediated social negotiations. This offloading of one of the most fundamental biological functions onto social-symbolic mechanisms is perhaps the signature feature of being a symbolic species. Thus, because of symbols and with the aid of symbols, Homo sapiens has been self-domesticated and adapted to a niche unlike any other that ever has existed. We have been made in the image of the word.

References

Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (John Murray, London), 1st Ed.

Darwin C (1860) Letter 2743 Darwin, C. R. to Gray, Asa, 3 Apr 1860. Source: http://www.darwinproject.ac.uk/entry-2743.

Darwin C (1871) The Descent of Man and Selection in Relation to Sex (John Murray, London).

Deacon TW (1997) The Symbolic Species: the Coevolution of Language and the Brain (W. W. Norton & Co., New York).

Deacon TW (2009) Relaxed selection and the role of epigenesis in the evolution of language. Oxford Handbook of Developmental Behavioral Neuroscience eds Blumberg MS, Freeman JH, Robinson SR (Oxford University Press; New York) pp 730-752.

Lyell C (1863) Geological Evidences of the Antiquity of Man (John Murray, London).

Okanoya K (2004) The Bengalese Finch: A window on the behavioral neurobiology of birdsong syntax. Annals NY Acad Sci 1016:724735.

Wallace AR (1869) Sir Charles Lyell on Geological Climates and the Origin of Species. Quarterly Review, April, cxxvi: 359-94.

So engrained is the trope of the animal in the West that animal truisms are seared into our shared cultural memory: the Boy Who Cried Wolf, the Wolf in Sheep’s Clothing, The Grannie-Masking Wolf of Red Riding Hood fame, the proverbial Fox in the Hen House, Chicken Little, Foxy Lady, Sly like a Fox, Mary’s Little Lamb, Lassie, Bambi, the Cheshire cat… to say nothing of Puss in Boots, the Dog That Won’t Hunt, Eating Crow, the Golden Egg Laying Goose, the Baby Bringing Stork, the Early Worm Catching Bird, Pop Goes the Weasel, Three Blind Mice, the Three Little Pigs, and the Little Piggy who cried ‘Wee, Wee Wee’ all the way home. Animals have often served as potent metaphors, argues philosopher Max Black (1962:40-2; Blier 1987:206), because they force new forms of interaction and engagement delimited vis-à-vis long standing assumptions. These assumptions challenge credibility (are foxes any more sly than wolves, dogs, or cats?). Related assumptions also reify enduring perceptions of animal-human complements that individually and together heighten the attendant potency of animal symbolism.

Cary Wolfe’s Animal Rites: American Culture, the Discourse of Species, and the Posthumanist Theory (2003) offers a provocative look into the role of animals in human thought and action. Addressing subjects such as animal sacrifice and debates about animal-human difference (the human potential evinced by the use of hands and fingers, for example),Wolfe also investigates larger symbolic systems and practices involving animals. Wolfe’s 2003 anthology (Zoontologies:the Question of the Animal) similarly takes up the boundaries between the animal and human worlds, addressing an array of ethical and philosophical questions posed therein. These are framed around core theoretical engagements (Heidegger, Freud, Lacan, Derrida, Deleuze, and Lyotard among others) as well as questions of interest to the arts. The latter range from Western visual artists (William Wegman and Joseph Beuys) to popular culture (Jurassic Park and Monty Roberts, the “horse whisperer”). One could easily add to this interest not only recent humanities explorations addressing evolution such as the Gottschall and Wilson anthology, The Literary Animal: Evolution and the Nature of Narrative (Rethinking Theory, 2005) but also recent explorations in the field of Environmental Anthropology (Dove and Carpenter, eds., 2007). The co-joining of popular appeal and attendant religious uproar as related to animal portrayals in recent animated films is also striking. Among the recent examples are Avatar (2009) and Happy Feet (2006) (the latter a striking complement to attempts to turn a group of “gay” penguins in a Berlin zoo “straight”). Interesting too arenews stories highlighting of the “split” of the “gay” male penguin pair in New York Central Park’s zoo — as well as accounts of Boston’s female swan couple, Romeo and Juliet, whose very names reveal the central role anthropomorphism assumes in our dealings with the animals around us.

Theory as Trap

When Alfred Gell writes of art as a trap (“Vogel’s Net: Traps as Artworks and Artworks as Traps”-1996), he addresses issues of human-animal engagement (the primacy of prey, the thrill of capture) as well as the enduring dialectic between the ordinary (the animal trap) and the richly articulated (works of art), here heightened by the provocative idiom of animal-human encounter within the context of visual engagement.

Similar issues of human and animal cross currency come into play in various disciplinary and theoretical perspectives. One key early scholar whose work focused on animals in part is Mary Douglas (1957, 1970, 2002) whose writings pointed to core questions of taxonomy (animals seen to transcend “standard” classificatory boundaries) along with issues of pollution and taboo (forbidden animals). In 1982 as a young faculty member at Northwestern University, I co-taught a course with Douglas entitled “Art and Culture.” One of the studies she addressed in class in considerable detail was R. Blumer’s 1967 essay “Why is the Cassowary not a Bird,” a work she republished in her 1973 anthology Rules and Meanings: the Anthropology of Everyday knowledge.” As Blumer argues in this article (1973:167), among the Karam (Schrader Mountains, New Guinea), the reasons why these large emu-like birds are not classed as “birds” has “…no simple, single answer to it apart from the very general statement, ‘The cassowary is not a bird because it enjoys a unique relationship in Karam thought to man.”[i] Oceanic art scholar Douglas Newton would take up complementary portrayals of these unique birds in art shortly thereafter: “Why is the Cassowary a Canoe Prow? (1973).”

Allen F. Roberts similarly has explored an array of issues concerning animal identity and representation in African art (1995), addressing among other questions why so few animal species became the subjects of African visual and performative engagement, and offering an answer that is as poignant as it is simple: namely, some animals are “good to think” with.[ii] Whether as a frame for larger philosophical engagement or as political tropes framed around African creation myths, royal divination icons, religion, and healing arts (see also Blier 1990-1, 1995, 2004), animals serve as a striking idiom through which human values and broader theoretical engagement are grounded. Some animals, such as elephants (Ross 1995), are particularly provocative subjects of both thought and art.

Theories, like animals (and of course humans), have lives. Theories can be said to die not only when scholars prove them “wrong” but also offer viable alternatives. Hence the Lévi-Straussian Structuralism of Douglas and others of her generation was eventually superceded by Post-Modernism (Post-Structuralism), Neo-Evolutionism, or Environmental perspectives, among these the explorations of Cary Wolfe, or Gottschall and Wilson (2005 ), or Dove and Carpenter (2007). British artist and African art connoisseur Leon Underwood published a short illustrated poetry book entitled Animalia, or Fibs About Beasts, Engraved on Wood and Ensnared in Verse (1926). I bring the work into play here, not only because its title is reflected in my own, but also because in many ways the issues he raises are germane. Theories, like objects (and art works, artifacts, or persons) ensnare us with provocative “fibs” (partial views, unique vantages, half truths). In some ways, the primacy of theory lies not only in issues of universality (science) but also its “art” value (perspectives that press one to think about things in a new way). Theories, like animals, matter, in my view, not necessarily only because they are “right” or “wrong” but also because of the challenge they pose to us to think about (and through) things in a new way. The death blow to a given theoretical frame often can be sensed when everyone finally “gets it.” At the risk of pushback, I wonder how much it matters if a given theory is “right” or “wrong” as long as it is something “good to think with,” a frame (trap, ensnarement, vantage, partial view) that elicits new insight, even if it may have been proved outdated or false.

Localities: Place/Time Trap

Many portrayals of animals in African art (masks among other forms) are notable in part for their striking elements of anthromorphism, among these earrings (on Bamana Chiwara masks), royal regalia (Ife terracotta sculptures), performative elements (Dogon monkey maskers as pick pockets), and roles in human-deity communication (divination – hornbills, spiders, crocodiles, and pale foxes, to name but a few). This is true not least in Ife, the ancient capital of the Yoruba in southwestern Nigeria. In this important center, one comes across numerous animal species on sale in the local animal market near the Ogun Mogun temple complex in front of the palace. One can find here not only domestic animals for sale but also an array of wild animals – some of which are sold alive (chameleons, snails and certain birds), others sold as parts – the skin of leopards, crocodile teeth, the meat and/or horns of varied antelope species among these. I visitedthis market on numerous occasions and observedthe sale of these animals in this setting near the main temple of Ife’s hunting god, Ogun. Knowing the importance of diverse plant and zoological species in Ifa divination and other contexts, as well as the enduring significance of related beliefs in this center, makes clear the iconic complexity of animal portrayals in ancient Ife art. Local knowledge (place, time, social frame) is critical to our understanding of these works (Geertz 1985).

The corpus of ancient Ife animal sculpture offers a rich template from which to examine similar concerns. Here too, there are also notable differences in how animals (even within the same species) are portrayed. Some are identified with regalia fit for kings. Others are secured with cord leads suggesting imminent sacrifice, offerings which in some respects complement human ritual deaths. In the end what is especially remarkable in these animal personifications is their primacy in the early art corpus and the remarkable detailing with which they are rendered, attributes which also conform with some Ife canons on human portrayal as well. In some ways, portrayals of animals are among of the most complex and interesting artistic exemplars because of expectations and the fact that models are not always readily available.

Several days before the end of my first research trip to Ife, capital of the Yoruba in southwestern Nigeria, I learned that not far from the city there lived a mudfish, some five feet in length and residing in a spring and adjacent brook which was part of a local healing shrine. Years earlier an Ife acquaintance had been treated successfully here for a painful ear infection and fever. I decided to visit this amazing mudfish that purportedly would come out of the water and onto land for food treats. Like others of the lungfish species, the mudfish possesses ancillary lungs that enable it to survive out of water for weeks and months at a time. After accessing this shrine on foot via a long winding dirt track, I interviewed the priest healer in charge and learned something of the temple’s history. A devotee of Mami Wata (a water goddess linked to European and west Asian trade, who brings material and other benefits to believers) from the Urhobo in the Niger Delta area to the south, the priest had moved to the Ife area in his youth. From him I purchased a small bag of the mudfish’s favorite cookies and waited expectantly at the edge of the spring as he tossed the crumpled cookies on the water surface while offering prayers intended for the ears of this remarkable being. Soon the spring’s surface began to shimmer with tiny bubbles and myriad mudfish and other aquatic denizens gulped down the proffered sweets. Alas, the giant mudfish, already satiated from a healing ceremony before my arrival, chose not to come to land. What little I did discern of this fish nonetheless reinforced the uniqueness of the species and why such animals figure so prominently in early Ife art.

Mudfish drawing by Blier

A particularly striking thirteenth to fourteenth century C.E. terra cotta mudfish depiction historically was housed in the city’s Omitoto shrine in Ilorin quarter — under the supervision of Ife’s powerful Chief Obaloran, head of the Ilode ward where the Obatala temple is located. The terra cotta mudfish long associated with his family is known locally as Orisa Ito. During the annual Odun Ose festival dedicated to Omitoto, held every year in October at the beginning of the dry season and related harvest-New Years activities, the sculpture (which most likely dates to the late thirteenth or early fourteenth century) was carried during the related rituals by a ritual specialist dressed in plantain leaves. While the exact origins of this procession are not clear, the term odun refers to year, and ose identifies the traditional five day Yoruba market week, underscoring the likely importance of this event (and the mudfish referent) with the cycle of time, and specifically the yearly transition from dry season to rainy season. The mudfish, with its unusual ability to estivate (that is, to survive for months in a hibernation-like state in dried spring or river beds using its ancillary lungs), seemingly comes “back to life” after rains again fill its resident pool or river bed with water. As such this animal is a particularly apt referent to the cycle of seasonal transition. Omitoto, who is also identified (Fasogun n.d.1) as having “carried igba-iwa to Ile-Ife,” was a central figure in the earth’s beginnings at Ife.[iii] This same igba-iwa vessel is said to have held the necessary Ife offerings conveyed to the heavens each year in exchange for well-being and plenty (Fabunmi (1985:194). The identity of the mudfish at once with the earth’s creation, and with rituals which preserved life more generally, also is in keeping with the mudfish’s unique identity with seasonal transition — not only the beginning of the dry season, but also the arrival of the rains. Indeed this unusual fish is said by some to vomit water at this time into the dried river beds and springs, thus assuring their renewal to meet the needs of local inhabitants.

Mudfish are among a group of unusual animal species that figure centrally in the corpus of ancient arts from Ife and other Yoruba centers. A variant of these mudfish forms is conveyed through the image of a human form with outward extending fish or snake-form legs or feet. This is a form that in Benin royal art has come to be identified with both the ruler Oba Ohen (who was paralyzed) and with Olokun, god of the sea. The striking bi-morphism of these “fish” — as denizens of both water and land — provides them with unique symbolic potency, in relationship to (among other things) ideas of autochthony.

Pennant wing nightjar drawing by Blier

Two unusual birds share similar visual and symbolic potency in ancient Yoruba art. The first is the Pennant Wing Nightjar, a small owl-like bird species that migrates to this area during the rainy season, where it mates, during which time senior males grow long wing streamers double the normal size, that ripple in the air in flight, in a manner suggestive of snakes and or lightning. These birds figure in several ancient Yoruba area art works (a Janus terracotta man-bird figure from Ife and several bronzes from the Niger River area to the east that seem to be linked to this center). In both cases they appear to be identified with the ancient Ife thunder god, Oramfe. There is a somewhat complementary bird, the African Paradise Flycatcher (okin in Yoruba). Both birds are notable for their bi-morphism and for their rarity and distinctiveness, suggesting a possible conceptual link between them. During pairing season, senior males of the African Paradise Flycatcher species grow long tail streamers double the normal seventeen-inch length of their tail feathers. These streamers, which ripple as the birds fly, are incorporated into royal Ife and Yoruba crowns.

Both avian species suggest the importance of both anomaly and change in the animal world as a reference to ritual and political primacy. These three animal species all have unique local primacy (and symbolic power), yet all three appear to have early foreign roots, most likely in ancient Greek, Coptic, and Early Medieval European idioms of fish legged Sirens (the early source for Mami Wata here it appears[iv]) and snake winged Harpies (see below) as also do early Gorgon motifs in this area– human heads with snakes emerging from the nostrils or ears). These motifspoint to notable visual and iconic similarities with, among others, twelfth to early fourteenth works from pilgrimage sites in Great Britain and France. Most likely complementary formsenteredinto the area of central Nigeria around the same time as part of richly colored Coptic textile forms from Egypt from this era.Arriving here perhaps around the same time were early forms of Nubian jewelry (see below), works that also appear to have left its imprint on one of the thirteenth to fourteenth century fragments from Ife, a ram head pectoral from the site of Oke Eso.

drawing by Blier, click to enlarge

drawing by Blier, click to enlarge

drawing by Blier, click to enlarge

Since the ram, like the mudfish and bi-morphic birds, is so closely integrated into the fabric of Yoruba belief, does it really matter where these motifs originally came from, when they arrived, or how? Or to the contrary, since all symbolic forms ultimately are delimited locally (their meanings, contexts, qualities of empowerment specific to place and time) — one wonders whether these local perspectives necessarily carry more symbolic or theoretical weight than others that are in play.

Bibliography

Blier, Suzanne Preston

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• 1990 “King Glele of Danhomé: Divination Portraits of a Lion King and Man of Iron (Part I),” African Arts 23.4: 42-53, 93-94.
• 1991  “King Glele of Danhomé: Dynasty and Destiny (Part II),” African Art 24.1: 44-55, 101-103.
• 1995  African Vodun: Art, Psychology, and Power. Chicago: University of Chicago Press.
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• Forthcoming. Past Presence: Ife Art in Yoruba History.

Blumer, R.

• 1967   “Why is the Cassowary Not a Bird? A Problem of Zoological Taxonomy Among the Karam of the New Guinea Highlands” in Man, Vol. 2, No. 1 (Mar., 1967), pp. 5-25. Republished in  “Why the Cassowary is not a Bird” in Mary Douglas Rules and Meanings: the Anthropology of Everyday knowledge” (1973: pp 167 ff)

Douglas, Mary

• 1957 “Animals in Lele Religious Symbolism,”  Africa, XXVII, 1:  46-58.
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Drewal, Henry John  et al.

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Fabunmi, M. A.

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Fasogun, M. O.

• n.d.. “A Brief Constitutional History of the Ancient City of Ile Ife Ooye Lagbo (the City of the Survivors)” Seminar Paper presented at the Conference on Yoruba Civilization, University of Ife, Ile-Ife. July 1976.

Geertz, Clifford

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Gell, Alfred

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[ii] This idea extending back to Claude Lévi-Strauss among others.

[iii] Perhaps in part for this reason, Fabunmi  suggests (1969:.5) that Omitoto-Ose “…held a very important office in…government [performing her role] with great credit.” He adds: “Some Ife historians say that she had no issue but she adopted Obaloran as a son and brought him from his home in Iloran to her own home at Ilode where the present Chief Obaloran still lives.” Fasogun however identifies  (n.d.1) Omitoto as one of the original thirteen chief-priests of Ife (Chapter-Op), suggesting that this chieftaincy only later was taken on by Obaloran.

[iv] A later era diffusion of the “siren” motif in southern Nigeria has been advanced by Henry J. Drewal, whose Education Ph.D. (Teachers’ College, Columbia) rather than more standard academic degree program (Art History or Anthropology for example) seems to a more comparative rather than a more  historical or theoretical vantage.

To understand portions of one’s own culture demands a lifetime; to become familiar with another’s depends upon a host of enthusiastic interpreters, attentive listening, experiencing a multitude of unfamiliar activities, a receptive heart, and good fortune. Throughout my life, a major focus has been the contrast between Northern (Western) and African perceptions (neither of which is homogeneous) and their relationships to wildlife and how these motives and practices have played out in conservation policies and through political-economic power. How have such policies affected local or indigenous populations? How thoroughly have they disrupted earlier relationships? What about the impacts on ecological interactions? Undoubtedly, my concern is part of a legacy from a childhood and youth spent equally in temperate North America and tropical Central Africa, of immersions in dissimilar cultural and environmental settings, an early absorption of three languages, of childhood roles and adventures under conditions favoring explorations in both cultural and environmental domains, and of subsequent career choices allowing the time, resources and the connections to follow these interests and to write about them (Marks, 1984,1991,2005, nd.).

This passion for different world views and for wild animals came as I reflected upon earlier experience while temporarily stationed among the Inuit in the Bering Straits in 1962. While there, I realized that my Inuit associates were not seeing the same animals that I was taught in graduate studies. My hosts on St. Lawrence Island possessed a more inclusive connectedness to the lives of neighboring creatures, expressive of longer term relationships, both spiritual and epistemological, upon which their welfare depended. These exposures and thoughts challenged me to expand my own awareness beyond what was considered then appropriate in the academy. Accepting this ordeal has taken most of a life time to observe cultural and environmental processes in just two small places. In the process of completing intermittent field studies with members of one central African society (Marks, 2008; nd), I am asked to share some musing with readers of On the Human. I will give some background and findings from this longitudinal study before posing some questions that may affect, if not implicate, us all.

In Zambia’s central Luangwa Valley, the Valley Bisa share their landscape with dense populations of large and small wild animals. These Bisa have been my hosts and teachers during intermittent stays spread over half a century. Like many of their neighbors, this society is matrilineal and organized into chiefs’ and commoners’ lineages. They are subsistence hoe cultivators, dependent upon rain-fed sorghums/maize, upon collecting and hunting wild products, as well as upon wage employment. Gender largely determines who farms, cooks, raises children, collects, hunts, or seeks wages. Residents’ numbers have doubled in fifty years (10,000 people in 2006), now skewed decidedly towards the younger ages while their communities are challenged by a weak government’s retrenchment in education and in other social services. Today most residents are experiencing growing poverty and persistent resource scarcities as shifting climate regimes affect their subsistence agriculture and as government edicts restrict their gathering and hunting of bush products. Unlike their neighbors, the Valley Bisa were a comparatively small and “marginal” group somewhat distanced from outside administration until recently (accessible by a rough, unimproved road since 1960), yet with sizeable portions of their land appropriated initially by the colonial and later the Zambian state as game reserve and as national parks respectfully. Today, they inhabit a narrow Game Management Area (a “buffer corridor” of some 2500 km2) surrounded on three sides by national parks with a steep escarpment on the fourth which separates them somewhat from developments elsewhere.

Since the 1980s, these imposed institutional boundaries, supported in the “mental furniture” of conservation officials, backed by considerable international funding and enforcement on the ground, have had a devastating impact on Valley Bisa welfare and culture. One way to illustrate this quandary is to list the names given to their domestic dogs, which, as the state has disarmed the local population, have become residents’ close associates in their conspiracy against the state’s limited economic vision for wildlife. Unlike the domesticated dogs found in many Northern societies’ households, which are brought into the family hearth, well-fed, and treated much as kin, the names of and condition of Valley Bisa dogs are symbolic of their despair and fragmenting social relations. Dogs are rarely fed or cared for and, in times of duress, are sold sometimes to safari operators for target practice. In 2006, these given-names reflect the recent individualization and social alienation taking place under increasing uncertainty and poverty, for the monikers were either derogatory or punitive evocative of unsanctioned sentiments (even in translation): “we have no relatives” [after no one came to inquire about a wife maimed by a crocodile], ”hatred”, ”not yours”, “no justice”, “remember me”, “not sure why I married this woman”, “if your marriage is unstable- you will travel”, “mistake”, “shut up”, “you offend the whole household”, “jealousy”, “rudeness”, “chaos”, “no appreciation”, “stinginess”, and “you will see.” To me, this litany is symptomatic of the depth to which a once brave and resourceful people have descended in their relationships with their neighboring animals and to each other.

Valley residents depend upon wildlife as an important complement to their agricultural products and especially as a safety net in times of famines. For some of their men, the hunting of and protection from wildlife are both necessary and customary. The necessity comes from the endemic presence of the tsetse fly, which prevents livestock husbandry, and from the need to protect human life and crops from wildlife competitions both in its large and smaller forms. Wild meats are important supplements in diets and the produce of a few men selected by their lineages. Gender roles dictate that women engage in “mundane” agriculture and serve as the structural core of villagers, while most men assume the expansive and chancy activities of hunting, trading, and local or migrant labor.

Valley Bisa relationships with the wild animals around them are diverse, complicated, and, in some cases, seem contradictory. While wildlife competes with cultivators and collectors for food, some animals reciprocally become important sources of meat and of power when used as medicines and in witchcraft. This dialectic is the basic social organizing principle in which matrilineally-related women, identified symbolically with subsistence agriculture and the “community”, are contrasted to wildlife and its potential destructiveness of human life and sustenance. Men (mainly marrying into the group) with their wide-ranging activities are identified with wild animals, hunting, trade, and the bush (Morris 1998). Local people employ familiar concepts to classify wild animals including their grouping, relations between these categories as well as the same images to interpret behaviors, “spirits,” and power. Their folk classifications express utilitarian and anthropocentric values. Historically, relationships with and knowledge of wildlife were the domain of few men belonging to specific lineages.

Whereas “mammals” are presumed to have their own autonomous reality, local hunters’ categories encapsulate lineage interests, needs, and uses as well as expressions of personal fears, histories, and experiences. Historically, hunters managed and addressed their roles tangentially through beliefs in spirits, through the tangible uses of culturally mediated rituals and prescriptions, and through following the normative distributions of its products and procedures. Lineage elders used unseen agents (spirits, ancestors) and observable agency to monitor the compliance of their subordinates while imaginatively structuring and legitimizing the ethical order in times of crisis and uncertainties.

Linguistically and historically, the Valley Bisa did not separate themselves from other forms of life nor do they typically objectify “nature.” No indigenous word exists for the Northern idea of “nature,” or “environment” as such, although their noun “nchende” is sometimes translated as such by outsiders pushing a conservation ethic. The vernacular meaning of this term includes people, place, and the resources (“fipe”-baggage, goods, or “properties”) necessary to sustain people within a particular site. The term to convey something of the meaning for a “natural resource” is “ifilingwa waleza” (literally God’s gifts). The ideal of wildlife “conservation” requires a whole phrase- “kusunga ifilingwa waleza” (caring for God’s gifts). Yet, this indigenous term denotes more intrinsic and spiritual meanings than the English term, as it assumes that humans and the other lives around them constitute a seamless whole. Both are integral parts: no “nature” exists outside the morality of the human community, for reciprocal obligations extend outward from the village embracing other forms of life as well as spirits. The bush becomes responsive and responsible to residents as their ancestral spirits reside there as former embodiments of the current community. Causality embedded in moral principles and human intentionality are the bedrock explanations for why “good” and “bad” things happen; the latter might happen even to “good” or innocent people because someone, somewhere has violated ethical expectations and norms. The “how” and the “why” questions of life are often embedded in the same search.

For most Valley Bisa, their recent transformations have been precipitous and traumatic, brought on by many dynamics seemingly outside their influence, and expressed through the consciousness of increasing scarcities and decreasing welfare. Within this synthesis of progressive factors is a steady inflation in the national economy since the late 1970s, the death of a long-reigning chief in 1984 with an interregnum until 1990, a weak and truculent government unresponsive to local needs, uncertain rainfall regimes and climatic shifts. In addition, the AIDS epidemic and a doubling in population size (since the 1960s) mean that the majority of residents are young, with little formal education and facing local resource and productive land scarcities as well as few opportunities for employment. This demographic shift has brought its losses in cultural and local ecological memories as many residents reject the earlier “limited” communal worldview of ancestors for conversions to Pentecostalism with its individualistic expectations and anticipated rewards. Under donor pressures, government has further aggrieved local welfare by legally commoditizing the value of wildlife to generate revenues, thereby privileging access by safari hunters and international tourists rather than local users. Towards this goal, the administration employs large cadres of wildlife police officers to arrest those killing wildlife without formal licenses and harassing others while dispossessing residents of the firearms formerly used to feed and to protect themselves. Officials offer no proactive protection to residents or compensation for their losses. While enduring a high level of arrests and losses, residents resort to “hidden transcripts” (secrecy) and earlier devices (snares, downfalls) to deliver their protests, to protect their properties, and to acquire their animal protein. Another way is through the recent husbandry of domesticated dogs as co-conspirators in protection and for acquiring prey.

I do not pretend to present a final, definitive picture for these cultural dynamics or for the biological commons in this central Luangwa Valley. What I have witnessed will continue in various shapes and versions as current cultural tragedies and policies continue in that part of the world. Finding the words to match the meanings and expressions for my experiences in this distant valley has, for me, become an instructive hunt, if only a mental one. Nevertheless, this quest has enlarged my range in curiosity, taken me across new conceptual terrain and provided different “targets” of opportunity. The pursuit to reconfigure “the place of nature within the space of culture” essentially becomes one of redefining “ownership”, “possessions”, and “belonging” and remains necessarily elusive as the perpetrators in time often become victims (Buell, 2001). Comments of fellow trekkers, as well as those of others, are welcome as what I have learned leads me to some inferences about the acquisitive structures and presumptive natures of our (northern) societies. I conclude with a few of these reflections.

For an indeterminate past, wild animals were around and with us, in our minds if not in our stomachs, and vice versa. According to our myths, these associations were essential for humans evolving and for reflective definitions of our humanity. Humanity’s place within nature became a topic of modern European philosophers, who agreed on human superiority even as they differed on the specifics as to what humans had that other animals lacked. It is assumed that only humans have language and practical intelligence that allows imagination, speculation, and deliberation about death and what comes after. With European exploration and colonization worldwide, and later its own industrialization, wild animals began to disappear from human life as environments became fundamentally transformed under the egis of ideas about hierarchy, dominance, and utility. Today most descendants of and operatives within these European worldviews must travel far to witness the diversity of wild animals, even if these animals are now found in contrived surroundings elsewhere; otherwise, they remain surrounded by domesticated types bearing human utilities as pets and food. As human livelihoods and wants have become the major impact on the evolutionary trajectories of most forms of life, I wonder if animals, particularly in their “wilder” forms, will continue as a major epistemological category in human development and thought. If animals are no longer the standards, what might take their place as holistic contexts on life recede and comparisons become increasingly reductionist? (Thomas, 1963; Lippit, 2000)

Scott Atran and Doug Medin (2008) show how the remarkable breadth in biological and ecological knowledge of some indigenous people compares with that of modern literates in the United States. Some of these smaller groups, whose worldviews include spiritual and ethical links to their environments, strive to maintain “sustainable” resources within livable environments, often while in conflict with more powerful and imperial groups claiming privileged (but truncated) worldviews based in distant and “unsustainable” cultural appetites for material resources. Other indigenous groups, such as the Valley Bisa, resist the odds by persisting with their own claims and identities despite its high cost. What I find remarkable about this discrepancy in biological knowledge is that many people in the Northern Hemisphere, even those working for organizations proclaiming their mission as protecting the environment, seem oblivious to the limitations of their own perspectives and prefer to remain in the dark about the high “hidden” human costs in their own overseas activities. My view is that we would learn a lot from listening and learning from people who know about formal, and even informal, restraints and from long term perspectives that our histories have repeatedly yet to teach us. Unfortunately, we seldom venture beyond our invented environments and the comforts of our insulated lives, including the pets that bear our marks, as seekers rather than as tourists.

What I have described for the Valley Bisa is not a unique event, for similar episodes have occurred in the past and continue into the present. In many respects, these incidents are reminiscent of persistent biological drives submerged in imperial cultural demands for resources and territory. In this case, it would be a basic biological need for territory and resources sanctioned as a cultural necessity controlling (managing) disorder, unknowns, and diversity spatially (Sheets-Johnstone, 2009). In a recent remarkable book, Anderson (2004) depicts how colonists in North America used their domestic livestock to undermine indigenous Indians of their rights to land and resources, which once obtained, settlers then transformed into commodities that they considered more manageable. Anderson (2004:246) concludes this tragic story thus: “Indians found room in their world for livestock, but the colonists and their descendants could find no room in theirs for Indians.”

My experience leads me to think that those who strive to preserve biological diversity in terms of their own worldviews, restrictive in its visions of cultural diversity, are imperially pushing their own control of “nature” rather than broadening our common understanding about what “sustainability” of life might be about. For the Valley Bisa, the story begins with wild animals, a cultivated and cultured landscape; it ends with domesticated animals bearing the imprints of their makers, with the human vision dimmed, the land cartelized by new proprietors none the wiser. The national park might just be “America’s best idea,” according to the recent Ken Burns’ documentary; yet a closer look at this cinema graphic shows that it is really about our violent history and displacement of indigenous peoples, about our heroes and villains, about the cultured versions of our interpreters and scholars, about our technologies and acquisitiveness, about our class-based disparities in wealth, about our cultural concepts of work, play, and leisure, as well as our definitions of a “good life.” How can such an idea be expanded and disciplined to serve a more universal ideal?

Some References

• Anderson, Virginia 2004. Creatures of Empire: How Domestic Animals Transformed Early America. New York: Oxford University Press.
• Atran, Scott & Douglas Medin 2008. The Native Mind and the Cultural Construction of Nature. Cambridge: MIT Press.
• Buell, Lawrence 2001. Writing for an Endangered World: Literature, Culture, and Environment in the U.S. and Beyond. Cambridge: Harvard University Press.
• Crist, Eileen 1999. Images of Animals: Anthropomorphism and Animal Mind. Philadelphia: Temple University Press
• Diamond, Jared 2005. Collapse: How Societies Choose to Fail or Succeed. New York: Penguin
• Goodman, Nelson 1978. Ways of Worldmaking. Hassocks UK: The Harvester Press.
• Hughes, David M. 2008. Requiem for the Zambezi Valley? Conservation and protected areas under climate change. Policy Matters 16:108-115.
• Lippit, Akira M. 2000. Electric Animal: Toward a Rhetoric of Wildlife. Minneapolis: University of Minnesota Press.
• Marks, Stuart A. 1984. The Imperial Lion. Boulder, CO: Westview Press
• _________ 1991. Southern Hunting in Black and White: Nature, History, and Ritual in a Carolina Community. Princeton, NJ: Princeton University Press.
• _________ 2005. Large Mammals and a Brave People. New Brunswick, NJ: Transactions (2nd edition)
• _________ 2008. On the Ground and in the Villages: A Cacophony of Voice Assessing a “Community-based” Wildlife Program after 18 Years in Zambia. under review
• _________nd. Life as a Hunt: Thresholds of Identities, Images, and Illusions on a Central African Landscape. under review
• Morris, Brian 1998. The Power of Animals: An Ethnography. New York: Berg
• Posey, Darrell A. (compiler and editor) 1999. Cultural and Spiritual Values of Biodiversity. London: Intermediate Technology Publications for the United Nations Environment Program.
• Sheets-Johnstone, Maxine The Descent of Man, Human Nature and the Nature/Culture Divide. Public Lecture given in symposium “Darwin Across the Disciplines” at Duke University, November 6, 2009.
• Shepard, Paul. 1978. Thinking Animals: Animals & the Development of Human Intelligence. New York: Viking.
• Sinton, John. 1993. When Moscow Looks Like Chicago: An Essay on Uniformity and Diversity in Landscapes and Communities. Environmental History Review 17(3):23-41.
• Tambiah, Stanley J. 1990. Magic, Science, Religion, and the Scope of Rationality. Cambridge: Cambridge University Press.
• Thomas, Keith 1983. Man and the Natural World: A History of the Modern Sensibility. New York: Pantheon Books.

The use of hounds in hunting excites great passions. Hunting deer is particularly hated by those who are opposed to it and ardently loved by those who support it. If you wept as a child at the death of Bambi’s mother, you know what it is like to be hunted. On the other side, the Hunt supporters have believed sincerely that very little suffering is involved in hunting with hounds. They regard this method of culling red deer not only as necessary for the protection of the environment but also as an entirely natural process. Wolves chase deer, the argument runs, so deer should be adapted to being hunted by hounds. The battles have raged for the best part of a century in terms that have changed not one bit.

In 1997 I submitted to the National Trust my report on a scientific study of the welfare issues involved in the management of red deer on Exmoor and the Quantock Hills. Among other things, I had been asked to examine the evidence for stress induced in red deer by hunting with hounds and to compare this with the stress resulting from other culling methods. The law is stringent about the use of animals in scientific work. Like hunting, scientific research involves great companionship, much skill and the thrill of the chase. However, as attitudes to animals changed, scientists have had to temper their enthusiasms for their work with concerns for the welfare of the animals they use. The Animals (Scientific Procedures) Act of 1986 stipulates that: “If procedures used in research involve pain or discomfort, the investigator must consider whether the knowledge that may be gained justifies the stress and pain inflicted on the animals.”

Pain is clearly defined in terms of human subjective experience. So, for that matter, are fear, distress and suffering. Nevertheless, those who must obey the existing legislation on animal welfare have to project these unpleasant states into animals. They are required to make the same sort of assessment of another creature’s condition as they do implicitly and routinely when dealing with a fellow human being. In humans each unpleasant state is associated with observable behaviour and with identifiable physiological processes. A profile of these characteristics may be built up and considered when taking any particular case of questionable animal welfare.

Weighing suffering against human benefit is inherently unsatisfactory because they are not measured in the same terms. What can be done is to find an acceptable space in which suffering is kept to a minimum and humans maximise what they can get out of the use of the animals. When I started my investigation of the hunting of deer with hounds, I supposed that here again we should probably finish up with some notion of acceptable space. Hunting undoubtedly gives great pleasure to those who partake in this activity as sport. To many people, witnessing a hunt is to feel part of English history. I had supposed that, if the suffering of the hunted deer were contained, the positive aspects of stag-hunting could be supported. However, the science led my Research Associate, Dr Elizabeth Bradshaw, and me to a very different position.

Science can make contributions to the hunting debate at various levels. The movements of red deer may be monitored by radio-tracking. This method involves fitting a collar on the animal which contains a small transmitter emitting a regular signal which can be detected at some distance by a receiver. Apart from long excursions by the stags before and after the rut in the autumn, red deer on Exmoor spend 95% of their time within about half a mile of the same place. The ancestral habitat of red deer is woodland and, in such habitats, wolves do not chase them for long distances. Instead wolves rely on stealth, short bursts of speed and ambushing to catch the deer. Further, red deer are not equipped with sweat glands, easily over-heating when chased, and their muscle fibre type is not that of an animal adapted for endurance running. Armed with this knowledge, it is startling to discover that the average hunt with hounds last about 3 hours in which time the deer has been run 12 miles. The use of a standard technique in behavioural biology and the gathering together of some well-established facts about red deer suggested already that hunting with hounds is not natural. However, even this did not prepare Dr Bradshaw and myself for the astonishing changes in the physiology of the hunted deer which we discovered from their blood after they had been hunted.

The absolute levels of stress hormones are as high as have ever been found in red deer and do not differ from animals with very serious injuries. The carbohydrate resources for the muscles are totally depleted in animals that have been hunted for long periods. Acidity of the blood, resulting from great exertion, is very high at an early stage in the hunt. At an early stage in the hunt the level of haemoglobin in the plasma jumps to eight times what is found in undisturbed animals and then continues to rise. Much of this is probably due to the break up of the red blood cells. In longer hunts extensive leakage of enzymes from the muscles occurs. In some deer these levels are so high that they are likely to be due to actual damage to the muscles. In short, many of the physiological changes are seriously maladaptive and would not be expected to occur in normal conditions. The pattern of the data is entirely consistent with the view that the hunted animals are extremely frightened, pushing themselves as much as they are able and risking a great deal in their attempts to escape.

If deer are to be culled, the only realistic alternative to hunting red deer with hounds is to shoot them. We compared the suffering resulting from stalking with that produced by hunting with hounds. The critical issue is the frequency of wounding. While just over 11% of red deer are wounded when shot, the majority of these were then quickly killed. A maximum of 5% of shot deer are likely to escape wounded. At the time of assessment, which occurred immediately after death, the physiological effects of wounding by shooting are comparable to those of a long hunt. An important missing dimension is the length of time for which a deer has to endure its suffering. In this context, it should be appreciated that half the hunted deer escape. Some escaping deer may die from the effects of the long chase. Others are likely to experience the consequences of the long stressful chase for days afterwards.

Of the 130 or so red deer killed annually by the Hunts, we believe that all experience an unacceptable level of suffering because of the stresses and strains put on them. At least a further 100 that escape would suffer because of the distance travelled before they escaped. This makes a conservative total of 230 deer a year presenting a serious welfare problem. If the 130 or so animals killed by the Hunts were culled by stalkers instead, then on the basis of the 5% wounding estimate we obtained, less than seven deer would suffer because of their injuries. These are broad calculations but the great reduction in numbers of suffering animals is obvious.

Hunting with hounds can no longer be justified on welfare grounds given the standards applied in other fields such as the transit and slaughter of farm animals, the use of animals in research and so forth. This is the key conclusion of my report to the National Trust. The Trust had to weigh this conclusion against other issues, including their wider responsibilities, considerations about the social and economic benefits of hunting and the problems of conservation. At the time of writing this article, I was uncertain what the Council of the National Trust would decide. The result of a ban on hunting could be an increase in indiscriminate and inexpert shooting which might increase the proportion of deer injured from shooting and also reduce the overall red deer population on the Quantocks. The judgements involved are not easy ones and lie outside the realms of science. Even so, the application of orderly method has led to findings which are likely to change the perception that many people have of hunting.

Before the study was carried out, it was possible to argue that views about suffering in hunted deer were subjective and open to debate. I was convinced that supporters of the Hunts were sincere in their belief that stag-hunting was not cruel. This position is, I believe, no longer tenable. Both those who hunt red deer and those who are concerned more widely with the welfare of these animals will need to take the new evidence into account.

Addendum

This article was written before the outcome of my report to the National Trust was known. The day after the report was published, the National Trust banned the hunting of red deer with hounds on their land. I was vilified by those who supported the hunting of mammals with dogs and the Stag-hunting organizations commissioned a second study by Professor Roger Harris and colleagues, hoping that this group would disprove what we had discovered. However, they obtained exactly the same physiological results as we had obtained. While they made no claim to study the welfare aspects of hunting, they concluded incorrectly that the deer ceased to run when they had exhausted their stores of carbohydrate. Later still, Professor Harris and I collaborated on a report for a Government Inquiry into the hunting of mammals with dogs. This Inquiry led eventually to an Act of Parliament that introduced radical curbs on the hunting of mammals with dogs in England and Wales.

Sir Patrick Bateson is a Fellow of the Royal Society, Emeritus Professor of Ethology (Animal Behaviour) at the University of Cambridge, and former Provost of King’s College, Cambridge. An earlier version of this essay appeared in Times Higher Education on 11 April 1997 and is used here with permission.

We humans spend a remarkable amount of time, money, and energy to benefit others, including family, friends, and strangers. Why do we do it? Do we ever care about others for their sakes and not simply for our own? Is our ultimate goal always and exclusively self-benefit, or are we capable of caring about another person’s welfare as an ultimate goal? These questions are asking about the existence of altruistic motivation in humans.

The orthodox answer to such questions, at least in Western thought, is clearly stated by La Rouchefoucauld: “The most disinterested love is, after all, but a kind of bargain, in which the dear love of our own selves always proposes to be the gainer some way or other.”

The empathy-altruism hypothesis offers a very different answer. It claims that empathic concern (other-oriented emotion felt for someone in need—sympathy, compassion, tenderness, and the like) produces altruistic motivation (a motivational state with the ultimate goal of increasing the other’s welfare). Over the past 35 years, other researchers and I have attempted to test the empathy-altruism hypothesis using laboratory experiments and have, overall, found quite strong support (for a partial review see Batson & Shaw, 1991; Batson, forthcoming provides a more complete review). Altruistic motivation does seem to be within the human repertoire. Of course, fundamental questions remain: What produces empathic concern? Can we give a plausible account of the evolution of empathy-induced altruism? What are the practical and theoretical implications if empathy-induced altruism exists?

In everyday life, empathic concern seems to be a product of (a) perception of another as in need and (b) intrinsic valuing of that other’s welfare. Contrary to what is often thought, empathic concern is not a product of perceived similarity of the other to the self. We do not simply feel for ourselves in the other. We can feel empathic concern for a wide range of others in need, even dissimilar others, as long as we value their welfare.

In terms of evolutionary history, I do not think that reciprocal altruism, inclusive fitness (kin selection), or group selection in its various forms can account for empathy-induced altruistic motivation in humans. Rather, generalized parental nurturance now seems the most likely evolutionary basis of empathic concern—even for strangers. Human parental nurturance is far more flexible and future-oriented than the parental instincts found in most—perhaps all—other mammalian species. It is need-oriented, emotion-based, and goal-directed. And it can be generalized well beyond our own children—in the case of pets, even to members of other species. If parental nurturance is the prototype for empathy-induced altruism, then the intensity of tender, empathic feeling for strangers should vary with perceived similarity to progeny, not perceived similarity to self. Is this true?

Colleagues and I sought to address this question with an experiment (Batson, Lishner, Cook, & Sawyer, 2005). Undergraduate women were asked to read a pilot article for a new feature for the Daily Kansan, the local university newspaper. The feature was called “Helping Hands,” and was to run articles in which students described their volunteer experiences in the local community. Some of the women read about a female student helping Kayla, a university student much like themselves, with rehabilitation exercises after a severely broken leg. Others read about exactly the same volunteer experience, except that Kayla was a child, a dog, or a puppy. This variation produced four experimental conditions:

Does evolution explain our behaviour? The short answer is: No. And you may well concur with that answer but ‘out there’ there is an increasing constituency of thinkers claiming quite otherwise. Along with the claims that the brain explains the mind and activity in one bit of brain or another corresponds to love, joy, conscience, the self, or whatever,  and that much of our behaviour is explained by a gene for this or a gene for that, is the increasingly popular notion that our behaviour has an evolutionary explanation. Indeed you can hardly open a newspaper without encountering some manifestation or other of ‘evolutionary psychology’.

Evolutionary psychology assumes that  human behaviour is  being shaped, indeed determined, by processes of natural selection: those modes of behaviour that favour the replication of the genome will preferentially survive. We behave as we do because we are designed to optimise the chances of our surviving long enough to replicate our genetic material. Men who sleep with a lot of women, traders who aim to maximise their returns on their investments – or at least attempt to – are simply responding to the fundamental biological imperative to make the world safe for their genes.

You may think that you chose your mate because she was kind, and witty, and shared your view of the world. Forget it: you were attracted to her because she had a waist/hip ratio (the ratio of the circumference of the waist to that of the hips) that approximated to the 0.7 – a figure which is associated with good health and fertility. As for your political or religious beliefs (if you have them), they possess you because they improve your chances of survival or that of the group to which you belong. You haven’t chosen them; they have chosen you. If you think this is somewhat tendentious, consider the  recent claim is that evolutionary psychology can explain why pink is associated with femininity and blue with masculinity. Women in prehistory were the principal gatherers of fruit and would have been sensitised to the colours of ripeness – i.e. deepening shades of pink. Men, on the other hand, would have looked for good hunting weather and sources of water – both of which are connected with blue.

Not everyone is persuaded by the terribles simplificateurs who preach the gospel of  evolutionary psychology;  and I include myself among them. The initial attractiveness of the notion that it is our genes, and not our thoughts, or our conscious agency that guide our lives is understandable. Has not Darwin demonstrated that human beings were manufactured by the same processes as gave rise to chimpanzees, sea slugs and centipedes? And are we not living now because we have bodies and behaviours that maximise our individual or collective fitness? What makes us think that human beings engaged in the manifestly biological act of choosing a mate are any different from other creatures engaged in the same activity?

Well the facts of everyday life, for a start. The enormously complex events that result in  two individuals deciding to share their lives, consisting to an important degree of a very long sequence of conversations,  has no counterpart in the pair-bonding processes even of our nearest primate kin. This seems so obvious that you may wonder why anyone gives evolutionary psychology the time of day. It is easy, however,  to persuade of the truth of this pseudo-science if we describe  animal behaviour in anthropomorphising terms and human behaviour in ‘animalomorphic’  terms. Words such as ‘mating behaviour’, ‘courtship’, and so on,  shuttle back and forth between the human world and the animal kingdom.

We can see this linguistic pincer movement closing on the gap between humans and animals at work everywhere in evolutionary psychology. Let us look at the first strategy because it is particularly effective. Indeed, we have got so used to re-describing what goes on in ordinary human life in such a way as to make it sound like what goes on in ordinary animal life, that we no longer notice ourselves doing it. Here are a couple of examples. Supposing you invite me out for a meal. Having learnt that the credit crunch has turned your house into a mound of negative equity, I choose the cheapest items on the menu and falsely declare that I am full after the main course, so as to spare you the expense of a pudding. A chimpanzee reaches out for a banana and consumes it. Evolutionary psychologists would like to say that both the chimp and I are doing similar things: we are exhibiting “feeding behaviour”. This identity of description, however, obscures huge differences between the chimp’s behaviour and mine. Here’s another example. I decide to improve my career prospects by signing up for a degree course which begins next year. I have a small child. I therefore do more baby-sitting this year in order to stockpile some tokens. Daisy the cow bumps into an electric wire and henceforth avoids that place. It could be said that both Daisy and I have been exhibiting learning behaviour. Again, I think you will agree, the difference between the two forms of behaviour is greater than the similarities.

Those who wish to obliterate the gap between humans and other beasts not only try to make human behaviour beast-like. They also describe animal behaviour anthropomorphically, making it seem to be human-like. We are all familiar with Walt Disney- like descriptions of animals that impute to them all manner of abstract or factual knowledge and institutional sentiments for which there is evidence only in human beings. This exemplifies a wider error that I have christened the Fallacy of Misplaced Explicitness that enables thinkers to speak of squids classifying the contents of the world, wasps grieving for their young and even artefacts such as thermostats making judgements.

One of the motives behind this is a feeling that, since we are animals in so many respects, unsentimental honesty requires us to say that we are just like animals in all respects. Like animals we are ejected from our mother’s bodies at birth and like animals we die of physiological failure; like animals, we eat, defaecate, copulate, fight and so on. But this is beside the point; for it does not follow that we eat, defaecate, copulate, fight. etc., like animals. It is a mistake, by the way, to look for our uniqueness solely in higher manifestations of humanity – such as religion, art, science and creativity. It concedes too much and overlooks that our uniqueness is there in every aspect of our lives: all of the biological givens are utterly transformed in us. We do not even defaecate like animals. Or not by choice, anyway. We  insist on a certain amount of privacy – often in facilities with light switches that are connected to fans driven by artifacts based in the mighty science of electromagnetism to take away the pong. And we are the only beasts who manufacture toilet paper and argue over the respective merits of different brands of it. Even human dying is profoundly different from animal dying, except at the very end, when we become more like other stricken beasts. And when it comes to mating, we are the only beasts who make love. Every seemingly animal need or appetite – for food, water, and warmth – is utterly transformed in humans. And many of our strongest appetites – for example, for acknowledgement of what we are in ourselves, for abstract knowledge and understanding – are unique to us. Only humans think about the distinctive features of their own species.

Even evolutionary psychologists can occasionally see what is in front of their noses and notice that we are a bit different. They acknowledge  that it was no mere coincidence that the organism  that saw how all organisms came about and   wrote The Origin of Species was a human being rather than, say, a chimp or a centipede. Undaunted, evolutionary psychologists look  for ways to fill the Great Ditch separate humanity from animals and human behaviour from animal behaviour. The land-fill they need to make their approach to human behaviour seem half way plausible is provided by the concept of the ‘meme’,  introduced by Richard Dawkins over thirty years ago,  but now ubiquitous, like Dawkins himself.

The meme is a notion designed to cope with the fact that human lives are filled with, and are shaped by and shape, cultural phenomena that have no counterparts in the natural word. It is supposed to be analogous to a gene. While the gene is a self-replicating unit by which biological characteristics are transmitted, the meme is a self-replicating unit by which cultural characteristics are transmitted. Dawkins gives as examples ‘tunes, catch phrases, clothes, ways of making pots or building arches’. The enormously influential philosopher Daniel Dennett believes that human consciousness is a huge complex of memes.  The key feature of memes for evolutionary psychologists is that they replicate by occupying human minds, which accept them as passively as does a brain invaded by a virus. We do not choose our memes; our memes choose us. They are advantageous to themselves but not necessarily to us; for their whole raison d’être – not only the reason  for their existence but the reason that they exist at all – is that they are able to find minds in which to replicate.

This is a desperate attempt to ‘save the appearances’ in the face of a theory – the notion that evolution determines our behaviour – that has difficulty accommodating them. Just how desperate is manifested in some of Dennett’s examples of memes: the SALT agreement, faith,  and tolerance for free speech. It is difficult to think of ‘tolerance for free speech’ as something that infests my passive mind as a unit. On the contrary, it is a principle that is argued over within and between people, and whose scope is likewise debated. It requires active, conscious assent on each occasion. Even the simpler memes – such as a tune in one’s head – are often discretionary: I bet the tunes in my head and the tunes in yours are not the same; or the same in my head from day to day or hour to hour. And many so-called memes – ideas for example -  far from being passively acquired, indeed inescapable and   unchosen as genes are, require hard work.

Meme theory, which sidelines human agency – and pictures the human mind as something between a junk yard and a lumber room – is the reduction to absurdity of evolutionary psychology. It is an example of what happens when science gives way to scientism; when evolutionary theory spawns evolutionary psychology. It is not Darwinism but Darwinitis or Darwinosis that leads us to believe that our behaviour is determined by evolution.

I am an anthropologist and primate sociobiologist who seeks to understand, step by Darwinian step, how apes could have evolved to imagine and care about what the lives of others might be like.  I believe that such questing for inter-subjective engagement laid the  foundations for significant later developments such as language and cumulative culture. My focus then is on the prequel to what became the main human feature film, worlds with symbols, words and story-telling, realms beginning to be explored by psychologists like Marc Hauser, ethnographers like Polly Wiessner and literary scholars such as Brian Boyd, Joseph Carroll, Jonathan Gottschall and Lisa Zunshine.  What follows is my take on how humans became such “other-regarding” apes.

“Nature red in tooth and claw”, “selfish genes”, and “rational actors” notwithstanding, humans are a peculiarly other-regarding, “pro-social” species. We routinely share and behave in ways that benefit others and find it pleasurable to do so. Our closest relatives among the other apes, chimpanzees and bonobos, with whom we last shared common ancestors some seven million years ago, and still share nearly 99% of  DNA sequences, also descend from highly social, manipulative ancestors and possess similar cognitive capacities, yet they are far more single-mindedly self-serving. In this respect, other apes are far more nearly “rational actors” than humans are.

Time and again, anthropologists have drawn lines in the sand dividing humans from other apes, only to see new discoveries blur those boundaries. By now every one of the Great Apes has been observed to select, prepare and use tools, crafting natural objects into sponges, umbrellas, nutcrackers, even pointed sticks for jabbing prey. Such traditions are transmitted across generations so that researchers in the new sub-discipline of “Primate Archeology” (Haslam et al. 2009) are excavating stone mortars chimpanzees used thousands of years ago.  Other apes are also born able to scan and imitate the faces of their caretakers, much as human newborns do, and they exhibit rudimentary capacities for attributing mental states to others. Chimpanzees and bonobos who exhibit considerable empathy in some contexts, also sometimes help one another (Warneken and Hare 2007) and may  occasionally share food although, in the wild they usually have to be badgered first (Fruth and Hohmann 2002). Apart from language (no one challenges human exceptionalism on this score) remaining outliers  distinguishing humans from nonhuman Great Apes mostly have to do with how much further along the continuum of other-regarding impulses humans fall.

Glossary
Alloparent – Group member other than genetic parent who helps the mother rear offspring (from Greek allo- for other than).
Allomother – Male or female group member other than the mother who helps rear offspring.  This catch-all term can include the genetic father when in the absence of DNA data no one knows who the father is.
Cooperative Breeding – Cooperative breeding refers to species with alloparental care and provisioning of young, and has evolved in roughly 9% of birds and 3% of mammals.  Shared care (without provisioning), as well as cooperative breeding (alloparental care plus provisioning) occurs in many species of monkeys and prosimians, but humans are the only great apes to rear young this way.  Note that “cooperative breeding” does not mean individuals always cooperate; a lot of competition can go on both within and between cooperatively breeding groups. SBH

Humans are far better than other apes at attributing mental states to others and appear to care much more about what others think and feel. Children as young as two years old who have not yet learned to speak are  already concerned about what others think. They are capable of both pride and embarrassment, eager to know how others perceive them. Right from an early age humans are also eager to share with others.    Every human society ever studied is characterized by food-sharing and carefully considered gift-giving. Furthermore, when a human does something nice for someone else, the  brain centers stimulated are those associated with registering  pleasurable sensations.  So what brought about this striving for inter-subjective engagement?

In retrospect, there are obvious benefits to attributing mental states to others and intention-reading.  No other apes coordinate behavior to achieve common goals the way humans do.  But Natural Selection can not favor traits simply because they might be useful and enhance fitness down the line. So how did individuals in the lineage leading to the genus Homo evolve these peculiarly other-regarding impulses? Traits like mutual tolerance, giving impulses and mental attribution had to already be there before higher levels of cooperation,  social learning, teaching, cumulative culture and above all language could develop. Furthermore, as psychiatrist Peter Hobson and  linguists like Tecumseh Fitch and Tom Givón stress,  questing for intersubjective engagement provide important motivations for developing forms of communication that go beyond the signaling of other animals — “watch out”, “there’s danger on the ground” or “food over here”. If even before they master language, children wonder what others think and feel and are capable of inventing  imaginary friends with emotional lives of their own, it is because — as  poet Daphne Merkin  puts it — we humans “long …to find our singular passions reflected in a larger pond than the selves we swim in.”

So just how on  Darwin’s earth did Natural Selection come to favor individuals incrementally better at monitoring the intentions and feelings of others? How did the peculiarly human quest for inter-subjective engagement get started? An important first step is to break down nonhuman-human ape differences into their component parts so as to better understand exactly what traits are involved.

In the most ambitious comparative study to date undertaken at the Max Planck Institute for Evolutionary Anthropology, 106 chimpanzees of all ages, 32 orang utans and 105 two and  a half year old children were subjected to a specially designed  battery of tests allowing Michael Tomasello’s team to compare their sociocognitive capacities (Herrmann et al. 2007). In terms of spatial memory and object permanence, or the ability to discriminate quantities or understand causality, chimps and orangs perform in the same range as human toddlers. The big differences are in the social realm where children test significantly better at learning how to solve a problem from watching a demonstrator. Children are also better at understanding communicative cues like pointing, and better at attributing mental states to others so as to understand what they are intending or trying to accomplish – what psychologists call Theory of Mind.  So to explain nonhuman versus human ape differences, we need to identify selection pressures favoring increases in mutual tolerance, social communication and intention-reading.

To my mind, existing explanations for other-regarding impulses –such as the need for in-group solidarity in the interests of defeating out-group enemies — leave unaddressed these initial steps towards prosociality and fail to explain why other apes such as  chimpanzees did not spend the last seven million years becoming more cooperative as well. This is one reason why in Mothers and Others I focus on one of the most obvious, albeit too often overlooked, differences between humans and other apes, the peculiar way that bipedal apes living in small-scale hunter-gatherer societies rear young.

Among chimps, gorillas, bonobos and orangs, mothers are extraordinarily possessive of new babies, in fur-to-fur, body-to-body contact, not out of touch even for a moment, for the first six months or so of life. Infants are suckled for up to seven years. Once weaned however, nonhuman ape  youngsters  provision themselves. By contrast, human babies born in traditional societies are passed around among other group members from the first day of life, and in some groups are held by allomothers (typically kin — fathers, older sibs, aunts, cousins or  grandmothers) for much of the time, even suckled if the allomother holding them is lactating. Within months, long before children are weaned, infants are fed as well by allomothers who deliver soft or pre-masticated  foods,  kiss-feeding the baby by pushing the mash in with their tongues.  Supplementation of children’s diets continues for many years.

Even though human infants are bigger at birth and take nearly two decades to mature, allomaternal provisioning means that mothers can wean babies sooner than other apes. With toddlers buffered from starvation, mothers breed again sooner, potentially increasing lifetime reproductive success but also forcing mothers to become even more dependent on alloparental assistance. In Mother Nature I proposed that increasingly contingent maternal commitment produced the curious combination of passionate maternal love with ambivalence evident in human mothers.  Although rarely seen in primates with exclusive maternal care, high levels of  ambivalence in mothers short on social support is common in other cooperatively breeding primates. Humankind’s deep legacy of cooperative child-rearing also had implications for the cognitive and emotional development of youngsters growing up dependent upon allomothers (the main focus of Mothers and Others) as well as  implications for allomothers (Burkart et al. in press).

None of us has a machine to go back in time and observe how infants in the Pleistocene were reared. But what we do have is a growing body of information about exactly who cares for immatures not only among other primates, but also among humans  living as hunters and gatherers. Furthermore, it is increasingly clear that in traditional societies with high child mortality, alloparental care and provisioning is not only useful, but critical for child survival. Homo sapiens species could not have evolved without it.

Space does not permit here a review of what sociobiologists are learning about the demographic implications of cooperative breeding and the conditions under which it is likely to evolve — particularly in highly social species with helpless young confronting environmental challenges such as unpredictable resources.  Nor does space permit me to review here information from neuroscience, endocrinology and the emerging field of comparative infant development relevant to understanding what the psychological implications of multiple caretakers were likely to be.  But here are a few of the highlights. The presence of a supportive kinswomen like a grandmother is correlated not only with increased child survival, but also with increased maternal sensitivity to infant needs, and infants growing up with nearby grandmothers are more emotionally secure and develop cognitively at a faster pace. Having older siblings around also enhances development of social skills.  As cognitive psychologists quip, “Theory of Mind is contagious” you catch it from older caretakers. Furthermore, forming attachments to multiple caretakers enhances perspective-taking and conditions children to integrate different perspectives.  Even though historians of the family, social workers and psychologists have long known about cognitive and emotional advantages from growing up in an extended family, only recently have evolutionists begun to recognize that such alloparental assistance would have been essential for our ancestors to rear surviving young.

Now consider information from other primates that cooperatively rear young.  Experiments with tamarin monkeys by Marc Hauser’s team at Harvard, and with marmosets by Judith Burkart and Carel van Schaik at the University of Zurich, reveal that these tiny-brained cooperatively breeding monkeys are far readier to pull a rope delivering food to others than are large-brained chimpanzees in comparable tests. These generous “other-regarding” impulses come into play even if the recipient is  unrelated. In the wild as well, cooperatively breeding tamarins and marmosets tend to be mutually  tolerant  and helpful to one another, as well as unusually good at coordinating their behavior in subsistence tasks.  Among these tiny-brained and distant primate relations, relations, the father (or another male the mother mated with) carries the babies (usually twins) much of the time, except when the mother is nursing them.  Allomothers also provision the babies around the time of weaning such that in some species of tamarins, 90% of the infants’ first solid food comes from alloparents. Chronic food-sharing spills over into generosity in other realms as well. Adults routinely vocalize to call the youngsters’ attention to novel, or particularly palatable, food items while intervening to prevent them from eating toxic foods — behavior that comes close to teaching (Rapaport and Brown 2008; Burkart et al. in press).

In Mothers and Others I summarize such evidence and invite readers to join me in a well-documented thought experiment. Take a primate with the cognitive and manipulative potentials and rudimentary empathy and  Theory of Mind typical of all  Great Apes, and rear that creature in a novel developmental context where his  mother’s commitment is contingent on how much social support she has and she and her infant depend on care and provisioning from multiple caretakers. The resulting phenotype will be a youngster adept at perspective-taking, far more so than any other ape under natural conditions would be. Then subject this novel ape phenotype to novel selection pressures such that infants best at monitoring the mental and emotional states and intentions of others, and also best at learning from them, are going to be those best cared for and best fed. This then leads to directional selection favoring traits like enhanced mutual tolerance, social learning, social communication and perspective taking — precisely the traits that comparisons between humans and other apes require us to explain.

No one knows when cooperative breeding got started in the hominin line. Evolutionary anthropologists Kristen Hawkes and James O’Connell have hypothesized that our ancestors began relying on alloparental care and provisioning by the beginning of the Pleistocene, 1.8 million years ago, and I concur with their reading of the paleontological evidence. If correct, this means that long before behaviorally modern humans capable of symbolic thought, art and language emerged, and even before big brained anatomically modern Homo sapiens with fully sapient 1350 cubic centimeter brains in the last 200,000 years,  these cooperatively breeding ancestors would have been well on their way to  “emotional modernity”. Emotionally modern apes would have grown up keeping track of others and  inordinately interested in the feelings and lives of others, even those out of sight, or far away.

Acknowledgements: This essay is based on a lecture originally presented at the Darwin Festival, University of Cambridge, July 5-9, 2009.

Selected References, including recent additions since Mothers and Others

• Burkart, J.; E. Fehr; C. Efferson; and C. van Schaik.  2007.  Other-regarding preferences in a nonhuman primate: Common marmosets provision food altruistically. PNAS 104: 19762-66.
• Burkart, J.; S. Hrdy; and C. van Schaik. 2009 (in press). Cooperative breeding and human cognitive evolution. Evolutionary Anthropology.
• Fruth, B. and G. Hohmann. 2002.  How bonobos handle hunts and harvests: Why share food?  In Behavioural Diversity in Chimpanzees and Bonobos, edited by C. Boesch, G. Hohmann and L. Marchant. Cambridge: Cambridge University Press, pp. 231-243.
• Haslam, M. et al. 2009. Primate archeology. Nature 460, 339-344.
• Hauser, M.; M.K.Chen; F. Chen and E. Chuang. 2003. Give unto others: Genetically unrelated cotton-top tamarins preferentially give food to those who altruistically give food back. Proc. Roy. Soc. London (Series B) 270:2363-70.
• Hawkes, K.; J.F. O’Connell; N.G. Blurton Jones, et al. 1998 Grandmothering, menopause and the evolution of human life histories.  PNAS 95:1336-39.
• Herrmann, E.; J. Call; M.V. Hernandez-Lloreda J.; B. Hare and M. Tomasello. 2007 Humans have evolved specialized skills of social cognition: The cultural intelligence hypothesis. Science 317:1360-66.
• Hrdy, S.B. 1999.  Mother Nature.  New York: Pantheon.
• Hrdy, S.B. 2005.  Evolutionary context of human development: The cooperative breeding model. In:  Attachment and Bonding, a new synthesis, edited by C.S. Carter et al. Cambridge: M.I.T. Press, pp. 9-32.
• Rapaport, L.G.  and G.R. Brown   2008. Social influences on foraging behavior in young primates: learning what, where and how to eat.  Evolutionary Anthropology 17: 189-201.
• Sear, R. and R. Mace. 2008. Who keeps children alive? A review of the effects of kin on child survival.  Evolution and Human Behavior 29:1-18.