Monday, January 6, 2014

Intelligence in Societies: Mainly in Insects

Intelligence in Societies: Mainly in Insects

Extended Abstract
Concepts of intelligence
Formally defining intelligence.
Constraints on intelligent systems.
Intelligence and apparent teleology
Intelligence and Information Processing
Intelligence in non-atomic systems
Intelligence and communication
Intelligence and subsystems
Intercommunication between component modules of intelligent systems.
Learning and intelligence
Coupling between subsystems or components of intelligent systems
In loosely coupled systems, coupling may be behavioural, even casual.
Examples of tighter coupling include compulsory parasitism or symbiosis
Parasitic or symbiotic vector relationships
Physiologically profound symbiotic relationships
Colonial sociable communities
Familial communities
Colonial eusocial communities
The colonial nature of multicellular or physically crowded communities.
Intelligence, language, and abstraction; dances and semiotics
Democracy and didactics in communal intelligence
Intelligence and subjective consciousness
Stages and aspects of sociality & associated intelligence
Hypothetically socially independent, parthenogenic organisms.
Sexually reproducing species
Mutually interdependent species
Rudimentary intraspecific tolerance
Interspecific and intraspecific flocking
Aggression inhibitors (Tolerance releasers?)
Episodic assembly
Rearing care and defensive behaviour releasers
The evolution of matching signal conventions
Eusocial colonies and familial groups
Systematic reaction to characteristics of the environment
Establishment of the colony
Maintenance and management of the colony
Conflicts of interests
Behavioural pathology in communities
Intelligence of and within colonies
Intelligence of individuals or castes within colonies
Intelligence of colonies
Culture and intelligence: elephants and apes, orchestras and Selenites
Evolution of intelligence

Extended Abstract

As compared to the behaviour of their individual members, behaviour patterns of communities raise questions concerning the very mechanisms and nature of intelligence.  If the concept of intelligence comprises more than one type of component then it is simplistic to reduce one’s view of intelligence to limited aspects such as at one extreme, information processing and storage, or at the other, subjective consciousness and purposiveness.  Currently it is not clear how many effects should be lumped together as evidence for “intelligence” in general. 
The concept of intelligence of communities is of particular interest because communities present a range of outgroups that we may contrast to intelligence in individuals with variously modular, unitary brains.  To object that one cannot compare intelligence in a unitary brain, with behaviour of a community of communicating individuals, is not cogent; the brain’s intelligence depends after all on the behaviour of communicating individual neurons. 
The most striking examples of putatively intelligent communities are eusocial insects, but other types of communities suggest wider ranges of underlying principles and lines of development.  Concentration on integrated brains or too narrow a range of types of community, may obscure seminal questions.
Brains are modular, largely in ways that reflect the components of their behaviour, and it is not clear how many sub-modules are the sites of intelligences in themselves.  When modules communicate, do they form a new functional intelligence (a new “mind”)?  If so, then how?  To what extent and in what way might modules retain identities when they form a team of minds in the one brain?  Our main difficulty in resolving such puzzles is that we lack physical measures by which to detect or quantify what constitutes a mind, or to distinguish between single minds, structures of minds, or fractions of minds.  Without measurable criteria discussion of the concept of intelligence reduces to simplistic information theory, observation, introspection, and thought experiment. 
Other information processing entities than unitary brains comprise recursive sub-modules.  Examples of such entities include communities and possibly entire species throughout their existence, with modules comprising individuals, groups, and super-groups.  Unlike brain modules, sub-communities are not connected physically by synapse or neuron, but they might behave intelligently in ways reminiscent of Searle’s Chinese room, where no component recognises the operation in which it plays a role, any more than a neuron in a human brain might do.  Conversely other types of communities might show little intelligence, but include at least some intelligent members who exploit their communities to achieve personal objectives.
Such examples show how community intelligence need not obviously be commensurable with that of particular members.  Even the intelligence of individual members, or classes of members, such as different castes in insects, or different professions in humans, might not be usefully commensurable.  Intelligence has too many dimensions for adequate one-dimensional measurement. 
That either communities or brains are modular does not imply that their respective modules are functionally analogous.  Communities need not have either anatomical or functional analogues to brains.  To faster, but not necessarily more complex minds the intelligence of long-lived communities or species may be imperceptible or qualitatively different.
Apart from community intelligence, aspects of the intelligence of insects differ from that of birds and mammals, including eusocial mammals.  Play activity seems practically non-existent, and mental generalisation is of a very low degree.  Insects are poor at passing on information except of special types, such as information concerning alarm or food.  In some ways the stereotyped behaviour patterns of individual social insects are reminiscent of the limited behaviour repertoire of neurons. 
How does one measure the relative intelligence of a system of components with stereotyped behaviour, as opposed to systems with putatively subjectively conscious components?  It is a messy business to compare the intelligence of a termite nest with that of a human community.
Politics offers one class of comparison between, if not actual measurement of, the intelligence of the individual and the community when the intelligence of individual members permits the establishment of rulers or mentors.  Political structures are of particular importance in the present context in that the field of politics is where individual intelligence and whatever passes for the intelligence of the community, affect each other directly. 


Read not to contradict and confute; nor to believe and take for granted;
nor to find talk and discourse; but to weigh and consider. Some books
are to be tasted, others to be swallowed, and some few to be chewed
and digested: that is, some books are to be read only in parts, others
to be read, but not curiously, and some few to be read wholly, and
with diligence and attention. 
Francis Bacon
This is not a balanced treatise on any aspects of intelligence in any useful sense of the word.  At most I present preliminary informal reading material to support a discussion on the subject of apparent intelligence as manifested in communities of organisms.  However, my only apologies are that the essay is too short by an order of magnitude, that this adversely affects its coherence, and that its construction has been so informal that I have not produced any supporting bibliography.  My biggest disappointment is that I have been unable to structure all the subject matter into a coherent conclusion or unifying principle.  As I point out, I am doubtful that the concept of intelligence is monolithic.  I am certain that we are not anywhere near being able to demonstrate the truth or error of this opinion as yet. 
Another point of irritation to some readers might be the way in which I speak as though colonies were comparable to individuals.  I neither do so purely accidentally, nor am I regenerate in the face of accusations of argument by false analogy; the human body – any metazoan body - is a colony of cells.  For our purposes it generally is a clone of cells, though that is a far looser definition than some might think; it also is arguably inaccurate.  Some of the attributes of a colony of separate organisms are fundamentally different from the attributes of a body, but some are not.  Both those that are logically analogous and those that are alien, are relevant to the points made in this essay. 
Please note that I am not being simplistic in this. I am well aware that to speak of an evolutionary strategy for example, is not the same as speaking of a conscious strategy in say, a chess player. I similarly realise that the decision of a swarm is not clearly the same as the decision of a brain.  Swarm behaviour however, certainly includes episodes that functionally correspond to decisions. 
Similarly, I do urge that readers remain alert for functionally valid analogies among the intelligence of species as one type of unit, colonies as another, and unitary brains as a third. All three are systems that use information and its processing in meeting the challenges of the environment. Even positivistically we are far from any cogent argument against dealing with each of the three as being intelligent in its own terms. 
Given the disputable nature of a lot of the subject matter, I expect and welcome discussion, argument and criticism.  As for the document content, anyone is free to use it in any good faith connection as they see fit. 
The question of what it means to speak of intelligence in systems other than individual, possibly unitary, brains (or machines) is of key importance if we are to make sense of the subject of intelligence at all.  The very idea of the intelligence of a community raises questions as fundamental as those raised by scripting or computer modelling of intelligent behaviour.  The intelligence of an insect colony is just one such dimension of study. 
It is tempting to argue that it is meaningless to speak of the intelligence of a group as anything but an analogy, and a foggy analogy at that, but the argument is treacherous at best, and in  my opinion is simply invalid.  Our brains are groups of neurons, but I have never heard any cogent argument to demonstrate that each neuron has a subjective consciousness.  It is hard even to argue that it has any intelligence at all, as opposed to information processing capacity. 
To argue that in the brain neurons are not a group, but are physically attached to each other, is neither valid, nor unconditionally relevant.  Much of the brain’s connectivity between either different parts of itself, or between itself and the rest of the body, is not at all by contact, nor even via neurons, but by chemical signals or even by other media.  It is perfectly possible (though a good deal more tricky than science fiction authors might suggest) to replace a synaptic connection with an artificial (say, electronic) intermediary, and without obviously changing the effect on the receiving neuron. 
After all, ever since Aristotle, it has been a commonplace that the whole is more than the sum of its parts.  We do not assume that because a car is made partly of bolts, partly of pistons, and partly of paint and the like, that it should behave partly like a piston, partly like paint and so on.  In fact, it behaves very like a car, even though within it there are pistons behaving like pistons and bolts behaving like bolts.  What is more, even though the headrests and windscreen wipers have little to do with the engine or the wheels, we would not easily be satisfied with a car in which the headrests and windscreen wipers did not work. 
To labour analogy still further, any organisational man will know that a number of ships is not the same as a fleet, nor a number of people the same as a company, even if no two elements in fleet or company ever make physical contact.  The organised aggregate has abilities that the individual lacks and generates liabilities that the individuals escape.  To treat the organisation in the same way as one treats an individual, or even to treat a small organisation in the same way as one treats a large organisation, or the other way round, is to tempt disaster. 
Furthermore, simply by rearranging the parts, it is possible to change the nature of such a structure radically, whether it is a device like a car, or an organisation like a company or a fleet.  The change might be favourable, irrelevant or dreadful, but the fact is that a dismantled or reassembled car can be a scrap heap, a stock of spares, a lounge suite, or a power plant, without there being a single component unaccounted for, and without there being any significant argument about whether what one now is looking at still is a car.  This crushingly demonstrates that Aristotle had a point about wholes and sums of parts, and as far as we know, he never even saw a car.  When one deals with kits of interchangeable parts, like Meccano or Lego, the concept becomes even more striking. 
Exactly analogously, a bee is not just a number of cells, and a colony is not just a number of bees, not even if one is a queen and all the others workers or larvae.  And as with Meccano or Lego, when one does have an insect colony, it is not just anyone who can tell the components apart.  Even in a large human company, not every member could put a name to every colleague, nor even recognise each one, and yet, not many would argue that each time a member dies, leaves, or joins the company, it is a new and different company. It might sometimes be, but not often.  
Other dimensions of intelligence include the intelligence of an entire species, as opposed to any member or community of that species, or of ecological interactions between species.  There is a lot to be said for the value of charting the direction of information flow and the nature of the information processing, to characterise the nature of the respective types of intelligence.  Such flow is far from the whole story, but it is better than hand-waving about what impresses one person and fails to impress another as defining intelligence. 
In this essay I do not explore the ramifications of the Gaia concept, interesting though they are, but I remark in passing that I do not regard it as practically or intellectually cogent, certainly not on this planet as things are at present. And the concept of information flow is part of the reason for my scepticism.   
All the populations that we discuss, whether species, colonies or individuals, behave in ways that from our point of view amount to algorithmic or at least systematic reaction to environmental situations.  How is one to discriminate between such behaviour and intelligence?   How is one to distinguish between human cells and independently moving colonies of driver ants or of myxamoebae in a slime mould “slug”?  Is the distinction the fact that our body cells stay together?  That holds no water.  We shed skin cells by the million daily, far more than the driver ant colony sheds non-reproductive members; some types of leukocytes move in and out of our mucosa all the time; blood and lymph cells range freely almost throughout our bodies. 
That same question becomes even trickier when we reflect that every distinct, non-trivial brain that we have been in a position to investigate, has different regions that deal, in more or less specialist ways, with more or less distinct functions.  In fact, there are good grounds for suspicion that brains are necessarily strongly modular in their architecture and function.  The analogy to eusocial castes is hard to refute. 
Brain modularity is of at least four major types, all of them of anatomical and functional importance.  The most fundamental type arises from the nature of the creatures with the most advanced brains: the vertebrates, arthropods, and more debatably, certain molluscs.  This is the segmented nature of the central nervous system.  Ancestral organisms originally had nodes of the central nervous system in each segment of the body.  Each module, each bodily segment, had its own segment of the central nervous system.  Their interconnection was necessary for overall function and control, but they had local function and control as well.  Their interaction in such coordinated functions as the walking of millipedes has impressed generations of biologists. 
The second type of modularity originated from a sort of reversal of the original segmentation process, the combination of small, repetitive modules into fewer, larger modules.  The brains of both humans and bees consist of multiple segments of their central nervous systems assembled into a more closely integrated structure. 
Vertebrates accordingly have a cranial brain and a spinal cord.  The two components are not cleanly separate, but each does have its own architecture and range of functions.  The brain evolved originally as a local enlargement of the spinal cord, associated with the location of particular sense organs, mainly sight, hearing, and certain classes of chemoception. 
The third type of modularity is elaboration of the functional enlargements of primitive segments of the central nervous system.  The major regions of vertebrate and arthropod brains are outgrowths of the segments that constituted the development of the front end of the spinal cord into a brain. In the vertebrate brain the most conspicuous enlargements are the cerebrum, the tectum, and the cerebellum. 
Furthermore, palaeontological evidence suggests that among some of the larger dinosaurs, most notoriously in Diplodocus, there was a secondary enlargement, a node of the spinal cord in the lumbar region.  This is consistent with the idea of local enlargements of the central nervous system as adaptations to meet local needs for control.   A possibly more significant extant example of distributed control is in the tentacles of cephalopods.  The muscular and sensory control are so elaborate that if the brain of the octopus had to control the tentacles directly, the necessary nervous connections would exceed the mass of the rest of the tentacle.  Instead, local modules control simple functions and sensory data, communicating with the brain only in terms of summarised data. 
At first sight this might seem alien, but it is not qualitatively different from the way the nervous system in the retina digests visual data before passing it back for secondary processing in other areas of the brain. 
However, note the difference between the function of the modularity of local nervous control in the cephalopod tentacle, and the spinal node of Diplodocus.  The latter almost certainly was necessary to overcome the problem of controlling hindquarters at such a distance from the brain as to cause disabling transmission delay.  In modern cephalopods, especially the largest squids, there is a similar problem, in that co-ordination of the control of the mantle is important, but that it may be so large that nervous controls signals to the various parts would arrive out of synchronisation. 
In this case the correction is applied by varying the thickness of the axons that carry the control signals to the various parts of the mantle.  The speed of the impulses varies accordingly, re-establishing the necessary synchronisation. 
These examples of contrasting problems focus attention on the relevance or otherwise, of the costs of communication between processing units, and how thy might affect the unity of the mental system.  
The fourth form of modularity is most familiar in the cerebral hemispheres of mammals.  Here we find that there are many functions that according to received wisdom are located in particular regions of the brain.  Arguably the most spectacular examples are various speech and vision processing functions.  However, we now know that such locations are far less precise than once was thought, and from person to person they vary noticeably in placement and extent.  In short, such modules are not as clearly anatomically defined as the first three types. 
Insect brains too are segmentally modular.  In fact, they are more clearly divided into parts above and below the digestive tract, than any division in vertebrate brains.  Various parts of their brains produce outgrowths that may be more or less conspicuous depending on their respective functions.  In this they resemble vertebrates, and it seems pretty clear that in every type of nervous system we know, the preferential growth of particular brain regions reflect the demands of storage and processing functions.  It also seems to me a reasonable speculation that in all the more elaborate brains, there are minor functional modules within large nodes, that are not clearly reflected in the gross anatomy.  The way in which zones of the brains of say bees, swell or subside at various stages of their lives, suggests that such anatomically indistinct regions have functionally modular roles, much as in human brains. 
There also is room for analogy between brains that differ in such ways, and communities that have larger or smaller populations of various specialist members.  Such differences affect both the personality of such communities and their skill bases.  
Apart from the problems of defining intelligence or intelligent systems, there are problems concerning constraints on the validity of the venerable analogy between colonies and organisms.  As people in this field use the various terms, there not only are problems of fuzziness such as may be dealt with in fuzzy logic, but also problems arising from false analogy and conflation of disparate, poorly defined concepts.  Informal writers for example have compared the queen of a colony of bees or ants with the brain, an analogy that is neither functionally valid nor conceptually useful. 
Other problems arise where we lack the basic techniques for scientific investigation, let alone quantification, of particular effects (for example, subjective consciousness).  Here we still lack fundamental formulae corresponding for example to F=MA in physics. 
Long before we could claim to have solved such puzzles we would have to deal with the questions arising from for instance, John. R. Searle’s 1980 “Chinese Room” challenge, as described in his essay “Minds, brains, and programs”.  Personally I reject Searle’s views of the impossibility of machine intelligence equivalent to biological intelligence, but I also reject the naïve AI claim (I do not like the term “strong AI” in this sense) that related examples of scripting are essentially equivalent to, and accordingly explain, the activity of human minds in dealing with, understanding, and solving such challenges. 
One of the major ways in which eusocial communities present assistance and challenges of their own in understanding such systems, is that they are models strongly reminiscent of the Chinese room, but contrast with the human brain in that their modules are physically far less closely interdependent, and that none of their members has the independent intelligence to substitute for the entire system.  (Remember Diplodocus and the cephalopod innervation as similarly internally interdependent examples!)  In contrast, either a single human speaker of Chinese and English, or a small team, could substitute for the Chinese room. 
Such points deserve emphasis because many people are over-reliant on simplistic analogy.  They will compare a particular part of a colony, or even the entire population of the colony, with a brain, commonly even with a human brain in particular.  In fact many metazoa exhibiting rudimentary but effective intelligent behaviour do not have brains at all; some of them barely have nerves worthy of the term, let alone definite central nervous systems. 
To argue that a termite colony is not intelligent in any meaningful sense because it has no definite anatomical analogue say, to the cerebrum or brain stem, would be to miss the point.  One cannot cogently dismiss the possibility of intelligence in anticipation by defining it in terms of familiar structures and pointing out, however validly, that the subject under inspection has no structure that conforms to the definition of those terms. 
In fact apart from pre-judgement on anatomical grounds one also needs to be very cautious about arguments along the lines that such a population has no functional analogue of particular mental or brain structures.  Apart from the trap of confusing anatomical structures with structures of intelligence, there is the trap of limiting our definition of intelligence to formal structures directly equivalent to those we recognise in our own mental makeup. 
For one thing we know only a limited amount about our own mental makeup and it is philosophically questionable in principle how comprehensively we ever can know it.  This makes it hard to recognise equivalent mental formalisms in alien isomorphisms.  What is more, there is no reason to suppose that our mental modelling of our selves and our world must be or could be comprehensive in certain important senses.  For example, we cannot show that any deeply different information handling or control mechanisms for functionally dealing with the world must be invalid or meaningless or not somehow mental.  Even if such chauvinism were justified to some extent, it would not follow that the alien structures were not of similar importance in their own terms and of value as outgroups in the study of our own mental structures.   
For purposes of recognising and characterising intelligence in taxonomically alien systems, we are reduced to searching for intelligence that we can recognise empirically, that is to say, by its behaviour.  As a rule it would make most sense to look for behaviour that:
·       Has selective value, evolutionarily speaking, or could reasonably be argued to have had selective value at some time.  (The fact that it might be currently counter-selective, say because of climatic change or conflict with recent immigrants, such as humans, is at best contingently relevant.)
·       Has homeostatic function.  This is closely related to the previous point, but is more specifically relevant to such things as food and physiological control. 
·       Effectively meets the challenges of contingency.  Such challenges need not be external circumstances, though those are the most obvious; contingencies within the organism or colony are just as important, and often more complex.  Examples include hormonal and neurosecretory feedback control.  The fact that there is no recognisable consciousness dealing with the challenges does not mean that the behaviour lacks any component of intelligence.  
These criteria certainly are far too limited, but they do exclude a lot of processes that otherwise might confuse the issue. 
Apart from examples that in our experience are generally biological, we are familiar with programmed behaviour in inorganic systems, such that if we observed it in a biological system, we should readily class it as intelligent to some degree or in some sense.  In strict terms programming a machine to pass the Turing test still is beyond us, but at the same time it also is too limited a test in the context of this discussion.  Human intelligence is not the only form of intelligence we should recognise. 
Such examples of empirically intelligent behaviour in programmed systems that are not biological organisms, may or may not in specific cases explain aspects of algorithms that the biological brain might use, but even if they do, it is by no means clear that they describe the nature of the mind that solves the problem and implements the solution.  They do not even prove that the brain uses the same algorithm.  In fact, even within the same brain, investigations of human brain processes suggest that the original solution of a problem, as developed by a naïve brain, might not closely resemble the practised solution of an experienced brain. 
Conversely, if humans achieved some sort of brain to brain communication as direct as the internal neuron to neuron communication, what sort of group intelligence would emerge?  And would the individual brains lose their individual awareness of being?  Whether they did or not, would they individually be aware of any global consciousness of which they were components?  One assumes that they could not be fully aware of it; if they could, then that would imply that the single brain could compass a larger meta-brain and it would seem to follow that building the meta-brain would add nothing to the capabilities of the system. 
Possibly however, such a single brain might be aware of its meta-brain in something like the way that a blind man feeling the foot of an elephant, may be well aware that the foot is not the elephant, nor perhaps, the only foot of the elephant, even if he could not conceive the full splendour of the elephant, tail behind and trunk in front. 
And if the entire connected system developed a subjective awareness, would it be aware of the awareness of individual brains within the system?  And what degree of interconnection is necessary to generate a such a consciousness?  Must it be neuron-to-neuron?  If not, tactile, pheromonal or vocal communication might suffice, and then we do not know whether insect, or even human communities have awarenesses outside those of the individual community members. 
These questions are non-trivial, but not at present based on concepts precise enough for us to pose them meaningfully. 
Another difficulty is so great and so poorly defined that I hardly mention it and certainly cannot discuss it cogently, or even comprehensibly.  Consider such examples of intelligent behaviour as we see in programmed systems, in eusocial colonies, or even in humans as perceived by their fellows. They get nowhere near the problem of explaining, much less explaining away, the nature or function of subjective consciousness, nor its relationship to functional intelligence, nor yet even whether or how subjective consciousness is necessary or relevant to functional intelligence. 
We simply do not know whether consciousness in this sense is important, or why, or whether it is simply a passive by-product of the way some part of our conscious brain works, or even whether other information processing systems generate similar internal subjective effects. 
Conversely we cannot argue that, as I at one time suspected, subjective consciousness is mentally irrelevant, just because, as I still suspect, it is a passive emergent attribute of a working brain.  A friend of mine once pointed out that if we have such a consciousness and are aware of it, then that datum of the awareness constitutes a material difference between two brains that otherwise would be functionally identical, particularly in their external behaviour.  This is more important than the academic quibble it might seem.  Such internal awareness might be vital to certain classes of maintenance and modification of the human mind by operation on itself. 
These problems of definition go beyond the problems of meaningful characterisation of a set, such as were discussed by Wittgenstein in his “Philosophical Investigations” (roughly in paragraphs 59 to 100) or Hofstadter in “Metamagical Themas”, topic “Variations on a Theme as the Crux of Creativity”.  Not only would it require Wittgenstein’s “family resemblances” to define some of the sets we discuss, but some of the sets would justify subdivision on the basis of the disparate natures of their elements, which of course, we know hardly, if at all. 
Firstly, no one has yet managed to devise even a prospect of a test for subjective consciousness, qualia, or what Searle calls “intentionality”, though other authors use the term apparently as interchangeable with intensionality.  Some prominent writers even claim that such things as subjective consciousness are obvious or logically necessary, but that certainly is not obvious to everyone, nor has its logical necessity been shown to be logically necessary.  Some obsessive empiricists stridently deny the existence of subjective consciousness, on the basis that we cannot define it in any way that is independently empirically verifiable or falsifiable.  This view I reject, pending proof, not that we cannot define it at present, but that we cannot in principle ever define it.  The fact that stone-age man could not define magnetism did not mean that nothing existed that caused two lodestones to move together or apart, even though not all stones attracted or repelled each other all the time. 
Some respected authorities (not all of them authorities on information theory or biology) go even further and dismiss the very concept of subjective consciousness as an illusion.  I shall not in this essay waste space on the question of what my subjective consciousness might be an illusion of, and to what it is an illusion, and what it might mean for something to appear an illusion to something that is not subjectively conscious, and why it should matter that it is an illusion, given that the illusion is concrete enough to affect the brain’s activity physically, as I mentioned my friend pointing out.  Personally I find the apparent intrinsic self-reference far less acceptable than the idea that most people choke on, namely the perceived indignity that it is conceivable that we might not really exist as aware minds, but only seem to our unaware minds to do so.  
Nor do I personally remember ever writing a program that gave any evidence of experiencing such an illusion. 
Some insist that the effects of subjective consciousness certainly are real, but are limited to biological structures; others contemptuously dismiss the idea that inanimate systems might lack similar internal processes, but as I see it, none of these groups have achieved anything beyond proof by assertion. 
Most people assume implicitly that for our purposes effectively all humans have both subjective consciousness and intentionality, but we base the assumption on subjective analogy from introspection, plus generalisation on behavioural and anatomical grounds, neither of which has been demonstrated to be inseparable from subjective consciousness.  The existentialist problem of our inability to demonstrate personality in other parties than ourselves remains intact, even without invoking sterile or meaningless solipsism.  We have no strong evidence that behind the eyes of a human, there are more dimensions of consciousness than behind a fine work of art or the screen of a computer that passes the Turing test. 
After all, though that is not the way the author typically puts it, a favourite theme in human literature, is the inability of people to demonstrate their intelligence to each other.  In essence they retail failures of inverted Turing tests.  Joseph Conrad’s short story, “Amy Foster”, is a particularly direct example, but the theme is ubiquitous and for example it recurs in the story "I only came to use the phone" in "Strange Pilgrims", an anthology of short stories by Gabriel Garcia Marquez.  
Far more importantly, as I have hinted already, we do not even know how many individual subjective consciousnesses exist in any intact, functional human brain, nor can we count them even in our own respective brains, nor can we tell how it feels when they split or combine.  Investigations of split brains and stroke victims have demonstrated the relevance of such questions, but the questions go a lot further than that.  We have little idea of whether the apparently “unconscious” or “subconscious” functions of our brains are in fact without subjective consciousnesses of their own, and I have no idea of how we can find out.  But it seems to me that such questions of whether we have swarms of consciousnesses in our skulls are very relevant to the idea of the swarm or sub-swarm consciousness among colonial animals. 
Conversely the Turing criterion, for what it is worth, stands firm.  How many kinds of (or aspects of) intelligence are involved here, I cannot say, but if, on objective, existing criteria, the behaviour of a system is logically indistinguishable from intelligence, then we must find new, objective, criteria before we deny that that behaviour comprises intelligence in any relevant sense.  Searlely denial of intelligence in a particular system just on the grounds that one has not identified its nature or its roots, ranks with Bouilland (Bouillaud?), the nineteenth century French doctor who embarrassed his colleagues by publicly denying that the phonograph was anything but a ventriloquist’s trick.  Whereas Bouilland could be corrected by friends who were in a position to demonstrate principles and working devices, we are not yet in an equally comfortable a position relative to the concept of intelligence.  Even so, proof by denial is not cogent until we are in a position to define what it is that we are denying. 
Now in this discussion, when we examine the putative intelligence of communities, I shall not ignore the presumable differences between “subjectively conscious intelligence” and objectively intelligent behaviour, but for lack of a means of distinction, I often shall treat them in the same way except where the putative difference turns out to be relevant. 
Conversely, when on the Turing principle one facilely accepts every arguably intelligent behaviour of a system as evidence of intelligence, serious difficulties arise on every hand.  If a colony reacts intelligently to deal with internal and external challenges, even though the behaviour of its members is demonstrably stereotyped, then why is a computer not intelligent when it controls a building or a super-critically configured aircraft?  It controls things in much the way that a human would, sometimes far better than any human, even though it has not a single clearly intelligent component.  And if we apply this criterion to a computer, then why not to a thermostat or hygrostat, or even a spring-loaded mousetrap? 
Why not in general to any system that displays feedback control? 
Accordingly I both decline to define the concept of intelligence rigorously, and I insist on using the term largely as though it were meaningful and monolithic.  I realise that this is unacceptable in the long term, but it is no worse than doctors who discussed the nature of disease in the days before human physiology or the germ theory were generally understood, and hence before there could be any deeply meaningful classification of disease.  Had they refused to discuss such matters before they achieved fairly advanced understanding of underlying mechanisms, they would have avoided talking a lot of nonsense, but they also would have made no progress in definition or treatment of disease, and we probably would have had no proper understanding of disease today. 
So I insist on continuing to speak of intelligence with no more fundamental knowledge or justification than a medieval doctor basing his theories of disease on the four humours.  

Concepts of intelligence

                                       . . .  On a huge hill,
cragged and steep, Truth stands, and he that will
Reach her, about must, and about must go;
And what the hill’s suddenness resists, win so.
John Donne
We are most used to thinking of intelligence in terms of our own largely conscious thought.  We think of internal representation and memory as essential for symbolic thought.  We also think of external symbolic communication.  We think of abstraction, explicit conception and manipulation of conception.  Imagination, motivation and conditional thought are also essential to intelligence as we see it in ourselves.  Both to deal with internal conceptions and external situations by reasoning with abstractions and analogies, we need pattern recognition, induction and deduction. 
And yet, how many of those are essential to intelligence in broader senses?  Certainly not many of them occur in non-human communities, and those that do, seem to fail to rival the development of the human versions by several orders of magnitude. 
But there are difficulties all the same.  We still cannot recognise intelligence as such.  Suppose that a tree, or a species truly thinks, but a hundred thousand times more slowly than a human, and perhaps in some radically different ways.  Do we criticise it as unintelligent because it cannot think fast, and perhaps cannot solve crossword puzzles?  Speed and literacy may not be relevant as long as the tree can orient its leaves toward the light, avoid contact with neighbouring trees and adapt to the prevailing wind and the location of soil nutrients. 

Formally defining intelligence.

When all night long a chap remains
On sentry-go, to chase monotony
He exercises of his brains,
That is, assuming that he’s got any.
Though never nurtured in the lap
Of luxury, yet I admonish you,
I am an intellectual chap,
And think of things that would astonish you.
W S Gilbert.     Iolanthe
In this discussion I simply do not formally define “intelligence”.  The concept is not understood well enough for either general agreement or cogent argument.  As I point out elsewhere, we still lack any way to recognise subjective consciousness, much less express its relationship to subjective intelligence.  This frustrates useful definition.  In the definition of intelligence it is fairly sure that there are whole ranges of concepts that get confused and lead to question begging and cross purposes in discussion.  Rival points of view, valid or otherwise, commonly depend on tacit assumptions that lead to conflation of ideas and to disputants talking past each other. 
As I instance later on, comparison of the intelligence of individuals with that of communities is fraught with traps.  Examples that we shall see include questions of complexity, redundancy, efficiency and speed.  Not one of these turns out to be an unambiguous measure of superiority of intelligence. 
The favourite positivistic criterion for intelligence is empirical evidence for problem solving and communication.  It arguably is the most practically useful, but it is not formally cogent.  For one thing empiricism without an adequate formal basis can hardly improve on (fallacious) inductive argument unless there is adequate statistical argument to support it.  There cannot be cogent statistical argument unless there is a cogent definition that supports cogent criteria for diagnosis of intelligence.  To a great extent I do in fact rely on such empiricism, but it is for lack of anything better, not because I mistake it for proof. 
For the present, pending major breakthroughs in the field, I personally am pessimistic about defining intelligence compellingly.  Instead I stop at proposing a few constraints on what we usefully might call intelligent systems.  Because of the poverty of our understanding of the field, it does not follow that whatever falls within these constraints meaningfully qualifies as intelligence.  However, the construction of such lists of constraints is as powerful a tool as we have at present.  

Constraints on intelligent systems.

The scale, properly speaking, does not permit the measure of the intelligence, because intellectual qualities are not superposable, and therefore cannot be measured as linear surfaces are measured.
Alfred Binet 
The following proposals are neither exhaustive nor compelling; I intend them as useful bases for informal discussion of constraints on what we might regard as evidence for intelligence. 
Notice that I do not include the concept of personality as a constraint.  It neither is clear that personality (individualism or any form of unique behaviour or attribute) guarantees intelligence, nor that it is necessary to intelligence.  In practice, sufficient complexity in a system almost guarantees individual behaviour patterns, simply as a consequence of the nature of complexity and chaotic systems.  And yet it would require some very peculiar arguments to justify regarding say, a hurricane as intelligent, even though people who deal with them often personify them, and though hurricanes are immensely complex and no two storms have the same behaviour. 
The concept of subjective consciousness also is not logically necessary for an entity to be intelligent to all other intents and purposes, so I exclude it from the list of constraints.  

Intelligence and apparent teleology

I find you want me to furnish you with argument and intellects too. 
No, Sir, there I protest you are too hard for me.
                      Oliver Goldsmith
It is not clear on what grounds to deny intelligence in any system that exhibits apparent teleology, even when the same system simultaneously exhibits several versions of possibly independent, sometimes conflicting, teleology.  In this essay I unapologetically refer to such behaviour as intelligent in certain contexts. 
Such terminology is not to be confused with anthropomorphism.  It is more like the empiric pragmatism of the Turing test.  It also is difficult to decide at which level apparent purpose grades into non-trivial teleology.  A gin trap might be seen as behaving with intent when it snaps shut on anything that triggers it, but we might reasonably reject it as a candidate for teleological behaviour.  After all, a door in the wind may behave very similarly.  In any such system, how one defines the intention, as distinct from noise in the system, is an arbitrary point.  The engineer might be interested in a sprung ratchet as a means of enforcing a single direction of movement, and any noise it makes is well . . .  no more than noise.  To the football fan with a rattle, that self-same noise is the point of the device. 
 We surely however must accept that the trapper who directs a pathway to pass over his gin trap, does demonstrate significant teleology. 
Contrast that trapper with the larva of an ant lion or of a worm lion fly (Rhagionidae or Vermileonidae, depending on the author).  Such larvae excavate pits in sandy soil, use them as traps for prey, throw up sand to keep the prey from escaping, and snap shut on them as soon as they get into the right area.  Now, you know and I know that there is a difference between the proximal mental processes in the two types of system, human and insect, but as we shall see in examining social systems, there may be more for us to think about than just proximal mental processes.  It also is reasonable to speculate whether a given species of ant lion, seen as a population in a given region and time span, exhibits teleology, even if the individual ant lion patently is not equipped for it.  
I think it should be clear that it is quite possible to imagine behaviour that seems very persuasively teleological in a system, and that in some such systems one is very unwilling to grant intelligence once one sees how the programming of a simple mechanism works.  In cases of the type I have in mind the apparent teleology evaporates on detailed inspection, as in mousetraps or perhaps Skinner-type conditioning.  Sometimes apparently teleological behaviour may be very impressive, but be based on simplistic mechanisms (e.g. in scripted systems in computers, or simple physics such as in the behaviour of heated oil droplets or soap-bubble membranes.)   Highly sophisticated behaviour in invertebrates, either individuals or colonies, often turns out to be stereotyped and looks suspiciously mechanistic on close investigation. 
Similarly, many systems that demonstrably engage in information processing are not intelligent in any persuasive sense.  Examples include apparently teleological, effectively target-seeking behaviour, which is necessarily controlled by information processing in simple mechanistic programs.  Common examples are stereotypic, innate behaviour.  J.H. Fabre wrote impressive early discussions on stereotypic behaviour in insects.  Later ethologists saw many examples in birds and mammals, triggered by releasers. 
Humans are by no means free of apparently similar behaviour. John Crompton criticised Fabre’s views on stereotypic behaviour.  He compared certain instances of behaviour in humans with stereotypic behaviour in insects and pointed out that it is very hard to distinguish them by external inspection.  For instance, Fabre spoke of the “Ignorance of instinct” that caused a hunting wasp to seal a disturbed cell in which there was neither egg nor prey.  John Crompton in his book “The Hunting Wasp” compared this with the experience of seeing a heartbroken woman in the Blitz, her house having been practically levelled by a bomb, but with the front door still hanging in a fragment of standing wall.  Having “entered” and examined what she could, she “exited” again, and carefully closed the door.  As a deservedly popular lay writer on mainly entomological topics Crompton cut no ice in the annals of ethology, but he did clearly illustrate the philosophical problems of such inter-specific comparisons. 
Furthermore, it is now well known that artificial stimulation of parts of the human brain can elicit particular behaviour patterns.  In some cases the behaviour is of a visceral, unconscious, involuntary type, though often coordinated, but in other cases it is fully conscious, such as the hearing of a tune or the recall of a scene.  These events are suspiciously evocative of the irritating repetition of a tune or a phrase “running through one’s head”.  It also recalls the (almost?) reflexive elicitation of appetitive behaviour by very stylised stimuli.  For example, even in humans the merest smell, or a few lines of suggestive sketch can stimulate hunger, sexual desire, or fear.  
Some such stimuli are extremely widely spread in the animal kingdom, so much so that they occur in aposematic signals.  The eye-spots in the wings of moths or mantids might appear unpersuasive to humans, but they have been observed to alarm small birds when suddenly revealed.  I personally have repeatedly observed large Calliphorid flies assume a pose of alertness as soon as a chameleon brings both its eyes to bear.  The flies then react so effectively as to avoid the tongue impressively often.  Is the intelligence of such a response any less real than the startle reaction of a human who helicopters when he unexpectedly encounters a representation of a snake or spider? 
Given the survival value of such reactions in the wild, are we to regard them as unintelligent?  If they are not intelligent in the individual, are they unintelligent in the species?  There most certainly are bases for objections to such views, but are the bases perhaps not too narrow definitions of what comprises intelligence?  Would they not exclude certain classes of behaviour that deserve to be called intelligence in entities that differ from say, the higher primates and molluscs?  For example, Protoctista, Cnidaria, plants, species, programs? 
For as long as we cannot cogently define, characterise, and measure intelligence, it is dangerous to exclude certain categories of behaviour from our characterisation and definitions.  Even in humans, the main virtue of IQ tests is that they are numerically convenient.  They also are notoriously non-cogent, however useful they may be. 
It also is dangerous to argue that teleology depends on an understanding of the mechanisms necessary for the success of one’s purposive actions.  Consider a dog that has learnt to drop a ball into the receiver of a machine that delivers a reward.  If we then move the machine to another part of the room, such a dog often will drop the ball in the original part of the room, ignoring the fact that the machine is now a few metres away.  If we argue that such a dog, in whom we had originally thought we had recognised intelligently teleological behaviour, now demonstrates a lack of intelligent teleology, then what are we to say of the vast majority of human drivers of vehicles, or computer users? Most are blackly, incuriously, ignorant, not only of how their apparatus works, but of the relevance of many of the learnt actions with which they control the system. 
In fact, it is a safe bet that many of the experimental ethologists who use such a slot machine in such studies, also have little idea of how the slot machine works.  What is more, concerning many such aspects of the operation that they confidently think they do understand, the beliefs of such people often are flatly wrong. 
Nor is such superstition and ignorance limited to humans low of intelligence.  I have seen a whole class of very intelligent, lively, gifted children, who when I challenged them to explain how a pocket calculator worked, eagerly informed me that one pressed the buttons.  It took me an extended exchange to awaken them to the idea that button pressing, seen as a computational primitive, left something to be desired.  If that is teleology, then why are we sneering at those silly dogs? 
I personally have seen an Argentine ant  worker Linepithema humile (formerly Iridomyrmex humilis) walking across a leaf, where there were scale insects tended by the ants.  Perhaps 40 mm away there sat a Hymenopteran parasitoid of such scale insects.  When the ant came within some 10 mm of the wasp, it apparently saw it, abruptly changed course, and charged at it at a much higher speed than its erstwhile relaxed walk across the leaf.  The wasp flew off, and the scale remained unharmed. 
Nothing could be more deliberate, more teleological.  The wasp wanted to avoid being killed by the ant, especially before it had finished laying its eggs in the scale insects.   The ant wanted to protect its important food resource, and to eat the wasp that threatened the scale.  And though I did not specifically see this at the time, the scale wanted the ant to keep protecting it and removing the troublesome honeydew that otherwise would attract moulds.  Simultaneously, from a slightly different perspective, the communities of ants, scale and wasps were intelligently, teleologically, pursuing their ends.  Very likely the injured plant also was emitting chemical signals to attract the wasp, so that it might attack the scale insects. 
Or so we might reasonably gather from the observations.  Nothing I saw argued the contrary.  If similarly I had been in a position to watch, say a sunbird chasing a rival from a favoured Tecomaria bush,  an orang utan chasing a monkey out of a disputed fig tree, a soldier guarding a magazine from an insurgent, or a shepherd chasing a jackal from his flock, I should have interpreted what I had seen in just such terms.  How am I to justify a different interpretation in the case of the ant? 
Indirectly I could base counter arguments on published studies of the brains involved, or on publications on comparative ethology, but even after that, assuming that I accepted the arguments of those publications, I am left with nothing better than arguments from analogy to convince me simultaneously of direct conscious purpose in the case of the vertebrate minds and its absence in the minute insects or their communities.  “I have a subjectively conscious mind with its associated conscious purposes and a brain that supports all those things, and therefore it causes me to behave in a similarly purposive manner; they have brains of related structure and physiology, and similarly purposive actions, therefore they have similarly related conscious purposes.  QED.” 
Persuasive?  Perhaps.  Cogent?  Certainly not. 
What is more, various learnt behaviours, especially those that for their highest perfection of performance rely on high degrees of coordination and speed, such as in competitive sport, seem to rely largely on different parts of the brain from those that support conscious behaviour. 
In learning such activity the beginner relies on the cortex, and usually performs relatively poorly.  After some training, often after a necessary interval of non-participation, there may be a discontinuity in the standard of performance.  Reactions become partly automatic and details of the control seem to be relegated to the cerebellum, and the entire performance becomes smoother without the baggage of conscious control.  And yet, some of those subjectively automatic activities may be fairly complex, not just single, reflexive movements.  Here again such modularity in the brain has suggestive resemblances to eusocial colonies with their relegation of particular functions to specialist castes. 
It also raises the question of the site of teleology in a given system.  Does the trap have the teleology, or does the trapper have it?  The jaw or the ant lion?  The goalkeeper, or his cerebrum, or his cerebellum?  Or the whole system?  Consider a human practising a physical action mentally (an effective and important form of exercise).  More commonly he might practise partly mentally and partly physically, for example shadow boxing in the absence of an opponent.  To a naïve observer such behaviour might not be obviously teleological at all, because the effort is expended on no obvious objective.  Compare this with someone performing much the same mental processes with a phantom limb after amputation.  Some of the distinctions raise quite troublesome philosophical questions in terms of empirical and subjective teleology. 
Other empirical examples where what we could defensibly describe as mindless behaviour seems functionally equivalent to teleology are frequent in the ontogeny of multicellular organisms.  There the growth of exquisite organ structures is controlled by chemical and physical gradients that affect the growth, multiplication, shape, proliferation, migration, and not least, the selective death of cells in ways that look stunningly teleological.  Less precise, but none the less amazing, are the formation of the reproductive phases, “slugs” of slime moulds, the Myxomycetes.  Their behaviour irresistibly suggests, practically screams, teleology.  By what criterion do we deny the teleology of the system?  That it is brainless?  That it is stereotyped?  That the individual amoebae are brainless, as brainless as neurons?  That they follow chemical gradients?
Maybe, but on cross examination we find that we are reduced to some painfully special pleading and assumptions.  If our individual neurons are not conscious of our intentions, and it takes a whole brain to be conscious in that sense, then until we develop a working teleolometer, our arguments will neither embarrass the empiricists, nor demonstrate that inorganic computers are not subjectively conscious. 
Then once more, what about the intelligence of communities?  More particularly, what about the intelligence of communities (such as the population of a species) in which some members never meet or interact with others?  How do we define intelligence that does not rely directly on the transmission of information?  How do we deny intelligence in systems that display apparently purposive behaviour?  Do we argue that in communities with temporal dimensions in which one member can affect the behaviour of another, but not receive feedback, there is no control?  That would be nonsense.  Not all control has to be closed loop control, neither in engineering, nor in physiology, nor even in communities.
Conversely, we have the problem of arguing whether any community is a tool user or tool creator.  This is an arguable point.  The bees’ use of wax and propolis, or their creation of honeycombs; do those constitute tool use?  Like the use of a pebble for tamping down the cover of its tunnel by some Ammophiline wasps, all this is of course stereotyped behaviour, but what fundamental difference does that make, apart from the fact that the creation of new tools or advances on old tools becomes a slow, slow business?
We accept the idea of intelligence in brains whose activities are made up of the activities of their unintelligent cells, even though such intelligence is not additive; when we contemplate a brain of 1010 neurons and an IQ of 102, we do not attribute an IQ of 10-8, to each cell.  Nor do we attribute an IQ of nearly 1012 to the entire human race as an entity.  The components of an intelligent community neither have to be individually intelligent, nor unintelligent; it all depends on the kind of relationship between unit and unit, and between unit and community.  

Intelligence and Information Processing

I have no doubt that in reality the future will be vastly more surprising than anything I can imagine.  Now my own suspicion is that the universe is not only queerer than we suppose, but queerer than we can suppose.
J. B. S. Haldane

Intelligence is not the same as information processing, as far as we can tell, but it is hard to imagine any system that is intelligent, but does not include information processing as a key aspect of apparent intelligence. 
One type of information processing is vital to every undebatably intelligent creature.  It has not been clearly recognised in any colonies other than metazoan brains, or possibly in colonies composed of metazoa with such brains.  This is explicit information abstraction, particularly abstraction in the form of generalisation from pattern recognition.  Every vertebrate that recognises danger or food from experience uses generalisation by pattern recognition; the stimuli that the threats or prey evoke are not every time identical. 
In invertebrates, such abstraction and generalisation certainly occur, but not always from personal experience.  The wasp that stings a spider will recognise the spider in any attitude or by its smell, and will know how to apply the sting to the critical spot, but her actions are innate.  The experience she draws on is the experience of her species, not her own.  It is encoded in her genes, not in what she has learned since her emergence from her pupa. 
However, the bee that learns which flowers are open where, and when, certainly does so partly by learnt pattern recognition.  This has been elaborately and repeatedly demonstrated.  She passes on the information to a clique of workers in the hive.  This too is well known.  Could one say that the hive has now learnt to find and recognise the resource by pattern recognition? 
From some points of view one certainly could say something of the type, but one must allow for the fact that then the hive has a memory no longer than the life span of that clique (though not necessarily as short as the life span of the clique's individual members.  For as long as members of the clique keep finding such a resource, they will recognise its pattern if it is a good resource.  They might even remember how to dodge the flower if it is one of the kind that keeps swiping at them when they land.  But when they all die before the information has been passed on, any learnt skills die with them and must be re-learnt seasonally.  In fact it is worse than that, because not many learnt skills do get passed on in invertebrate populations.
An interesting problem is best exemplified by the dancing language of the bees.  It is undebatable that in this language a worker bee conveys certain variables abstractly and symbolically.  Those variables include distance, bearing, and the quality and nature of the food. 
Furthermore, unless the source is particularly rich and stable, commonly only a clique of bees, a minority of the available workers, will follow the discoverer, and go looking for the food.  If it is a particularly fine source, say the nectar flow of a large stand of alfalfa or eucalyptus, they progressively recruit effectively the entire nectar collecting population of the hive.  “But it is an innate, stereotyped program,” one might object.  Certainly, but what does that mean except that the skill is the skill of the species or the hive rather than of the worker?  It does not mean that it is not a skill.  There are elements of the Chinese room situation here too.  
At a higher level, only humanity has demonstrated the ability to reason explicitly by analogy.  But I am always moved to caution by reflection on a paraphrase of Orgel’s truism: “Natural selection is cleverer than you”.  Argument from analogy has a bad name, but in fact all argument about the empirical universe is in essence from analogy, and much if it is valid.  It remains valid as long as one does not exceed the isomorphisms between the object of the argument, and the notation and representation in one’s mind.  Argument from analogy comes naturally to humans (granted, often invalidly) but validly or invalidly it is a sophisticated process.  It depends on abstraction of a putative isomorphism from one situation and application of that isomorphism to a patently different situation.  This is not something that one commonly sees in animal behaviour, still less in non-human community behaviour. 
If we could see such a thing in a hive or colony, we should certainly find it easier to recognise it as intelligence of a high order, and more importantly, intelligence that makes sense in terms of our own outlook on the world.  

Intelligence in non-atomic systems

And he asked him, What is thy name?
And he answered, saying,
My name is Legion: for we are many.
Mark 5:9  

Every system that one might reasonably call intelligent is non-atomic (or is tomic, if you prefer) in the sense that it has distinguishable components that so interact as to process information for necessary activities.  In fact I have failed to find any convincing information processing system, biological or otherwise, that does not operate by interaction between its components.  This trivial observation becomes very relevant in dealing with the intelligence of colonies. 
Conversely, it is not clear that a complex system must necessarily have some form of intelligence, nor that the respective roles of the components of a clearly intelligent system will be obvious. 

Intelligence and communication

I confined one of these Eciton hamata under a piece of clay, at a little distance from the line, with his [sic] head projecting.  Several ants passed it, but at last one discovered it and tried to pull it out, but could not. It immediately set off at a great rate, and I thought it had deserted its comrade, but it had only gone for assistance, for in a short time about a dozen ants came hurrying up, evidently fully informed of the circumstances of the case, for they made directly for their imprisoned comrade, and soon set him free. I do not see how this action could be instinctive. It was sympathetic help, such as man only among the higher mammalia shows.
Thomas Belt                 The Naturalist in Nicaragua 

It is not cogent for an outsider to insist that if an entity does not communicate recognisably it is not intelligent.  There are whole classes of reasons why it might not communicate detectably; for reasons of its own it might not want to.  Even if it did want to, it might not use a channel of communication, such as a particular band of frequencies of light, scent, or sound, that the observer might recognise.  It might not speak the same language; many animals simply do not share the same signalling systems.  For example, the growl of a cat simply means nothing to most dogs, whose growls are much deeper.  Conversely, a surprisingly wide range of mammals, though by no means all, share innate play signals.  However, some classes of signals come to invert their meaning in the course of evolution; in context, bared teeth in humans more often mean friendliness than threat, while in dogs and monkeys the default meaning is threat, stress, or defensive submission. 
On the other hand some form of communication probably is fundamental to intelligence as such, even if only because, as I propose elsewhere, it probably is not practical to implement intelligence in any form that we understand the term, unless the components of a system communicate with each other.  In communities this aspect is visibly essential; a crowd hardly constitutes a community in any useful sense if the effectively meaningful communication between the elements is insufficient. 
Even conceding this need, there is a tendency to think of communication as a transmission of passive, coded items of information, but there is more to it than that.  In intelligent systems the communication processes and media generally include part of the data processing mechanism.  In the interface between units, information may be emphasised, attenuated, inverted, combined constructively or destructively, even supplemented or newly generated by what, in human conventions and techniques are called clandestine channels.  An example would be when a bridge player conveys information to his partner by tone of voice or deliberately timed hesitations in his bids.  We see all these and more in the interaction between neuron and neuron, organ and organ, person and person, and most definitely between members of colonies.  Signals, whether visual, auditory, chemical, or mechanical, go far beyond simple single-channel one-way transmission. 
Just for one example, the same chemical signal emitted by an organism can carry information concerning distance or time as well as identification or stimulus to activity.  Such extra information is not intrinsic to the signal as emitted, but is the product of modification, of processing, that the emitted signal undergoes once it has been released. 
Signal context also is crucial.  A signal at low level might be attractive, at a higher level or repeatedly increasing in intensity, it could be a threat or even an attack.  A signal received in one physiological state could stimulate aggregation, while in a different state it could cause dispersal. 
There are many more elaborate examples of signal processing between transmission and reception, both within a body, between individuals, and within communities.  Without such information processing en passant as it were, it is hard to imagine how insect communities in particular could attain anything like their elaborate and apparently intelligent structures and control.  It also gives one pause to imagine the complexity of the intra-community control. 
It leaves one with an interesting speculation: the signalling structures within say, the mammalian or indeed the insect body, are of stunning complexity and are nowhere near being properly analysed as yet.  And yet within a community we have enormously complex communication structures too.  How confidently can we say that either of those two is greater than the other?  How meaningfully can we say that the complexity of the community is the sum of the internal and external signalling of its members? 
One way or the other, communication in several senses at least, could be argued to be a constraint that any claim of intelligence must meet, if only because so much of the function of communication is as a component or amplifier of intelligence.  

Intelligence and subsystems

                                       . . .  Say, from whence 
You owe this strange intelligence?
William Shakespeare -- Macbeth
Except for trivial examples for purposes of illustration, components of an intelligent system necessarily include subsystems of various levels of complexity. 

Intercommunication between component modules of intelligent systems.

For such systems to function, requires a high degree of intercommunication between modules.  The intercommunication may be one-way, two-way, one-to-many, many-to-many or more than one of those modes simultaneously.
The intercommunication may be broadcast, as with pheromones or alarm signals, or point-to point as in nerves or tactile communication.  It may be discrete (in that respect being in some sense digital) or analogue.  It might function by language, whether human and largely conscious, learnt and formally sophisticated, or rudimentary, instinctive, and stereotyped as in apes and some birds and insects. 
Such intercommunication need not be rigidly hierarchical in terms of the relative size of the components.  In fact, the relationship is largely a network of modules rather than in the form of a loop-free hierarchical tree.  However, hierarchical or not, the various modules generally are “information hiding” as in “good practice” in computer programming.  That is to say that components inside one module typically have no insight into the processing tasks or data of different modules.  Conversely, components in general might not even be aware of their own function in terms of the effect they have on other levels or parts of the “organism”.  They might not be aware of any organism or intelligent entity at all.  Here Hofstadter’s “Ant Hillary” in “Goedel, Escher, Bach” springs to mind. 
This “good practice” in biological modularity, whether in brains or in colonies, does not so much reflect an explicit design decision, as the fact that the typical module is neither equipped to inspect, understand, or even care about the insides of other modules.  In fact, one-to-many and many-to-many communication tends to have wide ranges of “unintended” effects, and many of the attributes of pheromonal or hormonal systems are adapted to minimise the harm done by those unintended effects. 
Such “unintended effects” are denounced as abominable practice in current software engineering.  Nature however, has not read the training manuals.  Natural selection works on all the effects of a genotype, a “design”, and anything really inviable goes extinct fairly quickly, whether it is at the level of a gene, an individual, a community, or an ecosystem.  Whatever is not too drastically inviable eventually gets sorted out in a stable, established ecosystem, but in a disturbed ecosystem or where there are new, potential niches, those unintended effects may be the basis for a new, drastic evolutionary development within one species or more, what some people nowadays call a punctuation event.  It usually takes a while for selection to adapt all the attributes to establish a new ecotype, but evolutionarily the process is rapid.  A lot of selection can happen in a few thousand years, sometimes even in a few decades.  
When the controls of a system that relies on internal communications are insufficient or get interfered with, the results may be disastrous.  In humans we see this in the complications following on hormonal treatments or abuse.  Among eusocial insects we see an analogous example of disaster when a hive of the much maligned “African Killer Bee” (Apis mellifera dorsata) is invaded by fertile workers of the Cape Bee (Apis mellifera capensis). 
The principle is as follows.  Unlike most Hymenopteran worker castes, unmated Cape bee workers can lay fertile female eggs.  Their own colonial control communications have adapted to the situation, but other subspecies, including dorsata, have not.   As a rule, the disturbances in the hive pheromones wreck the dorsata colony till sooner or later only Cape bees are left.  Without a queen, their hive balance is unstable and the colony is likely to perish entirely, though it may sometimes muddle on till a viable queen takes over and establishes a viable colony. 
As we have seen, intercommunication between components is not just there for simplistic control.  The signals themselves are the sites where much of some kinds of information processing takes place.  Probably a great deal of functional intelligence depends on that principle.  There are many examples.  Synapses may be excitatory or inhibitory.  Pheromones may vary in their stability or mobility, so that the signal permits the appropriate reaction to old or distant signals.  Receptors vary in their reactions to persistent signals, many actively destroy the signalling molecules that they receive.  They thereby permit habituation or clear the medium for successive signals.  In fact, whole classes of deadly poisons work by inhibiting the breakdown of the compounds that carry synaptic signals.  

Learning and intelligence

If the capacity for learning is essential to intelligence, it raises difficulties in formulating the concept of entity.  A given organism or indeed, a colony, might or might not adapt its behaviour in the light of experience, but a species might, through natural selection, adapt to circumstances.  Such a species might do so in ways logically difficult to distinguish from learning.  If the species is defined as a population sharing a common fertilisation pool and having dimensions in time and location, is it an intelligent entity?  Is the learning capacity of any component, either more important or less important than that of the total system?  Important or not, it certainly is not generally equivalent.  “Learning” by a species is not the same as learning by a particular colony, which in turn differs from learning by an individual, and differs even more so from learning by a neuron in an individual. 
At the same time, these entities are deeply different in their respective natures as entities, and perhaps even more different in the respective natures of their intelligence.  What “learning” means in one differs from what it means in all the others. 
Temporal factors also apply.  Many activities of long-lived communities or species are hardly perceptible to the faster, but not necessarily more complex, minds of their members or observers.  This would make a difference even if the learning process were logically the same. 
In human history it frequently happens that quick-thinking individuals first exploit community weaknesses or stock market patterns, then ossify their thinking.  Typically they fail to recognise that their views were far too simplistic to represent the complexity of the behaviour of the community in general.  Sooner or later new developments bury them, either within their lifetimes, or within a few generations.  It is not clear how far such contrasts indicate a fundamental difference between the natures of the intelligence of individuals and communities respectively.  One could argue that such examples suggest differences in speeds as well as types of learning, and that the communities had more simultaneous modes of learning than their individual members. 
Conversely, in many communities the individuals can learn far faster than the community can.  It is a long-standing source of ridicule among Young Turks, that as Max Planck wrote: “An important scientific innovation rarely makes its way by gradually winning over and converting its opponents: it rarely happens that Saul becomes Paul. What does happen is that its opponents gradually die out, and that the growing generation is familiarised with the ideas from the beginning.”  Insofar as the assertion has merit, this is by no means true only of science, in fact as a rule far less true of science than of prejudices in everyday life.  Some scientists certainly are unscientifically hidebound all their mature lives, but they should at least in principle have a discipline according to which they may proceed to explore or reject new ideas.  Among laity who have not developed such a discipline, novelties are likely either to become fads or unjustly get rejected. 
Inappropriate rejection of novelty by the way, is not only true of old fogies; young fogies often are much, much worse. 
Anyway neither age nor newness is any guarantee of truth.  Facile new superstitions are no likelier to be true than centuries-old superstitions.  However, traditionally the very stability of communal beliefs is what preserves wisdom in hard times.  Old beliefs may have survived simply because they have been seen to work.  The general problem of what to learn and what to unlearn is no better conditioned than the general problem of what defines evolutionary progress or retrogression, and for analogous reasons.  In vertebrate individuals, including humans, the programmed reduction of learning capacity with maturation is an important and universal process.
In insect societies in particular, learning is a cumbersome and vague process.  There might be a minor reaction to experience by individuals, a “change of mood” because of continual stimulation in the community, or genetic adaptation by the species under the influence of selection. 

Coupling between subsystems or components of intelligent systems

Coupling of components may be intimate to any degree and in several dimensions.  It is fundamental to any form of group or colonial intelligence or within a body.  Consider some degrees and aspects of the coupling of such components in nature:

In loosely coupled systems, coupling may be behavioural, even casual.  
Coupling between components of a loosely coupled system may be facultative, as when either conspecific or unrelated organisms combine in hunting, guarding, nesting, grooming etc (such a community exhibits intelligence in some form of apparently conscious interaction, not necessarily consciously or consistently aimed at the benefit of each other or the community as a whole.)  Conversely, though one does find adventitious, episodic examples, the behaviour commonly is consistent and continuously adaptive.  On what grounds do we deny intelligence in such systems?  What kinds of intelligence do we deny them?  

Examples of tighter coupling include compulsory parasitism or symbiosis

Examples include hedgehog fleas, that appear to be necessary to the health of the hedgehogs, scavenger interactions where scavengers remove matter that would be unhealthy for the creatures that produced the litter, cleaner wrasse, protective relationships between ants and plants or plant bugs or both. 
Such a community of multiple species exhibits apparent intelligence in some form of obligate interaction.  Often the participants display no apparent teleological, mutually beneficial intentions.  However, their relationship does so, even though the teleology is not apparently conscious.  Any disturbance of the short-term balance between the benefits of participants is likely to be fatal in the long term, in the absence of “enlightened teleology”. 
For instance, oxpeckers take no interest in the well-being of their hosts; for them a tick is a convenient means of rendering giraffe or buffalo edible.  They will greedily cluster round a wound and do their best to eat the host alive, bypassing the ticks.  Excessive demands on one partner might kill it, in some cases leading to the death of the greedy partner as well.  One could regard cancers as examples.  Conversely, modest, consistent demands are likelier to lead to adaptation that might develop into mutualism.  

Parasitic or symbiotic vector relationships
More complex relationships include vector relationships in worms, protozoa, Legionella, typhus.  The vector interaction comprises an intelligent system, in many examples harmful to some parties, in some cases beneficial to both.  

Physiologically profound symbiotic relationships
Examples include ruminant gut flora, and gut symbionts in cockroaches and termites (several levels and dimensions of endosymbiosis in the same termite or cow!)  Particularly stunning relationships include some types of mycorrhiza.  Another extravagantly baroque example is  “the medusa & the snail”, as related by Lewis Thomas.  Such communities of more than one unrelated, but physiologically interdependent species exhibit intelligence in some form. 
Examples of endosymbiotic relationships include eukaryotic cells. Various species contain plastids, hydrogenosomes, and mitochondria.  Such a cell itself comprises a community of genetically diverse organisms interacting as an intelligent structure.  Such communities challenge our concept of “entity” to the breaking point.  Hydrogenosomes in particular are almost the inverse of the medusa-snail relationship, in that the DNA relocation to the nucleus has left the endosymbiont sometimes without any DNA of its own.  

Colonial sociable communities
Non-eusocial communities are sometimes termed “sociable”.  They differ from eusocial colonies mainly in the following particulars:
Members may not be family members with overlapping generations, such as sibs;
There may be no co-operation in care of young; and
There is no question of castes. 
Examples include certain seabirds, colonial caterpillars, and wasps that nest in close proximity, often in regular arrays. Sociability has many advantages.  Arrays of nests may deter predators or decrease individual exposure.  Combined labour can achieve things beyond what individuals could; for instance sociable weaver birds construct huge nests that no one pair could build alone. 
In such a sociable community each individual selfishly seeks its individual comfort level, but the community profits and adapts accordingly, and on average, so do the individuals.  There is a gradation of sophistication of such nesting patterns, from mere mobs to spectacularly regular arrays.  In a still more radical adaptation to predators and cold, emperor penguins have dropped their mutual close range repulsion in favour of huddling. 
In our context, this last reversed adaptation is particularly suggestive.  At various levels of adaptation, the nature of the association changes, or in some cases even reverses.  Such change in the species is reminiscent of the process of flexible learning in individuals.  It is just that in communities, particularly a community comprising a species, the “learning process” is not trivially visible and may take many generations of selective adaptation.  However, a sudden change in the selection applied to the population may achieve changes within a few generations.  Examples include changes in the flight behaviour of game birds under pressure of hunting or road traffic.  

Familial communities  
Familial communities involve parent-young interaction.  There is some degree of care for offspring, the simplest form of which is selective oviposition, grading to elaborate parental care.  At higher levels there may be co-operation between siblings or between offspring and parents, such as when scrub jays, foxes  or carpenter bees share the burden of raising genetically related young. 
The components of the intelligently altruistic system are derived from those innate behavioural patterns that are necessary in non-social care for offspring.  I discuss examples in the section on inhibition of aggressive behaviour.  Familial altruistic behaviour grades into true social communities, for example, we get solitary mole rats such as the Cape mole rat, Georychus capensis, sibling assistance in family parties in the Hottentot mole rat, Cryptomys hottentottus, and then eusocial species of naked mole rats. 
Such familial communities appear to have been a stage in the evolutionary development of all the eusocial species we know.  It is not good to be simplistic about it though.  There are examples, such as some of the paper wasps, where the original family party is not a single queen and mate, but a small group of cooperating queens.   

Colonial eusocial communities
Examples include naked mole rats, sponge-living shrimps, termites, ants, wasps and bees, where there are non-breeding castes that are pheromonally and behaviourally determined.  Quite minor interference with such pheromonal controls can be fatal to the colony, such as the effect when Cape bees (Apis mellifera capensis) invade hives of African “killer” bees (Apis mellifera dorsata). 


The colonial nature of multicellular or physically crowded communities.
There are many examples in nature, of effectively sedentary organisms that live in close proximity, sometimes physically connected and reproducing vegetatively; many such creatures behave in orchestrated fashion in their feeding, defence or mating, even though their physiology and organisation is neither that of a body nor of a eusocial colony.  Examples include beds of mussels, bryozoans, barnacles, and corals.  Typically they all snap shut if a single member is alarmed, open together for feeding in synchrony, and release their gametes or young in synchrony. Their co-ordinated movements resemble those of shoals of fish or flights of birds.  Their growth patterns resemble the growth of organs in animals or branches or roots in plants.
Relationships between cells or organs within the body’s ontogeny, including cell death where it is necessary, irresistibly recall colonial interrelationships.  Consider tumours as counter-examples.  Where cells refuse to die on cue, one gets malformation and perhaps death of the colony.  Note how the nature of the genetic relationship affects the employment of cell lines in sacrificial roles.  Since the somatic cells in most organisms are all genetic clones, it is well worth a cell’s while to sacrifice itself if that improves the survival of the organism.  Consider for example cells in slime moulds or in embryonic tissues of multicellular organisms, and compare them with competing or cooperating team members in a colony. 
All such co-ordinated behaviour is closely related to the intelligence of communities, even though the range of the intelligence in the individual members varies from practically zero in the amoebae to birds with some of the highest intelligence in the animal kingdom, and yet the apparently intelligent behaviour of the various communities is strikingly similar.  Crowd traffic behaviour is in many ways similar in everything from ants to humans.  The reasons seem to have a lot to do with mechanical constraints, but that does not reduce their significance for the concept of community intelligence.  

Language, abstraction; dances and semiotics

. . . An expedient was therefore offered, that since words are only names 
for things, it would be more convenient for all men to carry about them 
such things as were necessary to express the particular business they 
are to discourse on. . .  I have often beheld two of those sages . . . who, 
when they met in the streets, would lay down their loads, open their 
sacks, and hold conversation for an hour together. . . 
Jonathan Swift                         A Voyage to Laputa

Pending anyone demonstrating the contrary in some way I cannot imagine, I regard information processing as one of the essential components of intelligence.  In information processing, communication necessarily entails the encoding and transmission of information to be processed.  In turn, this entails abstraction; some device must be put into a state that reflects at least the relevant parameters of the object that is the subject of the information.  It also must not present too much information besides.  In other words, the signal to noise ratio must be manageable, or the abstraction is likely to fail. 
Consider for example, the conversations of Swift’s Laputan sages, in which they displayed things instead of using words.  This would not have worked very well because of a specific reason: semantic ambiguity.  The abstraction would have been omitted and therefore the meaning would be ambiguous. 
Depending on how one looks at the act, to exhibit a knife in lieu of a word or a phrase could amount to presenting anything from several dozen bits of information, to ten to the several dozen bits.  In “holding conversation for an hour together”, the act of showing say, a knife, could mean “a knife”, “this knife”, “knives in general”, “iron”, “a mass of 250 grams”, “this colour”, or any of many other possibilities.  Further information must be added to indicate which of all the possible attributes of the thing is being presented for consideration, in other words, to filter out some of the critical noise. 
No doubt in practice sophisticated savants would add the necessary supplementary information by employing high-flown conventions and techniques of the charade, but in doing so, they would betray the ideal of displaying “such things as were necessary to express the particular business they are to discourse on” instead of using words;  linguistically and semiotically speaking, formalised gestures and grimaces are nothing less than words.  The medium also presents other difficulties, such as how to express relevant non-material concepts such as relationships or verbs, but those are minor compared to the abstraction problem. 
So it is with say, the language of the bees.  Unlike the Laputan sages, bees specifically use abstractions, even if their abstractions are not conscious.  Whatever consciousness a worker bee might have, if it has any at all, it does not react to the food that the worker has recovered as “this nectar”, “this pollen”.  It reacts instead to the abstractions: “such nectar, in that direction, so far away, at that time, to be identified by such a smell”.  Also, within the bee’s information processing mechanisms, as opposed to its foraging mechanisms, it is not pollen that gets passed on, but nerve impulses and synaptic signals that are as abstract as any signal passed by a human or an artificial signalling apparatus. 
The very fact that intelligent entities (apparently inescapably) are modular, implies communication, at least between the modules.  Otherwise the modules cannot non-trivially constitute an entity. 
Some kinds of communication, as we have seen, are so simple that they barely qualify as language in any non-trivial sense.  On the other hand, for non-trivial intelligence we need to convey concepts and data in non-trivial notation.  A few whiffs of hormone molecules really cannot suffice as non-trivial examples.  In the brains of the most spectacularly  intelligent mammals and birds, modules in the brain must convey complex, sometimes intrinsically abstract, concepts for internal manipulation.  “Think when we talk of horses, that you see them         printing their proud hoofs i' the receiving earth. . .”  Thinking thus, it is not little horses that we pass about the affected parts of our brains, but impulses and synaptic states.  Merely explicitly thinking about anything implies some level of ability for communication, abstract communication within the brain.  In the light of information processing practicalities, this implies at least the first requirement for abstractly communicating the same information outside the brain. 
Now this raises practical and philosophical difficulties in imagining recognisable advanced intelligence of species or social communities.  If all the extant specimens of a species are subjected to the same selection, then the species will respond by adapting as a unit.  Is that intelligence?  If it is, then is it intelligent for a snowflake to grow symmetrically, even though its various points are not communicating with each other?  To claim this is not persuasive, but the denial is not watertight.  Unlike non-contiguous parts of the snowflake, various parts of the species do in fact intercommunicate and adjust their response to selection by the redistribution of DNA.  Such responses could be seen either as very abstract indeed, or barely abstract at all. 
Obviously the problem is tricky.  There are several aspects that may be regarded in various ways, and some of the parameters are not well enough characterised for us to measure.
Within large mammalian brains, such as in naked mole rat workers, some such processes of communication between modules no doubt occur, and, at a lower level, possibly also occur within the smaller and less elaborate brains of termites.  There always is communication within colonies, even if it is only recognition of nest mates, alarm pheromones, or recruitment to activity patterns.  However, within the beehive is where we see the most advanced and explicit approach to language.  It is impressive in its own terms, being specific, semantically abstract, and fairly parsimonious. 
None the less it is not comparable to communication among humans, nor possibly to communication among some other mammals or birds.  For one thing, it is not a general means of communication, but highly constrained to particular subject matter.  It would fare poorly in a Turing test. 
If we are to find a truly advanced community intelligence, it must take some radically different form, possibly at some higher, more abstract level. 
And, in spite of our human intelligence, how much more intelligent are human communities than insect communities?  Certainly in some senses our communities can converse with apparently indefinite flexibility.  If we judge by the more sensible members of society, one would like to think our communities are very intelligent, but it is sobering to remember that majorities of the intelligent public concerned, supported Hitler, Stalin and some more recent political bodies that have highly invalid arguments in support of their policies.  To the extent that we as humans are influenced by the pathological, the influence of the intelligent in anything but the persistent improvements in technology and infrastructure, is debatable. 
One part of the problem with our community intelligence as opposed to our individual intelligence, is that the complexity is intrinsically greater than in a unit brain, and it lacks the equivalent of many of the controls in healthy unit brains.  Managing such complexity is currently neither practical, nor even foreseeable.  This is presumably why communities so often behave apparently more stupidly than most of their members, even when leaders are well-meaning and intelligent.  They are necessarily chaotic systems and we have not yet learnt how to damp their oscillations and excursions. 
How does one measure the relative intelligence of a system that involves stereotyped behaviour of its components, as opposed to systems that incorporate putatively subjectively conscious components?  It is a messy business to compare the intelligence of a termite nest with that of a human community.  Bear in mind that it does not follow that the intelligence of the community exceeds that of each, or even any, of its members. 
Possibly it is as well for animal colonies, that they have not yet achieved the internal complexity and communication characteristic of human communities.  As a rule evolutionary selection work more effectively on innate, stereotyped behaviour such as in insect colonies, than on the disorganised innate impulses and weakly-stereotyped rationalisations of human colonies.  However, it does not follow that human communities are exempt from such selection.  Lorenz speculated on personality traits such as degrees of aggression, being selected for in particular communities.  Possibly his views were simplistic, but like many of his speculations, they were intellectually fertile. 
The language of the honeybees is a useful model in several ways, though it is too complex for detailed discussion.  The nature of its abstraction is interesting at several levels. 
Warning: here follows a Just-So Story informal enough to drive any positivist, let alone any serious ethologist, to distraction, but it will have to serve as convenient illustration. 
The successful foraging bee returns from the field reeking of the food source, tired in proportion to the wind and the length of the trip.  The direction of the sun is her most important landmark and clue to her bearings over a distance of anything from a few metres to perhaps a few kilometres.  She cannot speak directly, but what she can do is to convey her condition to her hive-mates.  They in turn must accept her (necessarily) abstract message and encode it for comparison with their own internal condition. 
First she must pass the guards at the hive entrance.  They are there for several reasons, but a major one is to protect the hive from strange bees; in spite of their many virtues, bees have a practical outlook on honey: wherever you find it, grab it!  Accordingly, if they find it in any hive that lets them in, they immediately load up all they can, and take it home.  For the guards charity begins at home, so not many strangers get in. 
Mind you, the guards are not really perfect.  I have already mentioned the Cape bees invading the hives of “killer” bees.  The reason that they can do so is the slightly leaky protection of the guard bees. 
Now, it sometimes happens that a bee eats some fermented food, rich in ethanol, and the effect on bees is much like that of humans.  Any crapulent bee that finds its way home tries to enter, but it is likely smelling of the wrong mix of scents, so the guards will first shove her out, or if she persists, may bite or even sting her.  So we can say that her first signal on coming home amounts to: “I smell right and behave right; that is my password.”  
A frequent communication on the way in is for the returning forager to make sure that foragers on the way out get a good sniff of the flowers they have been collecting from.  She even might feed them samples.  If it is a kind of flower that the others know where to find locally, that might be sufficient to send them off to the right source.  Investigators have used sneaky tricks to fool bees into visiting the wrong flowers to prove that this scent and feeding trick is a non-trivial message.  It also proves that the bees knew where to find the wrong flowers; they must have memorised where to find many of the flowers in the neighbourhood of the hive.  This too is a non-trivial form of intelligence.  In fact, when bees are to be used for pollination of crops, it sometimes pays to feed the bees with syrup scented with the appropriate flower.  Then when the flowers open, the bees are ready and willing. 
Often the bees also mark a rich nectar source or a good source of water with their own scent glands to signal nest mates that they have reached the right place.  This is especially interesting because less extremely adapted eusocial bees that do not have the dancing language, use scent marked trails to guide nest mates to food sources.  
Honeybees also use their own scent marking to indicate that the food source is very rich, rather than just telling where it is.  When they do that, they tend to attract a lot more followers than just by doing their dance.  It is all part of the signal that adjusts the number of foragers to match the richness of the source.  If every bee that found enough to dance about would recruit the whole hive, that would be worse than useless, it would be disastrously wasteful of effort.  Instead she visits a food source several times before doing her scent marking.  That way she is sure that she won’t be sending her sisters after a temporary attraction.  It also is a good mechanism for assessing when to release the scent, instead of relying on the equivalent of a conscious estimate. 
When the forager gets into the dark of the hive, she cleans up, stores what she has brought, or passes it onto young workers who are not yet foragers.  If the source is a good one, plentiful and rich, and she has visited it enough times to move her to communicate the fact, the bee will start to dance.  She does this in the dark, on a vertical surface, usually a comb.  It is a complicated business with several variables.  First there is the type of dance.  If the food is just outside, say up to 25 metres away, she will do a round dance, roughly in a circle one way, then turning round and doing a circle the other way.  She may keep this up for a minute or so, or just a few seconds, with or without repeated encores.  By and large a specially good source moves her to longer, livelier dancing.  The only bees that follow her around are other foragers.  Young nurse bees and other non-foragers keep away, as well as one can in a crowded hive.  The dance carries no message for them! 
Between the scent and the performance, and the number of other bees that go foraging and repeating the message, each food source generally gets the most suitable number of foragers.  If there is a famine, practically any reasonable source gets recruits, but if there is a honey flow on, only the best sources set the bees dancing. 
At slightly greater distances to the food source, the circle that the bee dances becomes sloppier and no longer closes.  In fact it begins to take on a sort of distorted figure eight shape.  If the distance is much longer, say over 100 metres away, the dance is a proper figure eight with a good long middle stroke, not a wasp waist. 
At such distances the foragers are beginning to take note of the direction instead of just searching round the hive.  Here is one of the most remarkable aspects of the dance.  So far we have had fairly modest levels of abstraction and not much indication of anything but lot or little food nearby.  But for directing bees to distant sources we really need some good indication of direction and distance.  Otherwise we have not told the hive much more than that there really is food somewhere. . . 
The directional information is supplied by the direction of the cross-stroke of the figure eight.  The way it is indicated is so unexpected and so hard to detect that von Frisch richly deserved his Nobel prize for spotting and elucidating it.  What it does is to show what the bearing is relative to the bearing of the sun away from the vertical.  So at local noon in the high latitudes of the southern hemisphere, the sun would be due north. 
The problem remains that the bee is dancing in the dark of the hive.  How is it to represent the direction of the sun?  This is one amazing thing; the direction of the sun gets abstracted and represented by an alternative stimulus.  Usually the alternative stimulus is gravity.  It is as remarkable as humans graphically indicating a horizontal direction on a map hanging vertically on a wall. 
Much as we represent with a streak of ink on a vertical map, a direction to walk horizontally or up hill and down dale on the ground, the bee represents the journey with the middle stroke of the figure eight dance.  A dance with the middle stroke of the eight going straight up, means: “Fly towards the sun”.  If the stroke is at 45 degrees to the right of straight down, it means: “Fly away from the sun at an angle of 45 degree toward the right.  In other words, if the sun is due north, fly at a bearing of North 135 degrees.” 
There are various exciting exceptions to some of the conventions, but the most important is what happens when the sun is obscured by heavy cloud.  Then if there is any sizeable patch of clear sky at all, the bees deduce the position from the pattern of the polarisation of the light from the visible sky.  One amazing thing is their accuracy.  They can estimate the azimuth of the sun when it is barely 3 degrees from the vertical.  Given a fair sighting, their dances can direct their sisters to a mean accuracy of about one degree.  Even given a compass, an inexperienced human might do a lot worse. 
The accuracy of the direction is all the more amazing when one thinks how the distance is conveyed.  The fellow workers crowd round and feel her movements with their antennae.  The bee does not do the middle stroke as smoothly as possible, but waggles her abdomen from side to side as she runs.  Now comes another amazing thing.  When the bee is on the middle stroke of a serious dance, she begins to buzz with her wings, and the length of the buzzing run tells her sisters how much flying to do to get to the food.  Roughly speaking one second of buzzing run means one kilometre of flight.  Don’t take the exact figures too seriously; they vary.  The graph of distance as a function of the length of the run is not perfectly straight, and the bee does not dance just once.  Usually, especially for rich sources of food, she goes over the course repeatedly and her sisters track her time after time.  They base their conclusions roughly on the mean of the performances.  The message they get is remarkably accurate, as I think you will agree, considering how poorly humans typically estimate and convey such information without special instruments. 
Another very interesting fact is that novice foragers take a while, typically a day or so, to learn to interpret the dance properly.  In fact, in the dark of the hive novices have difficulty following the dancer with their antennae.  This is another dramatic example of how the innate mechanism of particularly complex behaviours often needs practice before it works at its best.  You might draw comparisons with sheepdogs, whose handlers speak of “teaching” the dogs to use the innate skills of the breeds, not “training” them.  Another analogy would be the learning of language by human children; large components of their learning seem to be innate, a mapping of the mother tongue onto an inborn universal human language. 
There are many amazing things about the communication system of the bees.  One is that the bee has an internal clock that allows for the movement of the sun.  She changes her direction of dancing even over a period of several hours’ delay when indicating which way to go.  Also, her information concerning distance is not the distance as a surveyor would measure it, but a measure of the work it takes to get there.  This includes allowance for up hill, down dale, crosswinds and the like.  But there is more.
However convoluted the course that the bee flew in finding the food, she flies back in about as straight a course as possible and on her subsequent trips, she also flies efficiently.  So do her followers in their turn when they respond to her dancing. 
At first it is hard to imagine how such an elaborate and abstract a language could have evolved.  Fortunately we get very suggestive clues to the history if we compare the honeybee language with the languages of some other species of bees. 
Think again about that buzzing run.  The bee in effect runs in the direction of the food. It is not the actual direction, but one that is isomorphic to the direction.  After all, remember that in the convention of the honeybee dance, the direction of gravity substitutes for the direction of the sun.  Honeybees do occasionally dance on horizontal surfaces in the sun, and some related species of bees always do so; and when that happens, they do the buzzing run in the actual direction of the food. 
Now, as it happens, some other species actually recruit followers with smells and other clues, but then fly off and lead their sisters in person.  Some species leave scent markings at various points along the way.  In the light of this, it is easy to imagine a honeybee ancestor doing just that.  Then, when the navigation got good enough, the scent marking could be reserved for just the target area.  Then maybe the bee got good enough in passing on the message, not to have to fly all the way.  Finally the pathfinder flight got stylised into just the length of that buzzing run that forms the middle stroke of the figure eight. 
Proof?  Of course not.  The foregoing discussion is an exercise in the assessment of a reasonable conjecture for a course of adaptation in information handling.  It is a course that involves no teleological leaps of ability or drastic genetic discontinuities on the way.  For the purposes of this informal essay, that is sufficient. 
Or is it?  Where did the bees in the first place get the ability to convey to fellow bees that there was something  they should be doing?  There are two aspects to this.  Firstly, sociable (nothing like eusocial) organisms commonly are adapted to feed together.  Except for creatures that live actually in their source of food, I cannot think of a eusocial animal that does not cooperate in finding food and bringing it back to the nest.  Those exceptions are the likes of the shrimps that feed on the sponges inside which they live, or thrips and aphids that live in the hollow plant galls that they themselves caused to grow.  The majority of those species that cooperate in foraging, lead their nest mates to food, if only by the smells they release in feeding or in storing the material they have collected.  Food and protection are the most obvious primary benefits of sociality. 
Secondly, decades ago studies of non-social species such as various kinds of flies, revealed some suggestive patterns.  Some of their behaviour patterns in feeding are suspiciously reminiscent of some of the bee movements.  In particular, they are delayed responses to food, say after a morsel of food has been removed.  If all the ancestral bees did was to be attracted to bees that had just been fed, that could well be the original basis for the whole development. 
There are other bee dances too, such as a buzzing run to indicate the direction of a desirable residence for a swam to move to, but many elements of such honeybee language have not been decoded yet.  Some of the putatively communicative behaviour might even have no operative significance; they may be vestigial movements prompted by assorted stimuli.  It is not a simple matter to decide such questions before one manages to identify the semiotic significance of a language. 

Democracy and didactics in communal intelligence

Loose systems last longer and function better.
            John Gall           Systemantics

Another thing to get clear is that the behaviour patterns that support foraging and communication in eusocial species are not in general automatic and deterministic, even though they incorporate a good deal of stereotyped behaviour.  In many kinds of eusocial colonies there typically will be several foraging recruitment efforts going on at any one time in a good season.  Which invitation will succeed in recruiting a given new forager, will depend on that forager’s own apparent whim and the intensity of the performance. 
In turn, in bees for example, the quality of the source, the intensity of the competition at the site, the exertion of the flight, all affect the recruiter’s dance one way or the other, so there is are continual changes in the numbers of bees that visit each particular feeding site. 
This sort of chaotic democratic process is characteristic of practically all co-operative eusocial endeavours.  In fact it is fundamental to eusocial control.  Note that such non-deterministic classes of algorithms need not at all necessarily be inferior or unsophisticated.  Even in human programming random resolution of conflicting demands for resources has proved to be quite efficient in most cases, and above all, has a very reliable worst-case performance.  In eusocial colonies the resolution is not truly random, but is more like what we might call the P. T. Barnum approach.  Whatever keeps catching the attention of colony members is likeliest to recruit them, and if they are disappointed, they are likely to follow rival recommendations.  Such classes of resolution are particularly suited to the needs of eusocial colonies because they are simple, flexible, and degrade gracefully except when a pathological agent exploits them. 
This is a prime example of why I speculated whether there are multiple types of intelligence.  Barnum influencing of the community looks as convincingly like a component of intelligent information handling as any synaptic or hormonal communication.  
Some of the most impressive examples of Barnum persuasion occur during swarming in honeybee colonies.  Swarming is a variable and complex process and the following remarks are nothing like a full or coherent discussion, touching only on a few illustrative points.  Typically, early factors promoting swarming include crowding, overfull stores of food, and the construction of queen cells.  Such stimuli lead to discontinuities in colony behaviour that provoke workers to go scouting for nesting sites and prepare the queen to stop laying and join the swarm. 
On leaving the hive, the swarm typically assembles at some intermediate point.  It may stay there for some time, typically several hours.  Sometimes it might take several days before the bees agree on where to go.  In deciding whether such an assembled swarm should move to one new home or another, the scout bees strut their stuff on the surface of the swarm itself, and as one group or another of the followers moves to prefer one option or the other, the swarm may vacillate more or less persistently.  If no clear choice emerges, the swarm might even split, each going to a separate new home, one swarm necessarily being queenless.  Such a splinter swarm lacks the queen pheromones, so they soon have to unite again with the other.  
Usually however, the end result of such plebiscites is a working solution, one way or another.  When ants move their nests, there will generally be whole trails carrying larvae, eggs and pupae hither and yon and, depending on the species, possibly even queens.  As one side or another prevails, individual ants might change their allegiance several times, doing and undoing.  This is more or less how driver ants, that are almost always on the move, decide where to assemble in their protective bundles or which trails to follow.  When nest building, eusocial insects might tear down each other’s structures to use the wax, paper, mud, or whatever it might be, for their own efforts. 
Once again the thing to note is that generally the ultimate outcome is not just any workable result, but a something that corresponds to that particular species’ style of doing things.  The comb, termite nest, anthill or trail activity is recognisable as typical of that type of colony. 
It all looks not only like intelligence, but personality and style as well.   When we as humans decide on a preference, who are we to claim that our cerebral neurons behave any more efficiently or sensibly than the individual colony members? 
Bee dances do not offer the only examples of interesting behaviour patterns and information transmission in recruiting foragers.  Ants also recruit fellow workers in various ways, and in particular, a recent observation is interesting in that the transmission of information is reminiscent of teaching, rather than simple telling and showing.  Foragers of the ant Temnothorax albipennis will lead fellow workers to a source of food, but the leading ant only proceeds while it is tapped on its abdomen by a follower.  Whenever the follower stops tapping and inspects its environment, the leader stops too, and waits for the follower to resume.  The whole procedure considerably slows down the leader ant, so it has an associated cost.  It follows that the mechanism is an important adaptation for efficient recruitment of foragers so that they can find their way to the source of food and back again. 
In some ways this is not radically different from the way that some species of bees (not  honeybees of the genus Apis) wait to recruit hive mates before going out again to the source of food. 
In much the same way as we have difficulty on defining intelligence when there is little reason to believe that activity is consciously teleological, so such behaviour leaves us in difficulties in trying to define unambiguously, the concept of teaching. 
There is yet more to it than that.  Many control systems in nature work, not on a basis of logical propositions, but on a basis of thresholds and majorities.  This is a very pervasive principle.  For one thing, it is fundamental to the control of cellular activity and growth; when things go wrong in cellular control in a an embryo, it causes anything from minor birthmarks to horrific abnormalities.  The slightest, subtlest changes affect the form and character of tissues and organs, the fit of teeth and joints, the shape and texture of bones, leaves, and flowers.  Such logic is the basis of hormonal control, physiological life histories and rhythms, circadian and other timing rhythms, and the movements of cellular organelles and of gross organs such as muscles. 
Above all in our context, such logic is the basis of the reaction, communication and control of individual neurons and of neural systems.  Unlike the deterministic threshold logic of most electronic devices, brain neurons fire only when triggered by a not very deterministic combination of trains of input pulses from various synapses or sensors.  Some inputs might be inhibitory and others stimulatory.  Sometimes particular combinations of inputs are necessary, such as in the eye, where retinal receptors detect features such as edges, horizontal or vertical lines, blocks of colour, or particular textures. 
And such mechanisms are irresistibly suggestive of control in communities, all the way from bacterial colonies, through the coordinated behaviour of slime moulds, the control of foraging or bivouacking driver ants, the recruiting of foragers or swarms in bees or ants, flock manoeuvres in shoals of fish or flying birds, even unto the behaviour of human communities.  Sociology has a lot to learn from such complexity, control, and information theory, before it can assume its place as a predictive and constructive applied discipline for control in human communities. 
The question arises why such an unobvious and complicated mechanism should be so pervasive in biology, while human control engineering predominantly relies on either analogue control or finite state control, increasingly often nowadays, in the form of explicitly Boolean logic. 
The most obvious suggestion is that threshold and majority controls could most simply evolve in cells that already contained mechanisms that supported such functions.  The next is that such mechanisms could work in many ways and perform wide ranges of functions.  They also are fail soft; they often continue to work under unfavourable conditions.  They are versatile in that they sometimes continue to work validly when presented with unfamiliar challenges. 
Another point is that such mechanisms also lend themselves well to genetic control and natural selection.  It is not easy to imagine the functional equivalent of electronic circuits being specified genetically in such a way that the resulting brain contains more information than the specifying genome.  Nor is it easy to see how such a system could undergo random modification without gross failure.  And yet, that sort of adjustment is just how natural selection routinely adjusts the logic of thresholds and majorities.
A point of particular interest is that in communities of largely independent units, like ants in eusocial colonies, instead of somatic cells fixed in the body, individual members seem to function in ways reminiscent of neurons.  The resemblance still is tenuous, but if it develops to a higher degree, it may be the basis for the formation of a recognisable community intelligence, literally comparable with the kind of mind supported by a unitary brain, and for the same reason. 

Intelligence and subjective consciousness 

MIND: A mysterious form of matter secreted by the brain.  Its chief activity 
consists in the endeavor to ascertain its own nature, the futility of the attempt 
being due to the fact that it has nothing but itself to know itself with.
   Ambrose Bierce  --  The Devil’s Dictionary
It is not clear what the relationships might be between apparent intelligence and subjective consciousness, but it is hard to imagine subjective consciousness in a system that has no relevant and elaborate form of intelligence.  Subjectively, subjective consciousness seems to me to require not only intelligence, but meta-intelligence, by which I mean intelligence aware of (at least its own) intelligence, but I am not at all certain that my impression is correst.  Certainly there seem to be various degrees of self-awareness among animals of different levels of objective intelligence, and yet among at least the most patently intelligent mammals and birds, their behaviour suggests some sort of subjective consciousness.  And the clearest suggestions are to be seen in the most socially communicative of species. 
Such remarks are normally taken to imply anthropomorphism, but I reject that charge in this connection.  If anything, it is the converse of the usual forms of anthropomorphism.  In practice we have no more cogent evidence for subjective consciousness in fellow humans than in fellow animals.  This is a (rather weary) problem frequently raised by existentialists.  The fact that behaviourist experiments and scripting software can produce impressive demonstrations of apparently conscious behaviour, does not logically compel one to accept that all apparently conscious behaviour is unconscious and mechanical.  Just because one cannot demonstrate the fact cogently, sceptics have no need to refute the suggestion, but this still does not establish that the idea is illogical or even that it is not a useful basis for criticism of experimental models. 
To pile difficulty on difficulty, it by no means follows that because an empirically intelligent entity passes the Turing test, including (truthfully) producing evidence that it is empirically aware of its own mental processes, it has a subjective consciousness such as humans do.  A mechanical or electronic computer could in principle be programmed to access and discuss its own data as well as external sensory data. 
In other words, subjective consciousness might very temptingly suggest meta-intelligence, but meta-intelligence need not imply subjective consciousness. 
The significance of such meta-intelligence, and perhaps subjective consciousness in general, would extend beyond intelligence purely and limitedly applied to external problems.  It is quite easy to accept that meta-intelligence could be of functional mental importance, but it is commonplace to assume that therefore subjective consciousness also is functionally important, almost a basis, an essential aspect of intelligence in any meaningful sense.  That this is so, is not obvious.  It is not clear that subjective consciousness is not an emergent, possibly non-functional, effect of the way that our brains function as intelligent systems, while say, an electronic device might pass the Turing test at an arbitrary level without any subjective consciousness at all, as far as we could tell. 
Various writers have pointed out that subjective consciousness seems to be uncommonly closely associated with short-term memory.  This might of course be no more than the consequence of the fact that our speech processing and idea retrieval and co-ordination depend on that part of the brain.  There might be a lot of other consciousnesses in the brain that never connect directly with the subjective consciousness that has contact with the outside world.  Speaking purely for myself, I often get a vague impression that there is a consciousness behind my primary consciousness, and that that is what feeds thoughts, words, intentions and so on to that accessible consciousness.  Uncompelling though such subjective anecdote might be, it remains suggestive in a field where suggestiveness often is as useful as anything one may reasonably expect. 
For instance, assuming that what we might call the short-term memory consciousness is indeed the main, definitive, subjective consciousness, that would have some important implications.  They are not definitive, conclusive, unique implications, but they open important possibilities.  Suppose that the short-term memory consciousness is somehow intrinsic to the short term memory as a region of the brain.  That leaves open the possibility that subjective consciousness is generated by the logical control functioning of a brain region.  If this is indeed the case, then we have no basis in logic for denial of subjective consciousness in the brains of animals, including insects. 
It also leaves us with serious questions concerning the possibility of subjective consciousness in social communities.  If regions of the brain that access and process data for immediate use are responsible for our subjective consciousness, then if particular sub-communities are responsible for logical control of the community and its data, how do we logically deny their having a literal, genuine consciousness of some sort?  Or even a number of partly or wholly independent consciousnesses in a single community?  Equally we cannot logically assert it, to be sure, but the importance of the implications, or the importance of the implications if we can show the speculation to be wrong, or even the importance of our contemplating the problem and exploring means of investigating the alternatives, forbids us to dismiss the concepts too facilely. 
In general then, discussion of subjective consciousness and its biological significance is fraught with difficulty.  We need a major scientific breakthrough, analogous to the first advances in the study of electricity.  Such a breakthrough would have to be on the scale of the first recognition, measurement and generation of electric current, before we could study the matter meaningfully.  At present we are no better equipped to discuss the nature of subjective consciousness, than we were to discuss the nature of disease before the invention of the microscope.  (And don’t be impatient, the germ theory of disease took some 200 years to develop after the first microscopic observations of bacteria and cells.) 


Stages and aspects of sociality

If an animal does something they call it instinct.  If we do exactly
the same thing for the same reason they call it intelligence. 
I guess what they mean is that we all make mistakes, but that
intelligence enables us to do it on purpose. 

Will Cuppy
One major value of the discussion of the intelligence of communities is that it throws the fallacies of composition and division into stark contrast.  Works such as “The Mind’s I”  by Hofstadter and Dennet have elegantly illustrated some of the problems, and Hofstadter’s “Ant Hillary” in Goedel, Escher, Bach” showed some others.  In nature we have cases where an intelligent system, such as the brain, consists of neither particularly nor obviously intelligent units, such as neurons. 
Conversely it is hard to tell how much intelligence a human community has, and how to compare it with the intelligence of any component human, let alone each component human.  What about idiots and geniuses who happen to be members of the same population?  Can we meaningfully compare the intelligence of the community with the mean, modal, or extreme intelligence of its individual members?  Does one measure community intelligence in terms of the technology that a few ingenious members have implemented and the rest hardly understand at all?  Or in terms of the infrastructure that a major sub-community have established?  If so, does it matter whether the infrastructure was imposed by a few in authority, or by conscious co-operation, or spontaneously by not consciously intelligent mass behaviour? 
If the community is less intelligent than its sentient members, then how does the intelligence of a honeypot replete or of a mobile worker ant or bee compare with that of its community?  How does the intelligence of a worker or queen termite compare?  How does the intelligence of a hive compare with that of a worker? 
The replete really is just a worker ant that has developed a huge crop that fills its abdomen with stored syrup.  Once it has achieved that stage it requires little more intelligence than is necessary for accepting or dispensing honey, plus hanging from the roof in a suitably comfortable chamber.  The queen started with enough intelligence for her mating flight, plus establishing a nest and feeding the first generation of workers.  Once she has settled in though, she has little to do but lay eggs.  We know that worker bees’ brains change in shape and size as they change tasks.  It is not implausible that these other job-changing insects have equally flexible brains. 
Apart from the question of the relative intelligence of different castes within a colony, how does the intelligence of a worker bee compare with that of a fly?  Houseflies are perhaps less inclined to get trapped behind glass because they are less stereotyped in their positive phototaxis, but not many of their behavioural patterns are nearly as elaborate as those of bees.  And yet the behaviour of either insect shows little sign of insight or of anything but stereotype. 
This problem expands to match other dimensions.  How does the intelligence of a species compare with its members or with communities of its members?  When two species form an association vital to both, such as a specialist pollinator and its associated specialist plant, then relatively, how intelligent is say, the yucca moth, the yucca, or the association of the two?  How intelligent is the mitochondrion in the human, relative to the neuron, the brain, the nation, or the species?  It is not obvious how far one should stretch such boundaries.  Do we include domestic animals as members of our communities?  Inquilines?  Pests?  Crops?
It is extremely difficult to compare the intelligence of different species meaningfully, let alone the putative intelligence of incommensurables such as individuals and communities, or devices and organisms.  We desperately need a cogent science of the systematics of types of intelligence, and measures of the various types of intelligence before we can talk sensibly of such matters. 
We also need some measure of entity-hood.  Formal fuzzy logic hardly begins to address the problem. 
Now consider some  levels of sociality and associated intelligence.  Note carefully that they are not all independent and that they do not form a clearly ordered sequence.  The problem is that apart from the poorly defined nature of intelligence, it is a multidimensional phenomenon or range of phenomena.  

Hypothetically independent organisms. 

When in that House M.P.s divide
If they’ve a brain and cerebellum, too,
They’ve got to leave that brain outside,
And vote just as their leaders tell ‘em to.
But then the prospect of a lot
Of dull M.P.s in close proximity,
All thinking for themselves, is what
No man can face with equanimity.

W. S. Gilbert  Iolanthe

One might expect that populations of socially independent, parthenogenic individuals should be unintelligent both as entities or communities, but it is soberingly difficult to find clear examples.  Even in many kinds of bacterial community there are interactions that look suspiciously like altruism or organised control or competition.  Bacteria not only are brainless, but lack the necessary complexity to form somatic structures the way the eukaryotic cells of metazoa or metaphyta can. 
The principle extends still further.  Viral communities seem to work in unison to cause effects such as diseases in hosts, or symptoms that aid the spread of the pathogens.  For just one example, rabies viruses, lacking any clear socially interactive behaviour, cause neurological damage in hosts, such that it causes varied and elaborate behaviour that propagates the virus.  They are by no means the only pathogen to cause analogous behaviour that harms the host but propagates the parasite.  There are whole classes of such behaviour among pathogens. 
Conversely, many other diseases, most notoriously myxomatosis, rapidly evolve into a less malignant relationship with the host, not out of obvious altruism, but because such aetiology maximises its infective success.  Selection for the success of pathogens works purely on effectiveness, not specifically the harm or the health of the host. 
How intelligent are such pathogenic species as entities?  Where does the information processing of such a species reside?  As individuals the microbes display nothing resembling intelligent behaviour that is not stereotyped and mechanistic.  And yet species that do not have any such effective aetiological strategy, tend to die out or at least become less successful as pathogens.  From the point of view of the empirical observer, how do we logically distinguish this from intelligent behaviour of the community? 

Sexually reproducing species

The popular type and exponent of obstinacy is the mule,
a most intelligent animal.
Ambrose Bierce  --  The Devil’s Dictionary

Most forms of sexual reproduction entail apparently intelligent behaviour, usually in the persons of its individual members, or they could not bring their gametes together.  In microbes such behaviour is stereotyped, even mechanically controlled, and also in many metaphyta and metazoa. Any sexually reproducing species also must exhibit intelligent behaviour as an entity, or its members could not exhibit effective reproductive behaviour. 
To illustrate just how extreme the contrast may be between the functional intelligence of a species and that of its members, consider the flowering plants.  The individuals are at most arguably intelligent, but the majority employ sexual reproduction, including strategies that are intellectually baroque to put it mildly.  Such selection does not come cheaply in evolutionary terms because within any species the reproductive apparatus tends to be very conservative.  This makes sense because the apparatus is an example of where two sets of components must not only work individually, but must work in combination.  If they do not, the line stops there. 
This principle that systems comprising complementary components are generally conservative, is of vital evolutionary significance.  We see it in major ways.  Firstly, without complementary mechanisms we simply do not get complex organisms.  Just to form bodies requires cells of considerable complexity; no prokaryotes have managed it yet, let alone any viruses.  Even for eukaryotes the development of cells that can be assembled into bodies is a challenging task.  The mechanisms necessary for the formation of bodies of metaphyta and metazoa are startlingly complex and specific.  They rely on elaborate cell structures plus elaborate molecular coding for intercellular matching, signalling, binding, and release.  Each of these four is at once complex and vital; let even one fail in just one type of tissue, and the consequences can be anything from serious disease in the adult, to death of the developing embryo. 
In assessing the non-triviality of the problem of the evolutionary development of such complementarity, reflect that the best current estimates of the length of the period it took for the original metazoa and metaphyta to evolve, exceeds the length of their entire subsequent history so far. 
Another example of the problems of the development of complementary biological mechanisms is the development of the eukaryota from prokaryotes.  We do not know how long it took, but it clearly was not abrupt.  
Still more radical was the problem of the development of intracellular organelles such as the ribosomes.  Once having formed they remained almost unchanged for probably more than three billion years.  And what is more, they look like staying that way, short of some really aggressive genetic engineering for as yet obscure purposes. 
The conservativeness of sexual parts cannot compare with those truly ancient examples, but it none the less is pervasive and stems from the same logical problem.  It is in fact so pervasive that among fungi and flowering plants the comparative anatomy of the sexual parts present far and away the most important taxonomic features.  Among insects the situation is not quite so extreme, but still strongly marked.  Insect genitalia are of great importance in recognising many groups.  Though it is not wise to generalise too simplistically, there are good reasons for this. 
The requirements of a sexual system vary with the nature of the organisms.  Plants are generally sessile and depend on mobile agents to move the pollen to the stigma.  Commonly these are birds or insects, which means that not only must the mechanics of pollination work properly, but the movement of the pollinator must be guided as well.  These factors limit the rate at which flower architectures can be changed without losing their effectiveness. 
In insects and in general among animals with small brains and highly stereotyped repertoires of activity there are different reasons for highly conservative mating anatomy, physiology and behaviour.  Although their behaviour is not strictly rigid, they cannot be allowed too much initiative in so critical an activity.  Often they get just one attempt and if they miss that, they have just become selective failures.  At the same time the challenges are formidable, far too much so to rely on the intellectual powers of their brains.  Instead they have developed highly specific senses, anatomical features and scripts for their mating, and the shorter their adult lives the more specific their scripts are likely to be. 
Usually the first attraction is chemical.  It serves for identification and attraction.  This sounds simple, but in fact recent work has greatly multiplied the range of compounds that insects are known to use for signalling.  In guiding mating activity, some wasps use both visual clues, plus at least seven successive compounds for control of seven successive stages of the mating act.  Amazingly, some orchids rely on those male wasps for pollinating them by trying to mate with them, and even more amazingly their flowers not only resemble the female of the relevant bee or wasp, but also produce all seven chemicals!  The very fact that the orchids had time to evolve and specialise in the use of such an elaborate system does imply that the wasps had been using the same system for a long time, probably many millions of years. 
The reason for the multiple attractants is that each one triggers a particular activity.  One might be a long-range attractant, another short range, the next a contact guide, another might stimulate genital contact, and so on.  In experiments it has been shown that omission of any one signal can interrupt the mating script at that point.  For all we know there are other insects whose chemical mating signals involve even more compounds.  The point is that where each signal is so specific, and their sequence so rigid, it enables the species to rely on the script for automation of the process, instead of the discretion of the insect.  Lorenz has remarked that in fighting fish the courting procedure is intense, not to say violent, but the actual fertilisation is apparently dispassionate.  This is true, and not only in those fish, though of course we have no idea whether such behaviour reflects actual states in the mind of the fish, if it has one. 
There are several variations of mating procedures among eusocial creatures.  In naked mole rats it is not unlike a lot of other mammals in which only alpha members of the community are allowed to reproduce. 
In many eusocial insects there is a nuptial flight and the males and females seek each other out and pair off.  In many of ants the flying reproductives form pillar-like clouds, much like the clouds of courting, non-social Empidid flies, and for similar reasons.  Being two-gender affairs, the visible clouds are not really leks, but like leks they attract all the locally available candidates. 
Most species of termites do not form such clouds, or not so long anyway.  Possibly this has to do with the fact that termites are more attractively edible.  Be that as it may, the participants tend to disperse as soon as possible and the females generally attract males by scent.    Once they have met and shed their wings, the pairs go off and the male joins the female in establishing a nest.  He typically remains with her for years.  Perhaps this is partly to ensure an adequate supply of sperm for a female that in some species might need to reproduce for decades. 
Honeybees are notorious for the single mating flights of drones.  It has long been known that the drone mates at most once in his life, and some authors have over dramatised the event.  They claim such things as that the female also mates just once and that she tears the entrails out of he single successful suitor.  In fact a young queen typically mates several times, once on each successful mating flight.  She certainly does typically return with mail parts trailing behind, but that is a consequence of the mating process, not any particular impulse to female sadism.  Instead, what happens is that at the moment of coupling, the mail orgasm contracts every muscle in his abdomen with such violence as to expel his genitalia, kill him instantly, and leave his corpse feeling peculiarly dense, almost like a little pebble in one’s hand. 
One effect of this is to leave the female with a clogged genital passage, so that she has to return to the hive and clean up before she can mate again.  This ensures that no subsequent suitor can displace the semen, so that no doubt the successful drone did not mate in vain, even if he shares sperm space with his rivals. 
In any case, since the queen does not mate again after her initial nuptial flights, it makes sense for her to stock up on sperm, since she needs it to produce workers as well as fertile daughters.  Incidentally this peculiarity of the honeybee mating process has led to a major debate concerning genetic conflicts of interest among the workers and their siblings.  On average they generally are less closely related than full sisters.  I do not explore the implications here, but mention the point as yet another illustration of the complexity of the factors that influence the strategies of the species. 
Sexual strategies are generally species-determined; even humans only observed the connection between sex and reproduction a few thousand years ago.  There are examples of intelligent animals practising sneaky sex, but there is no evidence that the proximal objective is anything but the mating.  In less intelligent species such as fish and lizards one encounters some examples of systematic sneaky sex, but those seem to be just as stereotyped as direct sex.  There also are many cases of where one male will frustrate the sperm or kill the offspring of another, but the elaborations of sexual strategies are too varied and too many to discuss in detail. 
The important point is that nearly all the sexual strategies in nature are strategies of the species, or at least of particular genetic lines, rather than of individuals or communities.  

Mutually interdependent species

A brother may not be a friend, but a friend will always be a brother.
             Benjamin Franklin
Not only do communities of species that become mutually interdependent exhibit their various individual apparently teleological behaviour, but the individual species that form components of the communities have their own teleological behaviour and so do the inclusive systems involved.  However, it is not at all clear that there is any question of individual teleology in the behaviour of particular participating specimens, or that if there is, it corresponds to the teleology of the species.  Examples include:
*      Symbionts such as cleaner wrasse or oxpeckers
*      Stranger-than-fiction symbioses such as “The medusa and the snail” discussed by Lewis Thomas in his essay of that name.
*      Endosymbionts such as mycorrhizae or organelles in eukaryotic cells.  A truly mind-stretching range of examples begins with the endosymbiotic bacteria that developed into mitochondria.  They tend first to lose genetic material whose function is redundant because the nucleus can supply it.  Some go further when some of the genetic material from the endosymbionts finds its way into the nucleus and in turn becomes redundant, after which it might get lost from mitochondria as well.  After such process have gone far enough, the mitochondria are left with very few genes indeed.  The details of the next question remain open to argument, but it seems that at least some classes of hydrogenosomes are descended from mitochondria, and most hydrogenosomes have lost all their genetic material.  Truly a grin without a cat! 
Now, this too is a very difficult concept to come to terms with.  It might at first seem simple, but it leaves us with difficulties in defining conflicts of interest, evolutionary strategy, and identity.



Rudimentary intraspecific tolerance 

How far you go in life depends on your being tender with the young, compassionate with the aged, sympathetic with the striving and
tolerant of the weak and strong. Because someday in
your life you will have been all of these.

George Washington Carver

Within any species a crucial step toward sociality is mutual tolerance for purposes of reproduction.  This is necessary, first for mating, second for sparing offspring instead of eating too many of them.  Innate tolerance creates evolutionary opportunities for developing altruistic strategies such as sib care in foxes, scrub jays, carpenter bees, and paper wasps. 
Mechanisms for such tolerance are varied.  Some species manage it by smell; they recognise unacceptable company by the fact that it does not smell like themselves.  This is very common in social species.  Others recognise their own neighbours as individuals.  This is not common in eusocial species, but sociable species, especially birds and some mammals do that. 
Another mechanism is that some species simply lose their appetite at critical seasons.  Some reptiles use such tolerance for seasons when they are likely to hibernate together.  Other species, particularly birds of prey, lose their aggression to other birds in the neighbourhood of their nests.  Thus we might find a hawk ignoring the nest of a dove in the same tree as its own nest. 
Territorial species such as some chameleons, may have young with different coloration from the adults.  Such adults, such as those in the genus Bradypodion simply ignore their own tiny putty-coloured young.  Conversely many creatures that exhibit no special parental care, actually eat any of their own young that they can.
There are other classes of examples, but the point is that eusocial species need just such tolerance in one form or another, or they could never have developed any successful sociality.  Many of them are after all not just territorial, such as naked mole rats, but predatory, such as social spiders and paper wasps. 


Interspecific and intraspecific flocking

It is easier to love humanity as a whole than to lover one's neighbor.
               Eric Hoffer 
Similar innate tolerance may arise advantageously between neighbours, such as Bembix wasps or solitary bees nesting close together in earth banks, or seabirds nesting in tight hexagonal array, or indeed, the flocking or shoaling of fish. 
Mammals and birds often form mixed herds or flocks for purposes of security, feeding and socialising.  There may be squabbles between neighbours, and problems with parasites, in fact, there are thousands of parasites of flocking species.  The hosts of such respective parasites range from locusts and fish to starlings and cattle.  In the (human) world wars, they also included convoys of ships trying to beat off submarine attacks. 
Seeing the dreadful costs of attacks on flocks, one can hardly believe that there are advantages to presenting such concentrated targets, but in most cases in nature (and in shipping convoys) the advantage of improved security exceeds such costs.  It is only when the predator is large and powerful enough to destroy the target wholesale that flocking is a bad strategy. 
The behaviour of the flocking community may be largely selfish, for instance, when attacked, each herring tries to hide behind its neighbours, and the results might temptingly resemble neurosis or other dysfunctional behaviour.  For instance, whales or large predatory fish herd schools of small fish into effectively solid masses for easy eating.  In such terms both communities, attackers and attacked, resemble individual entities, or even single organisms. 
The effectiveness of the swarm tactics can be calculated on operational research principles, or it can be seen directly in footage of birds of prey attacking flocks of bats or prey birds, or fish attacking shoals of pilchards or herring.  Not many attackers will venture into the flock.  The operative reasons for their shyness include the risk of injury and the counter-intuitively low probability of success.  Each prey organism avoids the attacker individually, while conversely the attacker needs to fix onto a single prey item and finds it difficult to do so. 
The true effectiveness of such flocking as a defence becomes most obvious when it is subverted and fails.  A notorious example is that if one marks a herd animal, such as a gnu, for study in an environment of heavy predation, it is difficult to do so without dooming it.  If the predators can recognise the marker, it enables them to concentrate on that individual.  Normally hunters will harry a herd till they can identify an individual by some peculiarity, and this is doubly effective because the most attractive of such peculiarities are those that indicate that the candidate prey is not well and is easier to catch.  However, in the light of the observation that even healthy animals fall to predators once they are marked, it is clear that as a rule, identification is more important than weakness. 
Conversely, there are potential advantages to the flock, that they might cooperate in their defence.  In its most sophisticated form this is strangely rare as a strategy, presumably because it is difficult to implement effectively, and it is costly to any individual if it relies on support and gets let down.  Such factors militate against the development of the strategy.  Buffalo weakly apply herd defence against lions, but if they did so systematically and aggressively, they would be effectively  immune to predators.  Musk oxen demonstrate the fact in their famous defence against wolves, in which the adults, especially bulls, form a protective ring around their young.  Except against humans with dogs, that defence is practically impregnable.  Presumably in the bleak, ice-age north, wolves wiped out most herd species that failed to develop such a defence. 
Among eusocial insects a different principle is added to the relevant variables.  The workers or soldiers are not generally reproductives.  They have little to lose by being killed in defence of their gonads and offspring, that is to say, the gonads and offspring of their reproductive castes.  Thus it is that we find ants, termites, wasps and bees aggressively defending their colony or attacking formidable prey, at the cost of their own lives.  Most of them do so in an apparently unorganised fashion, but even then it often is more organised than it seems.  The majority often will not bite or sting till they smell the attack pheromone released by the first hive mate that is injured or provoked.  By that time many warriors are likely to be in position, and they attack together from all angles.  This is far more effective than just having the first bee that gets into in position attack at once before the others can support it.  Bees and many species of ants are likely to attack in such a way.
Usually when social species attack one at a time, it is because individuals have little to defend, or they are very formidable, so that a single sting or bite is likely to rout the attacker.  Bull ants and some paper wasps are examples. 
Apis cerana offer an example of coordinated attack in their balling behaviour, by which they kill scouting giant hornets, as I discuss elsewhere. 
Notice that all these examples might be seen as illustrating the intelligence of the community; certainly there is very little evidence of any of them arising from the ingenuity of the individual, not even of any particular leader.   


Aggression inhibitors (Tolerance releasers?)

Notation is a tool of thought
A very important behavioural component in the origin and development of mutual adult tolerance and care for offspring, let alone sociability and eusociality, is context-sensitive inhibition of aggressive behaviour patterns.  Lorenz was the major pioneering writer on the subject, and his writings on innate, injury-limiting conventions for intraspecific fighting were seminal.  
Such tolerance is innate, and is widely spread, both within species and even interspecifically in non-colonial species such as hawks that will not attack prey birds nesting close to their own nest.  Bembicine wasps hunt flies and carry their prey into the tunnels in which they raise their young.  Parasitic flies wait around the nests for opportunities to oviposit on the wasps’ prey when they carry it in.  Observers often have wondered why these fly hunters do not simply catch those flies as well.  It might be worth investigating whether the wasps are not simply similarly inhibited from attacking flies so close to their own homes. 
In any case the importance of such innate inhibitions is that they can serve as a basis for co-operation in both interspecific and intraspecific relationships.   


Episodic assembly

Some therefore cried one thing, and some another:
for the assembly was confused: and the more part
knew not wherefore they were come together.

Acts of the apostles 19:32

Otherwise independent non-social individuals of many species assemble on occasion, and in many ways and for many reasons.  The simplest examples meet just for mating, but in practice this involves visual, chemical or auditory signals that might assemble many more than two, such that one sex or another has a choice of mate.  Males might display at an assembly point, such as a suitable tree for birds of paradise, or open space for grouse or ruffs.  Such assemblies are called leks. 
An interesting aspect of lek behaviour is that as a rule the assembly forms by positive feedback.  Some species of cicada males that feed in trees and sing to attract females, prefer trees where there already is a hubbub of rival males.  This might seem counter-effective, but firstly the females are prone to go where the boys are, and choose a male where the communal song is at its loudest, and secondly, the males in combination are subtle in misleading predators; an approaching noise causes the nearer males to soften their song or even go silent, so that a stalking predator’s direction finding equipment systematically loses the nearest target and fixes temporarily on a more distant target.  It is a fine example of how simple, innate activity on the part of individual members can in effect combine in a sophisticated collective strategy. 
Some species of firefly males assemble where they see the most light flashing synchronously in their specific patterns.  The females accordingly are attracted and can select mates.  Interestingly, other species of firefly females also assemble at such leks, but their objective is food; they flash the feminine come-hither, and grab and eat any male that responds.  This is obviously an ancient strategy, because research has demonstrated that there has been an evolutionary arms race, with victim species evolving ever more complex signals, and predatory species evolving correspondingly complex  responses.  
Another class of assembly is common among insects.  Some species will fly to a prominent object, where they will find others with similar behaviour.  For example, some beetles fly to the highest hill in the horizon.  Empidid flies of many species assemble over local items, such as a high nearby bush or rock.  Sometimes they even assemble in a cloud above a slowly moving human or large animal.  Some kinds of ants on nuptial flights assemble in a similar swirling cloud above the nest from which they emerged, until they have attracted their mates. 
A more direct basis for assembly is where there is food or a nesting site.  It is an impressive sight to see the huge dung beetles that specialise in elephant dung, with their antennae spread, circling in like so many vultures, not only to food, but to where they can find mates. 
By assembling according to common or complementary stimuli, these organisms are exploiting the principle of “Do you come here often?  Small world!”.  The seminal, but simplistic Milgram experiments suggested that human society was “shallow” in that the friend-who-knows-a-friend chain between any two people on Earth is likely to be short, probably less than nine. 
Importantly however, further work shows that part of the reason for this turns out to be because humans are not a random network of acquaintances, but inclined to homophilia; that is to say, birds of a feather flocking together.  Also, even in an otherwise random network, just a low frequency of long-range acquaintances shortens the maximum depth of the community drastically. 
By thus selecting or creating assembly points that attract participants from afar, the species not only increases the chances of successful mating, but also of mating with a low frequency of inbreeding.  Some of the assembly techniques more or less force outbreeding.  For example, some ant nests release only one sex of reproductives at a time and they accordingly are forced to find mates from other nests. 
Empirically such assemblies are most intelligent strategies.  The subject is a large one however, and there are many counter-examples in nature. 

Rearing care and defensive behaviour releasers

. . .And how can man die better than facing fearful odds,
For the ashes of his fathers and the temples of his gods,
And for the tender mother who dandled him to rest,
And for the wife who nurses her baby at her breast. . .

T. B. Macaulay  Horatius
Going beyond simple intraspecific tolerance, many of the behaviour patterns in sociable and social species are plainly developments of behaviour patterns that one might get in solitary species that exhibit rearing care behaviour.  For example, monkeys of many troop-forming species will compete to handle the young of other members of their troop.  Carpenter bees and paper wasps care similarly for young in their nests, whether those young are their own or not.  Some birds will accept the young of other species in their nests, though others will kill or eject them.  Mammals in their lairs have often been observed to feed young of other species. 
Defensive behaviour for protection of the colony is mainly distinct from defensive behaviour of solitary species in that it is less constrained.  For the genes of the soldier castes, it is more important that the colony survive, than that the individual survive.  In contrast, defensive behaviour in solitary iteroparous species usually is limited to conflicts which the defender might be expected to survive, because it is more important for the parent’s genes, that the parent survive to raise more young, than that any one threatened young survives.  For complementary reasons, in some ways the same, in some the opposite of the soldiers, semelparous species may be less compromising.  For the genes of the parent that will not propagate again, it is more important that the brood survive than that the parent survive.  Similarly, eusocial castes might exhibit cleaning or foraging behaviour similar to that of solitary species, but more specialised or intensive. 
Another interesting example of the use of castes for special defensive purposes is in the minor workers of some leaf cutter ants.  The major workers sally forth to collect pieces of leaf.  They are accompanied, or even ridden, by minor workers who do no leaf cutting.  On the return journey the minor workers ride on the leaf sections, “parasols” carried by the major workers.  What they are doing on the trips is not easy to tell, because their main duties are in the nest, tending fungus gardens and the like.  In spite of the cute speculations of early workers who studied leaf cutters, it since has turned out that the minor workers are not passengers, but “riding shotgun”. 
The function of these minor workers on foraging sorties is to repel parasitoid phorid flies.  The flies otherwise would lay their eggs in the major workers, so that their larvae could eat them alive from within.  This employment of the minor workers makes sense in simple economic terms.  To the nest, the cost of raising a minor worker is at most a few percent of the cost of a major worker.  They are so cheap that a minor worker need protect a major worker only slightly to justify its trip.  I have more to say on this point later.   


The evolution of matching signal conventions

Gnosin damazon, dedamakas panta
(Master knowledge, master all.)
There is a serious evolutionary problem to the evolutionary development of specific recognition signals.  I first saw this point raised by the late E. F. Whiteside, as described in the attached paper on the evolution of pheromones.  In summary, the question is: in any genetically determined, functionally vital, and specific signalling system, any deviation from the canonical signal must be seriously disadvantageous, because such a deviant signal would be ineffective.  If its signal did not match that of a mate, an emperor moth with the wrong sexual lure might s well be sterile because it would attract no males.  How does it come then, that we have more than one species of emperor moth, which in fact we do?  There are dozens known from Africa alone, and thousands of relatives world wide. 
As it happens, I can propose a mechanism in which classes of highly specific signals can evolve, as long as the receptor as well as the signal are determined by the same genetic system. 
This might be seen as part of the intelligence of the species as a community.  The principle is worth noting, but is not highly germane to the current issue; hence the attached article on pheromonal evolution.  


Eusocial colonies and familial groups

It is only to the individual that a soul is given.
                 Albert Einstein 
I know of no eusocial species in which a colony is not derived from a single nuclear family, though in some species the queen (mother) may mate several times, and there may be a succession of queens, sometimes even several reproductive queens in the same nest simultaneously.  The one clear class of counter example is not eusociality in the sense intended in this discussion, and that is in endosymbiotic or near-endosymbiotic cell colonies such as lichens, algae, sponges and humans. 
Such endosymbiosis occurs in numbing variety; it ranges from unrelated species of cells that interpenetrate to exchange materials or signals, to organelles in cells that simply were not suspected of being separate species till well into the mid twentieth century.  The most obvious examples are chloroplasts and mitochondria.  The suggestion that they might be endosymbionts at first was widely rejected, but once the evidence became incontrovertible it caused a sort of witch hunt, or perhaps a gold rush.
Suddenly zealots saw endosymbiosis everywhere and labelled practically every identifiable cell structure as a relict endosymbiont.  Flagella, cilia, ribosomes, nuclei, they all were proposed by various enthusiasts.  Some proposals were reasonable, some simply silly, but none detracted from the fact that cells and even multicellular organisms of various types, related or unrelated, could connect and combine in amazing ways for amazing functions. 
It also became obvious in retrospect that symbiosis and in particular endosymbiosis offered the most powerful means by which natural selection could defy the combinatorial improbabilities of simultaneous development of biochemical tools that had developed independently in unrelated organisms.  Simultaneous development of adaptations places an enormous selective burden on even the largest populations. 
We cannot explore every development of this type.  It is the subject of whole sets of books.  However, it is closely related to the subject of this essay.  There is good reason to suspect that the principle of combining independently developed functions involves as much of the mechanisms constituting community intelligence, as co-operation among family members. 
It would be conceivable that eusociality could arise between conspecifics of different parents, or even in multiple species, but obviously that sort of thing would increase the range of genetic conflicts of interest, so that there are probabilistic obstacles to the evolutionary development of a stable relationship of such a type.  In practice it is very difficult to find clear examples where this actually has happened.  Inquilines usually turn out to be either parasites, scavengers or commensals.  In the case of ants, some parasitic species of ant have graduated to behaving as actual predators.  
Conversely, all the most obviously social or eusocial species known, could easily be seen as having arisen from ancestors with highly developed nuclear family care.  


Systematic reaction to the environment

Societies need rules that make no sense for individuals. For example, 
it makes no difference whether a single car drives on the left or on the 
right. But it makes all the difference when there are many cars!
               Marvin Minsky

In the sense I have in mind, reaction to the environment falls into two main categories that are important in the current discussion.  Both might easily be overlooked because in solitary species we take such behaviour for granted, but there are intriguing analogies to social species when one thinks of them in terms of apparent teleology. 

Establishment of the colony

Brute force crushes many plants. Yet the plants rise again.
The Pyramids will not last a moment compared with the daisy.
And before Buddha or Jesus spoke the nightingale sang, and
long after the words of Jesus and Buddha are gone into oblivion
the nightingale still will sing. Because it is neither preaching nor
commanding nor urging. It is just singing. And in the beginning
was not a Word, but a chirrup.

D. H. Lawrence
There are many ranges of ways in which social insects establish colonies.  The most primitive are presumably those that most resemble the behaviour of solitary insects that patently exhibit parental care.  The most obvious examples of such insects include solitary bees and wasps and some kinds of cockroaches, but one could easily extend the category all the way to the even simpler behaviour of insects that lay their eggs only on suitable food plants or in suitably secure situations. 
Other forms of parental care include the raising of young to maturity, then dispersing them to mate and establish new colonies of their own.  Sometimes this requires the young to carry quite elaborate resources, ranging from internal body fat, to cultures of the necessary symbiotic micro-organisms.  Still others rarely send out reproductives on mating flights, but instead raise many active reproductives in the colony, and keep scouting for new nesting sites to take over on foot and without nuptial flights.  The Argentine ant for example, rarely has fights between nests, but forms huge colonial complexes, empires if you like, that include indefinite numbers of reproductives and workers in nests scattered over large areas and connected by trails.  When a nest becomes undesirable, there is likely to be a lot of carrying of brood to neighbouring or new nests. 
Not to multiply examples however, modern solitary bees and wasps include examples of behaviour that no doubt is functionally similar to that of the ancestors of many modern eusocial species.  Typically, the young, mature female will seek out a spot suited to building the home or digging the burrow where she will collect the food and lay the eggs.  Obviously if she never gets this right, her line ends there.  
In practice, there are thousands of extant species that routinely do get this right.  They include borers, miners, builders, colonists, parasites, opportunists in bewildering variety, but they all share one attribute: if they select the site unsuitably, or prepare or maintain it unsuitably, that too is the end of the line.  And such behaviour requires intelligence, in the sense that the parent must react appropriately to environmental stimuli. 
Nest selection is no simple task.  Safety from various enemies is one requirement.  Suitable moisture and temperature control are vital.  A large percentage of nests fail on such points.  It takes some doing for a human to judge the viability of a new nest, but as a rule the brood mother does pretty well. 
Here it does not matter a bit whether the parent is subjectively intelligent in herself or whether she is the purest automaton.  The same minimal amount of information collection and processing is necessary in either case.  In the context of this essay that information collection and processing is what matters, and in practice the behaviour of the parent can only be distinguished from subjective intelligence by such experiments as Fabre would perform to demonstrate the limits of their rational reactions to abnormal circumstances. 
This is a difficult field in which to maintain good perspective.  One is inclined to see the silly wasp demonstrate its lack of mechanical understanding of the situation, and dismiss its intelligence accordingly.  However, that is a treacherous line of reasoning.  Dogs for example, which we accept with good reason as being more intelligent than wasps or bees, will also reveal a drastic lack of such insight in analogous circumstances. 
However, none of that matters in our context.  The point is that the intelligence, if we meaningfully can call it that, in such a case is intrinsic to the species, not the individual.  Now, on that assumption we can move on from the solitary bees to say, some carpenter bee species, in which instead of keeping the larvae separate, they may be raised in communal cells and when they are mature may even assist in raising siblings. 
The wasp equivalent is in paper wasps, such as Polistes or Belonogaster, in which the queen establishes a nest in which the first young are likely to end up as workers.  In some species, when the first workers reach maturity the queen stops hunting, lets the youngsters forage and stays home to lay eggs. 
In most ants and termites the mated queen establishes a nest, with or without the male, depending on the species.  Usually this will be a burrow or a hollow under a stone, but many species live under bark, in leaves and other situations.
As I said, such behaviour requires decisions that would challenge human judgement, and what is more, it is judgement that must be made in the context of external circumstances.  In other words the behaviour must be flexible, adjusting to challenges of resources and adversity. 
Simultaneously however, there is a great deal of stereotyped behaviour of great sophistication.  Sometimes queens will carry necessary inquilines or microbes from the parental colony, from which they can establish the necessary cultures of food or other functions.  In at least the genus Carebara, the flying queen takes along a few tiny workers that cling on to the huge flying queen as best they may.  No doubt they are valuable, possibly absolutely necessary, in establishing the new colony. 
Such examples of behaviour have effectively nothing to do with the individual intelligence of the queen founding the new colony. 
Not to labour the point too heavily, this all amounts to intelligence in the species, if not always in the individual.  In their respective ways, each of the colony, the species as a whole, and individuals, may be seen in the light of dedicated, task specific, but flexible information processing structures.   


Maintenance and management of the colony

The work of construction seems to be a sort of balance struck between many bees, all instinctively standing at the same relative distance from each other,
all trying to sweep equal spheres, and then building up, or leaving ungnawed, the planes of intersection between these spheres.  It was really curious to note
in cases of difficulty, as when two pieces of comb met at an angle, how often
the bees would entirely pull down and rebuild in different ways the same cell, sometimes recurring to a shape which they had at first rejected.

Charles Darwin.                       On the Origin of Species 
The next phase, after establishing the nest, is maintenance.  In many solitary species, the need does not arise, because the adult abandons the completed nest.  However, there also are many species that build just one nest or burrow, in which they make separate cells that develop over an extended period.  Such homes may need a fair amount of maintenance during their period of occupation. 
However, it is mainly the social or eusocial species that really invest heavily in nest maintenance.  Termites are probably the most spectacular examples.  Though it is hard to get precise data, the mounds of some species may weigh tonnes and remain in use for more than one century.  Probably most species’ mounds are in use for a decade or more.  One of the major factors determining the longevity of a mound that survives the thousand natural shocks that termite mounds are heir to, is the longevity of the queen, and if that is too limiting, then of the secondary reproductives that finally replace the female that founded the nest. 
During its occupation a mound is constantly remodelled, with most of its concrete-like structures being temporary.  The most enduring parts are the outer walls, but even those get remodelled.  In the building season, usually after rain, worker termites tunnel to the surface and begin to construct new, soft, thin-walled chambers on the outer wall.  As these new shells harden, they get reinforced till they become the lasting new outer walls.  Then the old thick walls, progressively softened by the moisture of the internal environment, get perforated with new chambers and tunnels.  They soon become indistinguishable from the rest of the internal spongy structure of the nest. 
The mound also gets restructured internally as needs change.  Sometimes changes adjust the overall shape of the nest, for instance, in compass nests.  Sometimes there is tunnel digging or tunnel building to look for food or water supplies.  In some regions termites tunnel so deeply for water, tens of metres at least, that minerals in the walls of their mounds have been used successfully for mineral prospecting on an industrial scale. 
Furthermore, there may be substantial remodelling to establish or adjust food storage or other structures, such as fungus chambers and living space for queens, eggs and young.  The queen in particular cannot be shifted without a lot of serious restructuring of partitioning and passages. 
These last examples are particularly significant in the discussion of nest intelligence.   They entail both homeostatic adjustments to the internal environment, and modifications to accommodate new or seasonal requirements.  In other words, to meet the needs and “comforts” of their various sub populations, they require the workers to respond both to the internal and external environments, and to apply the appropriate negative feedback in their construction.
Nor are these static requirements.  The homeostatic requirements of the colony change all the time, according to season, weather, food supply, threats, damage, or changes in the surroundings, and anything of the kind requires building or remodelling activity.  It is easy to imagine the workers migrating from one part of the nest to another as things become too hot, or cold, too soggy or too crowded, but they do more than that; they move eggs and food, and thicken or replace walls to suit other things than just their own comfort.  In those species that have air conditioning requirements, the maintenance can be quite obscure, including convection chimneys whose functions humans simply did not at first understand. 
Special reactions are necessary on occasion, for example when it is time for the mating flights of their reproductives.  These are particularly demanding, because they should coincide with the mating flights of neighbouring nests.  The more neighbours that fly at the same time, and the shorter the period during which they emerge, the better the chance for outbreeding, and the better the chance that the flying termites will survive by overwhelming the predators that gather to feed on them. 
As an intelligent information processing system, such a colony compels respect, whatever the intelligence of its individual members. 
Ants are not far behind termites in these respects, and various species probably must accommodate even wider ranges of needs of tenants, eggs, larvae, pupae, and inquilines such as aphids.  Some of them also must suit the needs of the plants they live in.  In all cases they do not place items arbitrarily.  Unlike termites, they have immobile pupae to care for, and practically immobile larvae as well.  Some species also have repletes, gorged workers that form effectively a separate caste.  The job of repletes is to store food, and they hang upside down with bulbous abdomens filled with syrup.  They too are practically immobile and must remain in a part of the nest that keeps them healthy.  And like termites, some species of ants rely on fungus gardens that require special climatic conditions.  Ants apply less obvious air conditioning engineering than termites, but some species probably do apply the venturi tube principle to air exchange. 
Eusocial bees vary in their nesting behaviour.  Most hives are in hollows in rocks, earth, or wood, but some build combs in the open.  However, the hive does require adaptive climatic control depending on the region.  Such control can be quite sophisticated.  In cold weather workers ball around brood and the queen and vibrate their wing muscles to warm the core of the hive.  In hot weather they not only have teams of bees that stand in the entrance and fan fresh air into the hive, but water carriers will collect water and evaporate cooling drops of water in the current of air. 
Bees are less assiduous constructors than termites, but mainly from plants, they do collect gummy, resinous materials.  These they knead into “propolis”, a sort of all-purpose sealant and adhesive in adjusting the hive’s structure and layout.  Cerumen is a similar material with added wax. 
Using the propolis, they are likely to narrow the entrance to the hive to a comfortable size, and some, especially species of stingless bees, may form it into a rainproof, wasp-proof, intruder-proof spout as a nest entrance. 
The combs themselves, marvellous as their construction is, seem to be rather featureless, rather an exercise in industrial uniformity than anything else, but this is misleading.  At the edge of the combs one finds uncompleted cells, and they show irregularities in depth and thickness of their walls.  To complete the construction takes a great deal of remodelling, as described beautifully by Darwin a century and a half ago.  Larger cells also are built to accommodate drones, but they are sufficiently close to the normal size of comb cells, that it is easy for the inexperienced eye to overlook them.  Queen cells on the other hand, are much larger and of a totally different shape.  As usual, the variety and complexity within the hive’s repertoire is astonishing, even when one examines something that looks simple or uniform. 
Furthermore, the combs of honeybees usually are not hung randomly in the hive, but parallel to each other.  When this fails, if combs meet at an unsuitable angle, a great deal of reconstruction or reorganisation results.  It is another example of where the logic of thresholds and majorities settles things democratically.  Bees will take wax from the competing comb, and use it in their own constructions.  This usually leads to a lot of wasted and repeated labour before calm returns. 
In a sufficiently prosperous hive, the bees also will construct combs in every gap large enough to take a comb plus workers walking on it.  This complicates the construction of artificial hives.  They may look like simple wooden frames in a wooden box, but in fact they are very precisely constructed with spacers and measurements that will suit the bees’ comb-building tastes, without interfering with the apiarists’ harvesting and husbandry.  If the spaces are too wide, the bees build combs in positions that cause jamming and waste; if too small, they refuse to build and may abscond in search of a more suitable hive. 
When one goes further and looks not only at the combs, but also their contents, more complexity emerges.  They are not only used for brood cells, in which the larvae and pupae develop, but also for storage of honey and for pollen.  Typically the brood is in the middle of the comb, the pollen is stored around the brood region, and the honey around the edge.  This means that if the bees have to warm up the hive, they can do it round the brood in the middle.  Around the brood the pollen will come to no harm, while the honey round the outside is least prone to become too warm and runny. 
Space and time prevent anything like an coherent comparison of the various kinds of combs, ranging from the single paper combs of wasps, to the waxen combs of bees.  A point of interest is that though the honeybees have the most precise constructions, and though they suffice for all their needs, the various kinds of stingless bees make totally different separate honey pots in which they store their food. 
Maintenance and management of the colony comprises whole ranges of information processing controls and functions.  There is the question of the homeostasis of the environment necessary for the colony members and their functions.  There is the homeostasis of the nest materials and defences.  There is management of the costs of maintenance, of the efficiency of foraging and reliability of defence. 
The nest is not the only aspect of colony homeostasis.  There also is the matter of population homeostasis.  A colony with the wrong constitution of castes will be inefficient, insecure, or ineffective, perhaps all three at once.  A colony that forages too little in times of shortages will starve.  If it forages too much in times of plenty, the food will go to waste. 
In honeybees for instance, the hive workers begin to refuse to accept new nectar once the storage space gets too full.  For one thing, if they cannot find space to store the content of their crops, they simply do not have room to take on more; their crops become de facto storage depots, much like the repletes of honeypot ants, though on a smaller scale.  This is one of the controls on foraging.  Such control may redirect foragers to find water or propolis or return to other hive duties. 
Meanwhile the bees with the full crops are likely to metabolise the nectar they are carrying and convert it to wax.  The wax goes into building new combs if there is room, which in turn makes more room for what the foragers go out and collect.  After all, depending on the quality of the honey, it might take over ten parts of honey to supply material for one part of wax. 
This is a neat feedback system, but there is another dimension.  When the nest gets too full for new combs, that is one of the stimuli that sends scouts out to look for new homes for swarms. 
The ranges of community intelligence required for all these functions, and for invoking the functions in response to the appropriate circumstances, is not only imposing, but repeatedly astonishing.  They not only are astonishing in themselves, but in the apparently mindless ways in which such complex tasks and programs can be implemented.   


Conflicts of interests

The sum of the intelligence on the planet is a constant;
the population is growing.
A major, even a limiting, factor in the development of sociality as an ESS (evolutionarily stable strategy) is the conflict of interests that arises as soon as there are distinguishable participants.  Distinction in this matter might most obviously refer to genetic or psychological differences.  The latter are mainly obvious in large brained creatures such as mammals, but subjectively it is not easy to distinguish between conflicts of personal interest in queen bees, in fighter fish, and in mammalian sibling rivalry. 
Throughout evolutionary history genetic differences have been the most fundamentally important, though genetic rivalries often manifest themselves as personal conflict.  Investigations in the last century or so, and especially in the last few decades, have presented arguments for pervasive competition within communities, within families, between reproductives and their offspring, even within bodies. 
Most prominently, within Hymenopteran communities such as hives there is evidence for systematic competition between queens and workers, and between workers and workers.  The argument hinges on the differences in genetic relatedness between hive members.  These arise out of the fact that in the relevant species Hymenopteran males are haploid and the females diploid.  However, such fields are hazardous if one takes them simplistically at face value.  For example, the view of worker bees as competing with their queens for reproduction because of being less closely related to her than to each other or their individual offspring, is open to question even in the face of numeric predictions. 
The most serious problem is that no numerical argument has much cogency if it cannot be shown to have appropriate isomorphism to the system it describes or models.  What is the difference between the importance of genetic degrees of relationship within a colony, and the degrees of importance of the relationship between the cells in one’s various body parts?  Unless the worker favours the generation of reproductives more closely related to herself than to the offspring of the queen or those of other workers, there is no evolutionary advantage to frustrating the queen in any way.  There remains room for further analysis. 
In say, citrus and dandelion species, where, rather than zygotes arising from mating, maternal tissue most often produces cells that give rise to the next generation, one might see no conflict of interest between parent and young, except that the parent might “prefer” to produce flexible generations that are not parental clones, rather than only dead-end parthenogenic offspring.  Some animals, such as many species of aphids, do reproduce both parthenogenically and sexually in due season, and in fact do very well.  Citrus produces largely parthenogenic seeds from maternal tissue, plus a modest proportion of sexually produced seeds.  It is semantically hazardous to interpret such examples simplistically in terms of struggles between genetic lines.   
In humans conflict of interest arguably is more elaborate than in any other species; there is conflict of interest between cells and organism, between individual and individual, between family members and families, between factions within communities, between mind and genome, between mind and body, between individual and nation, between individual and ideal, or even between individual and species.  Arguably some of these conflicts entail major evolutionary penalties, such as when a self-indulgent person of high intellect or other desirable attributes chooses not to reproduce or becomes a parasitic despot. 
A great deal of argument and research have been expended on demonstrating the evolutionary justification, or advantages, of particular behaviour patterns of non-reproductives.  Although such arguments look very attractive in considering potential or prospective reproductives, such as subordinate females in naked mole rat or Polistes wasp colonies, they lose a great deal of persuasiveness in application to sterile worker or soldier castes.  Such non-reproductives have little option for selection within the colony, and their best strategy, insofar as the concept applies to their role, is to support the reproductive success of the colony. 
Conversely, it is much easier to make sense of the situation from the point of view of the evolutionary strategies of a queen or a dominant female.  Even if all the offspring share all her genes (in the ordinary sense, a most unusual situation) it might be more profitable reproductively, for a queen to dedicate some of her offspring to assisting in the production of larger numbers of more reproductively viable offspring, rather than futilely maximising the number of zygotes produced in the short term.  There are whole ranges of circumstances in which this is far more evolutionarily competitive.  In this respect the strategy of a queen is closely analogous to that of the zygote of a metazoan animal.  Naively it might seem to make more sense to feed and make more daughter cells to reproduce in their turn as quickly as possible, but in reality this is neither practical nor effective for competing with unicellular organisms.  Instead it pays to establish a smaller number of multicellular organisms that can compete on a more favourable footing. 
What happens in practice is that some of the daughter cells become somatic cells and never reproduce.  Suppose that the need is for one cell to split mitotically, such that one daughter cell becomes a reproductive stem cell and the other a somatic cell.  Which of those two daughter cells should choose to be reproductive?  The question is hardly meaningful.  The cells are not genetically distinguishable and if both became either somatic or reproductive then both would die without offspring.  Consider for example whether a muscle cell should try to favour its own life or that of its daughter cells over the reproduction of other bodily cells. 
Now, that muscle cell had had no say in becoming muscle instead of gonad.  Having been born in that situation, it is meaningless to speak of its options to favour its own type, and it would remain meaningless if that cell were part of a genetic chimera, which is what many of our bodies turn out to be in practice.  A surprising number of us are in fact mosaics of the cells of more than one zygote.  Would it make sense for those of our cells whose genotypes are not represented in our gonads, to turn cancerous so that they could produce more of their own type?  
Similarly, sterile non-reproductives in a colony may be seen as somatic cells rather than reproductive.  They can reproduce more of their genes by assisting in the rearing of related offspring and feeding and protecting related reproductives, than by favouring themselves.  In fact, if it were possible (which it seldom is) for a lost non-reproductive to join and assist a neighbouring colony of the same, or even of a similar, species, that would do more for propagating its genes than just dying.  Related species share by far the most of their genes. 
Such lines of thought lead to tricky problems when one considers behaviour such as slavery (“dulosis”) where ant species capture pupae of closely related species.  Workers that emerge from uneaten pupae then join the colony as fully conforming members of the infrastructure.  In extreme cases, as already mentioned, they make up effectively the entire worker caste.  What evolutionary pressures apply to such slaves?  


Behavioural pathology in communities

Turn on, tune in and drop out.
         Timothy Leary
This subject is far too large for serious attention here but it is important enough to demand cursory attention. 
Communities as entities, as well as their members as individuals, are subject to immense ranges of pathological behaviour, some of it interestingly reminiscent of the behaviour of creatures with integrated brains.  Drug addiction is one example, most notoriously in human communities and in ant colonies that have been invaded by various parasitic inquilines.  For example, larvae of  some Lycaenid butterflies (“blues”) enter ant nests and secrete narcotic substances that the ants eagerly consume.  It paralyses them, and the butterfly larvae then eat either ants or their larvae, depending on the species in question.  From the caterpillar’s point of view the animal food is more concentrated nutrition than plant material, and many Lycaenidae have largely become dependent on it at certain stages of their life cycle at least.  Some ants actually feed the larvae to the caterpillar.  This suggests that the caterpillar gives a particular pheromonal signal that moves the ants to do so.  As a speculation I suggest that it might be a signal that normally stimulates the ants to move the larvae about the nest. 
Just as in human communities, a heavy infestation of narcotic suppliers can have a harmful effect on the community. 
Actual poisons also may case behavioural pathology.  Apart from simple toxicity, in particular neurotoxicity, mercuric chloride poisoning provokes at least some species of ants, such as the Argentine ant, to begin fighting their nest mates.  This is interesting because Argentine ants do not normally fight other colonies of the same species, let alone nest mates.  Patently one pathological effect of the neurotoxin is on the social signalling and feedback mechanisms of the ants. 
The most pernicious and varied of such aberrations, with or without the mediation of drugs, occur most obviously in human society.  Examples include faction fights, riots, and sport hooliganism. 
There is not much evidence of play behaviour in invertebrates, not even in bees, whose workers go through various roles during their lives, and learn part of their skills on the job.  Quite apart from play in general, I can think of no example of anything resembling sport in invertebrates.  Bees and ants do have factional disagreements of a sort, where there are stimuli to change nests or split into new colonies, but these are generally the natural mechanism leading to what amounts to decisions on such matters. Such disagreements are more like eusocial democratic disagreements in the colonies than like social pathology.  The reason that group disagreements often are pernicious in human society is that human society and communication are so much more complex and less specific than in any other species.  Such complexity gives more scope for the development of pathology.   
Stress certainly can affect the behaviour and the health of a colony.  A colony that lives in a disturbance-free environment is likely to be less easily aroused and less aggressive than after regular disturbance.  Part of the reason is perhaps because of the persistence of traces of alarm pheromones after disturbances.  Part might be because of accommodation of the respective nervous and hormonal systems of the colony members to constant arousal.  In this respect colonies behave in ways that closely resemble the behaviour of many individual animals. 
Interestingly, the reactions of many colonies, as opposed to their individual members, to some kinds of disease or parasitism also resemble those of individual animals.  In beehives for instance, some kinds of infections cause the bees to clump and increase their temperature by vibrating their flight muscles.  The effect is much like a fever.  Fevers are surprisingly ubiquitous as a reaction to disease in the animal kingdom.  

Intelligence of and within colonies

For the same thing that might, perhaps, with some reason, seem
very imperfect if quite alone, may be very perfect in its nature
if it is looked upon as part of the whole universe.

It is by no means clear in what ways the intelligence of a community is commensurable with that of its members, or even in what ways the two are the same type of thing.  Apart from communities of cells that form complex organisms with the most complex of central nervous systems, we have no clearly recognisable, let alone unambiguous, examples of consciousness, let alone self awareness in colonies. 
And yet the nature of collective behaviour seems almost inseparable from collective character.  Every species of tree has its characteristic shape, often ineffable, but positively shouting its nature at those with eyes and experience to see.  A naturalist at home in his territory often can tell the species of a tree where he can make out no more specific detail than its motion in the wind.  He might see a flying bird at distances where he cannot make out anything at all except the motion and rhythm of flight, the area and direction of its course, and yet recognise not only the species, but its probable activity as well. 
As I shall point out again, these characteristics are not random.  Their causes may not be traceable in detail,  but like an individual’s voice, gait, and handwriting, they are real and they are the consequences of informational states in the colonies of cells constituting them.
The intelligence of individual colony members obviously forms the basis of some differences between the behaviour of communities of insects and of birds, jail or otherwise, but it does not do to be too dismissive of colonial intelligence.  As far as we can tell, after all, the intelligence of the individual birds stems from the behaviour of the communities of neurons in their brains.  Those individual neurons in turn are less impressively intelligent than individual bees or ants.  
There are may ways in which it is very difficult to say how to compare individuals with the communities in which they live, or with which they deal.  For instance, one normally expects the individual to adjust a great deal more quickly to intellectual pressures, while communities often have to wait for the powers that be to be replaced before there is any effective change of mindset.  And yet, one must be cautious in condemning the relative hidebound stupidity of the group.  For one thing, there is a certain robustness of a culture that can call on the greater redundancy of multiple members to substitute for each other. 
Then also, temporal factors are not always a matter of “fast mind good, slow mind bad.”  Many activities of long-lived communities or species are hardly perceptible to the faster, but not necessarily more complex, minds of their members or observers. 
It is easy to think of examples.  What good would quick thinking do a tree?  It is more viable to integrate one’s views to react to the cumulative trends of environments on the scale of the speed of change of those environments, than to commit quickly to reactions to temporary, superficial observations. 
Modern human politics is rife with examples where individuals lead their communities to exploit the weaknesses of other communities, then fail to recognise when the victim communities, typically on a time scale of decades, adjust so effectively to the challenges as to turn the tables.  It is an open question how far these contrasts reflect different complexities, different speeds, or even different types of intelligence. 


Intelligence of individuals or
castes within colonies

The intelligence of any discussion diminishes with the
square of the number of participants.
               Adam Walinsky
Since intelligence is a multidimensional variable and we have no precise and unambiguous measure for most of those dimensions, interspecific comparisons of intelligence are suspect.  Often they apply to such narrow aspects of behaviour as to be practically useless for comparisons of general intelligence.  Also, there has been little work on the intelligence of individual members of eusocial colonies of any species, and less on comparisons between such individuals and the individuals of related species that are not eusocial.  It is in any case difficult to investigate the intelligence of insects meaningfully, and the very concept of the relative intelligence of eusocial individuals is extremely tricky.  
It is open to speculation whether there are at least two classes of intelligence in colonies. One type might be where the colony has a level of intelligence of a higher order than any of its members, such as possibly a termite colony, or the intelligence of a human body relative to its component cells.  Another type could be where the intelligence of at least some individuals within the colony is greater than that of the colony. 
Whether a third type would be where the intelligence of the most intelligent member determines the intelligence of the colony, I cannot guess.  Comparisons of different types of intelligence is a very moot problem. 
Though I do not go into an analysis of the point (I do not know how to!) there are some deep problems to distinction between communities with intelligence based primarily on the structure of the community, and those with intelligence based on the intelligence of sub committees or of individual members.  As humans we might well be biased in favour of the latter, but then human communities should be far and away more intelligent than those of other species.  In practice human communities do not put up a very impressive showing in comparison to other communities.  In particular, the superiority in intelligence of human communities are nothing like as radical as the comparison between the intelligence of individual members would lead one to expect. 
A jaundiced view might suggest that the intelligence of human communities rather reflects the nature of the most selfish, obstructive, cross-grained, dominating, parasitic, vindictive and malicious of their members, rather than the most intelligent. 
This is not necessarily self-indulgent, simplistic misanthropy.  Humanity is adapted rather to life in small groups where everyone knows everyone else.  Instead we live in faceless masses in which social parasites can prey on socially supportive members. 
Structures of unselfish, mutually beneficial and supportive social interactions are metastable in most circumstances because they are vulnerable to parasitism.  Among humans it seems to be easier for betrayal to change them into distrustful, resentful, selfish relationships than the other way round.  In human families and intimate communities this can lead to intense embitterment.  In larger, anonymous, urban communities the personal embitterment may be less intense, but alienation, distrust and selfishness may become a way of life if the powers that be are not sufficiently firm and far sighted. 
These are not problems that arise in most social animal communities.  In eusocial communities the necessary structures just do not exist.  Not that one gets no strife in such communities, even deadly strife, such as queens destroying queen cells or younger queens, but such things are totally impersonal competition.  In a healthy hive they do no lasting harm.  There is no indignation or obvious resentment. 
In contrast, in many species of monkeys, biting is the standard form of chastisement, particularly of subordinate by dominant troop members.  However, in many such species the canines are large, dangerous weapons, and disciplinary nips involve only the incisors and are not much resented.  Biting with the canines on the other hand, invites instant retaliation, much as biting or kicking might do on a first world school playground where there are conventions for the use of fists above the belt only. 
Such conventions for the control of social interactions in communities are necessary in species in which personal recognition is essential.  In eusocial species this is not a consideration except for such things as recognition of the fact that individuals share the same hive smell, or possibly that worker bees share the same floral smell, or things of that type.  In human communities there are sub-communities such as families, gangs, neighbourhoods, and the like.  These may overlap or even demand conflicting or contingent loyalties. 
Loyalties and inter-group enmities are among the most important and basic controls of mammalian or avian social interactions, and in humans they are the most abstractly and variously developed.  Without following this line of thought too far, note the way that neighbouring gangs, schools, or sporting teams and their supporters will treat each other with pointless, automatic resentment rather than amiable rivalry, but when they are pooled for regional competition, the supporters unite almost seamlessly and almost unconsciously. 
Such hierarchical resentment and loyalty may be traced on all scales up to the international.  It is widely, almost automatically encouraged in education, but repeated experimental investigation has demonstrated that it also is innate and arises in totally meaningless and artificial associations even between strangers.  It makes patriotism one of the most powerful and resources of the cynical demagogue, not, as Johnson said, the “last resort of a scoundrel”, but, as Bierce observed, “the first”. 
Simply put, the population is a fertile medium for such exploitation. 
I do not intend this as an irrelevant Jeremiad or satire, but as an illustration of a mechanism of control and characterisation of community interactive control.  It depends on innate attributes of the community members, is rarely recognised consciously as such by participants, has little to do with any participant’s conscious intelligence, and yet controls the bulk of the population without appeal, often at the whim of ruling sub-populations, cliques with their own internal loyalties.  It has been like this for all human history.  “All were for the party.  None were for the state.” 
Furthermore, the smaller the human community, the more directly the intelligence of the leaders is likely to be reflected in the policy of the tribe.  In large nations one is likely to encounter the curse of the committee, encapsulated in many a quip about the standard of the intelligence of any community varying with some inverse function of the number of members. 
Quips are all very well, but there are material reasons for taking such relationships seriously, and some of the reasons favour neural networks in modular brains on the one hand, and on the other, eusocial insect societies.  They may not be as intelligent in some senses, but they also interfere less with each other’s intelligence.  Each member of such a community simply reacts to neighbours’ behaviour for local decisions, and to major signals such as sounds or pheromonal or hormonal diffusion for long distance signalling.  The signal may be unable to carry many bits of information and accordingly it may be simplistic, but that need not be a problem if the repertoire of reactions is small and suited to the community’s means of dealing with its environment.  When that is the case, then the effect is for the community to display startling sophistication in its reactions as an entity. 
When human communities try to give everyone a voice and to consider each voice before taking action, then everyone (in theory at any rate) should hear every opinion, weigh it and combine it with every other opinion, and finally produce a digested, hopefully optimal, conclusion as a basis for the policy to be implemented.  In practice the problems of choosing the right leader in each decision, the quadratic growth of two-way communication, and exponential growth in the number of sub-committees, the increased path length of lines of communication as the community size grows, the intellectual limitations of certain members, the disagreements among others, the conflicts of interest, lead to appalling results. 
The more passionate the sincerity of the reformer the worse the results one may expect.  As John Gall said in his brilliant book “Systemantics”: “Reformers blame it all on  "the system"  and propose  new systems that would, they assert, guarantee a brave new world of justice, peace and abundance.  Everyone, it seems, has his own idea of what the problem is and how it can be corrected.  But all agree on one point  -- that their own system would work very well if only it were universally adopted.” 
In practice no system of government follows such extreme logic.  For one thing our species is not as well adapted to such innate controls as the eusocial colonies are.  Pretty soon, no matter what bliss it might have been in such a dawn to be alive, it becomes necessary for the leaders to reduce their discussions to cabinet level, and the inevitable slide into indirect, party democracy or even despotism, begins. 
In short, there is not yet much foreseeable hope for communities of intelligent individuals being unambiguously more intelligent than the communities of unintelligent, or at least less intelligent, individuals.  The participation of too many independent brains exacts crippling penalties.  Their mutual interference might not affect their respective effectiveness, but often it certainly affects their communal effectiveness. 
Still, the intelligence of community members does have some effects.  Social structures based on more stereotyped, genetically determined behaviour are more subject to direct evolutionary selection, because counter-adaptive behaviour carries a more direct penalty for the genes responsible. 
In contrast communities of organisms with more flexible behaviour might be dysfunctional for contingent reasons that are not a direct function of a particular heritable trait.  Although they could not escape the consequences of severe dysfunctionality, they would not be not subject to the same precision of adaptive selection, but the consequences of the looser selection should include a wider range of social behaviour patterns. 
Such variability would increase as a function of the intelligence of the individual members of the communities.  There is some support for this among primates, but far and away the most spectacular variability is in human communities. 
Apart from the variability of human communities, there also is greater complexity of community structures and activities.  Some of those forms of interactions could be seen as components of community intelligence that are as different from the community intelligence of eusocial insects, or naked mole rats, as the brain of a primate differs from that of a fish.  The major part of the primate brain consists of extensions to the brain stem that simply do not occur in fish or amphibians.  Analogously, eusocial communities have nothing that corresponds to business, religion, art, science or many other activities that deeply influence human societies.  It is hard to see whither such developments will take human societies, but they are materially different from any components of community intelligence in extant eusocial species.  
There have been some speculations to the effect that eusocial individuals would be unusually intelligent in specialised activities appropriate to their castes, and unusually unintelligent in other respects, as compared either to other castes or to non-specialised individuals.  Sometimes this seems to be so, and certainly the brains of some eusocial insects have local enlargements that differ both from the brains of nest-mates of other castes, and from the brains of related non-eusocial species. 
This is very suggestive of course, but it is not much more than suggestive.  The ratio of brain size to body size is a useful rule of thumb for estimating intelligence in mammals, and it surely is not without significance in other animals, but it also is highly imprecise and untrustworthy as a measure of intelligence.  And in any case it is not based on any clear functional relationship. 
It is not at all clear that the size of the brains of say, crows and parrots, gives a fair indication of their intelligence compared to that of mammals of similar brain to body size ratios.  It is quite possible that different neural organisations might give greater compactness of brains with competitive levels intelligence.  Pigeons, not normally regarded as mental giants, have demonstrated behaviour that suggested startling intelligence by mammalian standards.  (Of course, we must be cautious.  It is not many decades since the high intelligence of pigs was formally recognised, and anecdotal evidence suggests that black rhinos are disconcertingly intelligent.) 
At the same time, absolute brain size remains inescapably important.  Obviously no single neuron could have a very high IQ, but less trivially, a little girl with a particularly vicious case of persistent epilepsy had “half her brain” removed (presumably a defective cerebral hemisphere).  She was very young and everyone was delighted with the speed with which her half brain adapted and took over the running of both sides of her body, leaving her superficially normal in most obvious ways.  Subsequently however, it seemed that there were in fact limitations to her learning ability, as though her brain just were not large enough.  Anyway, it cured the epilepsy. 
This case is of course very far from cogent, being abnormal, probably pathological, without controls, and open to rival interpretations.  However, it is sufficiently suggestive to justify suspicion of clichés such as that humans use just X% of their brains, where X an arbitrary small number, varying according to the source of the assertion.  Such clichés too, are very far from cogent, being abnormal, probably pathological, without controls, and open to rival interpretations. 
Anecdotal evidence of purposive behaviour suggests that some very small-brained animals are disproportionately intelligent in particular ways related to their adaptations to their ways of life.  For instance, I have personally observed that Cape chameleons (Bradypodion sp.) can rapidly trace an unfamiliar, unobvious and circuitous path to a perceived rival several metres away, and follow the route directly without preliminary trial-and-error exploration.  This is far beyond what say, a dog could do.  It is an innate skill that might reasonably be associated with short range navigational problems commonplace in shrubbery. 
Taking this even further, I have been astonished to see a noctuid moth (Sphingomorpha sp.)  fly through an open window into a lighted room at night, and when it was alarmed while still in flight, turn without alighting, and unerringly navigate its way out again by the same window a few metres away.  It was one window of several.  This was astonishing behaviour in an alarmed insect at night; as a rule one would expect automatic flight towards or around the lamp.  Even diurnal birds alarmed at night will usually fly blindly towards a lighted sheet.  What passed for mental processes in the moth, I cannot say, but the subjective effect was that it gave an impression of surprising purposive intelligence. 
Interestingly, in the laboratory at least some species of eusocial insects, particularly bees and ants, show intelligence thoroughly comparable with other insects.  This recalls Lorenz’s opinion that social animals are more intelligent than solitary species because they need the extra facilities for social interaction.  He mentions foxes as being less intelligent than wolves for example.  There is room for argument on either side of this point.  I mention it just for consideration in context. 
It also is worth noting that the brains of many species of worker ant are exceptionally large relative to the animals’ size.  I have already mentioned some reservations on the significance of this ratio, but it is interesting that their maze running behaviour is only a few times less than that of a rat.  Significantly however, unlike a rat, they do not obviously benefit from having previously learnt the maze when they have to learn to run it in reverse. 
Then again, both bees and many ants are good at navigating their way back after a convoluted outward foraging sortie.  What is more, species that do not follow trails routinely, tend to follow a direct route back to the nest, often amazingly accurately.  Even the trail-bound species tend to shorten a long-used trail progressively by cutting corners. 
Many insects show elementary learning abilities when confronted with the standard laboratory challenges, maze learning, Pavlovian responses, and the like.  However, insects are usually not able to generalise from learnt experiences, much less show any comprehension of the tasks they perform.  Still, occasionally there is an innate ability to perform surprisingly sophisticated tasks when the species is adapted to particular demands. 
For instance, bees, with their need to find flowers, and, having found them, locate or evaluate the nectar and pollen within them, are surprisingly good at pattern recognition and memory of patterns once they have found food associated with them.  They also can to some degree recognise categories of patterns, such as the same pattern in different colours.  Another very impressive feat is the worker bee’s ability to find the direct way back to the hive when having found food after a convoluted search. 
Especially in more stereotyped, but apparently intelligent behaviour, one is at a loss whether to ascribe intelligence to the species, the colony or the individual.  Consider the ability of some species of ants and bees to navigate by the sun.  Is this their own intelligence, or that of the colony?  In the latter case we should regard the “intelligence” of the individual ant as being analogous to the intelligence of an optic neuron in the human eye, specialised to identify a vertical or horizontal edge.  The ability as such is trivial, but it gives great power to the intelligence of the colony. 
The local enlargement of regions of the brains of various castes of insects is of course highly suggestive.  To the extent and in the ways that intelligence is a function of the amount of brain tissue available to support it, that phenomenon makes it extremely likely that such insects differ in the degrees and types of intelligence they exercise. 
There is more to it than that.  Not only are all the specialisations within a given species the product of effectively the same genome, but in some species, such as social bees, the same worker undertakes different duties at different stages of its life, in a more or less fixed sequence.  In the case of the honeybee, which probably exhibits this effect in its most advanced form, a typical sequence would be something like: nursing older larvae, nursing younger larvae, receiving and processing nectar from foragers, housekeeping and cleaning functions, comb maintenance, comb building, water carrying and foraging. 
Such functions vary with the demand in the hive, and often workers differ both in the duration of each duty in the sequence and in the degree to which they deviate from it.  Patient observation has shown that many bees and other social insects seem to spend a lot of time apparently loafing.  This should not be taken too seriously, on the principle that they also serve who only stand and wait, and idle bees consume little food.  Furthermore, during the hive phase of the life of a mature bee, once it is say, a week or two old, it “patrols” the hive, looking for odd jobs.  It deals with each job as it encounters the need for it. 
Even some of the jobs that sound very organised, are tackled in a very arbitrary fashion.  For example, comb building, that most precise of the engineering tasks that bees undertake, begins by the deposition of tiny pellets of wax where some of the bees happen to leave them.  Other bees might pick up the wax or add to it according to individual reaction.  As work progresses, more and more bees join in, adding, removing, and sculpting wax.  It begins crude and thick, and ends up exquisite and precise. 
This illustrates principles that are very important in many ways.  The fact that a large part of the colony’s survival and efficiency depends on such ad hoc recognition by individual workers, of when there is work to hand, shows how complex a society can be maintained by the apparent whim of individuals.  A great deal of the activity of humans in a free enterprise system is strongly reminiscent of that mechanism. 
It would be interesting to see whether it would be possible to have a lot of human workers build a structure or work of art out of say, clay or wax, or computer graphics, volunteers free to fiddle with the material as they please, perhaps for a maximum of half an hour each, no talking allowed.  It is commonplace that a singing crowd, as opposed to a choir, hits the correct notes with remarkable precision. 
Be all that as it may, some of those bees’ duties, such as the feeding of the young and the production of wax for the building of combs, depend on stages of physiological developments.  The main examples are the development of the glands that produce the bee milk that forms part of the food of the larvae,  and of the wax glands that produce the comb wax.  These strongly influence the degree to which the bees can alter the sequence of their functions.  Conversely, some individual worker bees specialise in particular duties more than some of their nest mates.
None the less, not only is the sequence of the employment of a worker bee fairly well marked, but some of the transitions have a tempting logic.  It is for instance after a stint of feeding the older larvae that the bees begin to feed the newly hatched larvae.  It is after comb maintenance experience that they join clusters for comb building.  It is reasonable to suspect that the comb cleaning and smoothing duties act as development initiators for innate comb building skills. 
Such ranges of behavioural development within individuals suggest intelligence and mental development of various kinds.  However, the behaviour, though flexible in duration and in response to need and circumstances, is largely stereotyped.  Consider pollen collection.  This is a vital activity because pollen is the main source of most body-building nutrients other than the carbohydrates in nectar. 
Collection of pollen depends on availability, on need, and it is a complex activity that varies with the plant that supplies it.  Different plants signal the availability of pollen and nectar in different ways, some of them by changes of colour and scent.  Some do the pollinator violence, for instance the stamens of some plants swipe the bee to daub it with pollen.  Bees visibly behave as though the experience were negatively reinforcing, and have been seen apparently trying to avoid the blow after they have had sufficient experience to teach them what is coming.  Furthermore, some kinds of pollen are not of value to the bee and are not actively collected. 
What is more, flower species differ in the nature of the duty they require.  Some, such as Eucalyptus macrocarpa, offer more nectar than any one bee can carry, for pollinating just one flower.  Strictly speaking, Eucalypts with such large nectar supplies are not bee plants, but adapted to pollination by vertebrates.  This does not discourage the bees though; they avidly pollinate many species of Eucalyptus and other Myrtaceae, including those with the largest burdens of nectar.  It does not follow that this is bad for the plant; such a flower is likely to be visited by several bees, and receive pollen from many other plants. 
It shows that the simplistic view of plants offering the least reward possible to make it worth pollinating them, is too naïve a view of the selection pressures on the species in the relationship.  If there is some form of intelligence in such relationships, it is obviously not trivial to characterise it in any predictive form, however facilely one might spin Just-So stories after the fact. 
Other major honey plants, such as alfalfa, give minuscule amounts of nectar from each of thousands of florets.  For each form of flower the bee has a particular strategy to exploit the flowers, for example working its way up one column of flowers on a raceme, and flying off to another raceme when reaching the top.  This may sound counterintuitive, but some research suggests that for some flowers it actually may be more energy efficient.  However debatable this point, the fact remains that there are many more possible inefficient than efficient algorithms for pollinating a complex inflorescence, and the bees use at least reasonably efficient algorithms.  
The relationship between flowers and specialist pollinators gets even more complex in some species, with for example, the flower changing in colour and in scent once the nectar has been removed and the stigma presumably pollinated.  It might then produce pollen, and get visited mainly by pollen collectors.  Later on it is no longer in the interests of either party that bees should disturb the flower, and the appearance changes again.  Foragers from eusocial bee colonies not only specialise in the type of collection they perform during a given few days, but in the time of day.  They visit the same spot and seek the same scent, and collecting the same type of food until the supply dries up.  During the rest of the day they are very likely to remain in the hive while other bees undertake other duties.  Interestingly, within the hive they often form groups, cliques if you like, of similarly scented bees that have been visiting the same patch or type of flower. 
Solitary bees often specialise in the species they visit, but cannot afford the luxury of moping around the nest between chores. 
Also, some species of flowers, including rich sources of nectar such as Aloes, might keep their nectar in tubes that the bees cannot reach into.  They normally might be bird or moth pollinated.  Often the bees will discover the fact and learn to chew through the side of the tube and rob the flower without pollinating it.  There is dispute about whether learning such tricks from either nest mates or from other species happens. 
Then again there is the point that bees soon learn to visit particular flowering plants How much of this behaviour is stereotyped,  and to what degree, is unclear, but it certainly is intelligent by some criteria. 
A vivid example of where behaviour adjusts homeostatically to match circumstances, is the way in which bees will ventilate their hive when hot, and bring water which they fan for evaporative cooling.  This is reminiscent in the way that a very hot cat will slather itself with saliva, cats having hardly any ability to sweat.  Such behaviour seems to be equally innate and stereotyped in both cases.  
There are other similarly illustrative analogies.  In cold weather bees will bundle for warmth much like emperor penguins, and they stock their combs in zones that reflect the need or tolerance of the contents for warmth.  As for how much more intelligently vertebrates behave, any poultry farmer will know how easily chickens may smother each other when large flocks huddle for warmth. 
Homeostatic shivering in vertebrates has direct counterparts in moths warming up the wing muscles in their furry thoraxes, and in bees that vibrate their wing muscles to warm a cold hive or cook an enemy smothered in a bundle of workers.
Then again, some eusocial insects have very high degrees of task specialisation between castes, and hey typically have castes that, unlike honeybee workers, do not much change function during their adult lives.  This occurs in many species of ants, and probably most termites.  Among termites the soldiers usually are physically well-marked castes, such as in the snouted harvester genus Nasutitermes, the soldiers that plug nest openings with their heads, both in some Camponotine ants and in Cryptotermes, and the various kinds of large-jawed, heavy-headed soldiers in many species of both ants and termites. 
For example, apart from the reproductives, some ants have two, some more than two obviously different castes.  Such anatomical specialisation of castes goes with behavioural specialisation.  In fact, many such specialised castes are simply unable to undertake the duties of other castes.  In fact, non-worker castes often cannot even feed themselves. 
Interestingly, in many species of eusocial insects, particularly ants, such as the Argentine ant, Linepithema humile (formerly Iridomyrmex humilis), there is just one multipurpose non-reproductive caste, while in say, Pheidole megacephala there are few large-headed soldiers and they seem not to be as active in combat as the workers are.  I suspect that such soldiers are on the way out, and that given time Pheidole megacephala will become an ant with just one non-reproductive caste.  I furthermore suspect that some species of non-specialist ants already have undergone such changes and have lost castes that were present in their ancestors.  In other words, their condition of having just one non-reproductive caste would be secondary. 
There are other examples of specialisation in function within eusocial colonies, apart from definite castes.  One example is the use of immature colony members for particular duties.  For instance, larval termites (“nymphs”, though real entomologists seldom use the term nowadays) of many species will wall up small enemies or undesirable matter by regurgitating building material over them.  Ant larvae of the weaver ants are the source of the silk used in binding the leaves from which the workers construct their nests.  Furthermore, it is practically universal among eusocial species that young adult workers begin with duties inside the nests, before going out for external duties. 
Also, some species of ants, such as Anoplolepis custodiens, have no clearly defined multiple worker castes, but just a size range of workers, typically a log-skewed normal distribution that yields a startlingly straight line on being subjected to probit analysis.  Such observation strongly argues that the distribution represents a dosage response curve that reflects random feeding of the larvae, such that those most frequently fed grow into the largest workers.   Such feeding could easily result from stereotyped feeding behaviour in worker ants.  This suggests a phylogenetic point of origin for the evolutionary development of multiple castes.  If that is correct one might expect little difference between any form of intelligence, between the larger and smaller workers, because they are no more than developments of slight variations of the same processes in the same organisms.  However, the argument is not very strong; even in honeybees various parts of the workers’ brains change in size during their adult life, presumably in association with their experience or changes in their duties or hormonal development or all three. 
As we have seen, even in eusocial insects with just one worker caste of more or less uniform size, such as honeybees, it is commonplace that some workers concentrate on particular duties to a greater degree than others.  It is only to be expected that where there are differences in size and shape as well, there would be much greater differences of bias in the types of work undertaken.  In species where there are distinct castes of different sizes, the types of work undertaken are strongly dependent on physical differences between the workers. 
A spectacular example that I mentioned in another connection, is in leaf cutter ants, species of Atta.  Here we find small workers riding shotgun on leaf segments carried by the large workers.  They are not just hitchhiking, but protecting their larger sisters from parasitoid phorid flies that otherwise would lay their eggs on them.  Note that these phorids are not the same as the ones that attack fire ants, but there is an interesting thing about the strategy of the Atta colonies.  Unlike the apparently frightened behaviour that paralyses the working production of a fire ant nest, this allocation of a part of the minor worker force to protection of the convoy of large workers, permits the colony to remain productive, full speed ahead and damn the parasitoids.  As I already have remarked, the cost to the colony of a minor worker is just a few percent of that of a major worker. 
In eusocial mammals, naked mole rats, there is noticeable physical difference between soldiers and workers.  There also is considerable, though not necessarily absolute, difference between their respective habitual behaviour.  It is plausible that there is a significant difference in intelligence, but it is clear that their behaviour patterns are not quite as stereotyped as in the eusocial insect castes.  It also seems that their specialisation is more reversible than in highly eusocial insects.  If the queen dies then in a healthy colony the changes in hormonal balance cause other members of the community to re-adapt to the role of the queen.  In some eusocial insects, such as many species of termites, existing immature larvae may develop into secondary reproductives of various degrees of effectiveness. 
Now, there is relatively little published material on the subject of the brains and intelligence of eusocial insects, and what there is, is largely speculative or limited in scope.  Not surprisingly there is a great deal of stereotyped behaviour, and wherever this is demonstrated it is likely to be characterised as non-intelligent behaviour.  From our point of view in this discussion, such non-intelligence is debatable.  What is more, even stereotyped behaviour may be fairly flexible, whether in insects or in humans. 
One example is in web building spiders.  On one hand, most species will repeatedly run out to attack a wax-covered insect when lured with a suitable tuning fork.  Conversely, though web building is a largely stereotyped behaviour, not only must the web conform to local circumstances, such as the available points for tethering, and the suitability of the site for catching prey, but in species that build orb webs and repair them every night, the repair activity seems to respond to the success of capture of prey.  If one places prey consistently in the same corner of the web, then during reconstruction, the spider progressively builds that corner of the web larger and larger. 
Is this intelligence?  Learning?  It certainly is flexibility, though it is not clear whether that flexibility is a “mental process”, or the modification of one component of stereotyped behaviour by another.  In either case, why or in what way is that not learning?

Intelligence of colonies

There was a cage with several apes in it.  In the cage there was
a banana hung on a string, and stairs under it.  Before long an
ape went to the stairs to get the banana, but as soon as it even
touched the stairs, all apes were sprayed with water.  After a
while the same ape or another one made another attempt, with
the same result: all apes are sprayed.  If later another ape tries
to climb the stairs, the others will try to prevent it.

Now they took one ape from the cage and put in a new one. 
The new ape saw the banana, and wanted to climb the stairs. 
To his horror all other apes attacked him.  After another attempt
he knew: if he wanted to climb the stairs, he would be beaten up. 
Then they removed a second ape and replaced it by another new one. 
The newcomer went to the stairs and got beaten up.  The previous new
ape took part in the punishment with enthusiasm.

A third old ape was replaced by a third new one.  The new one made it to
the stairs and got beaten up as well.  Two of the apes who beat him have no
idea why they may not climb the stairs.  They replace the fourth old ape, and
the fifth, etc.  until all apes which have been sprayed with water have been replaced.  Nevertheless, no ape ever tries to climb the stairs.

   One day a new, young ape asks, "But Sir, why not?"

   "Because that's the way we do things around here, my boy."

Anonymous (traditional on the WWW)
If it is difficult to compare or even define the intelligence of individual organisms, it is far more so with communities.  As I have remarked, it is hard enough to define the intelligence of a community at all.  There are grounds for arguing that if group intelligence means anything at all, then some kinds of intelligence and temperament in the group are not the same thing as in the individual.
For example, any teacher or drill sergeant knows that each group of pupils or rookies has its own character.  One gets them good and bad, affable and sullen, biddable and unwilling, stupid and talented.  Sometimes it is easy to tell what some of the sources of some of the attributes are; here there is a class member with high skills, there we find a couple of rotten apples, again, we find a class that has not met its prerequisite training goals, or one class comes from an educationally backward area, while another comes from a university town teeming with gifted children. 
And yet, that explanation is nothing like sufficient, let alone of predictive value.  Given the most consistent possible source of members, each batch differs from the rest.  In fact some teachers will swear blind that the years of good classes alternate with years of poor classes.  Similar differences appear with sporting teams, teams of workers, company staff, public servants, teams of dogs, and almost any type of community one might mention. 
Sometimes one can change the nature of such a community by replacing or removing a few members.  Sometimes, perhaps more often, a particular attitude, atmosphere, character, remains associated with the team indefinitely.  Often unthinking tradition will determine the strengths or the ultimately fatal weaknesses in an erstwhile dominant company until it goes under and its staff disperse into other institutions. 
Certain approaches that have been of limited value in exploring the human mind can be of even more limited use in studying eusocial communities;  The main approaches have been to examine the behaviour of the colony and the contribution of certain members of the colony, and to observe the effects of removing or interfering with the activity of certain members of the colony. 
So for example, we can see that say, comb-building and foraging in bees are worker activities.  We find it unsurprising that the worker brains and anatomy are the only ones in the hive that equip them for such activities.  In much the same way, we do not expect mature human retinal neurons to be interchangeable with say, neurosecretory cells in the hypothalamus, any more than worker castes are interchangeable with queens.  We also find that typically, soldier ants and certainly mature soldier termites, contribute nothing to the tasks of workers. 
Although objectors might point out that in fact neurons do have considerable functional plasticity, that is nothing like enough to suggest that mature neurons are anything like interchangeable.  And even if the occasional worker does lay an egg, or the queen does display a bit of incidental worker-like behaviour, it does not imply that their roles are not distinct. 
Conversely, by removing a source of particular pheromonal or behavioural control from the colony, we can tell how rigidly the behaviour is limited to a given population in the colony.  In some kinds of termites, losing the queen causes suitable larvae to become fertile, though they seldom can rival the function of the original queen.  Vigorous colonies of naked mole rats or of some kinds of paper wasps can replace the queen in a fairly short time. 
However, the fact that some members of the colony have particular functions and are specialised for those functions, does not mean that they understand their own functions, any more than a neuron in the human visual cortex knows what a face is, even though its function is vital to face recognition.  Most often any particular member almost certainly has no explicit understanding of its own role and actions, any more than a neuron in the human cerebrum could understand its own role in art appreciation.  “All” it does is to modulate its output according to the interaction of its initial state at any given time, with the rates of pulses that it receives from it various input synapses or external stimuli. 
Like the actions of insects in eusocial communities, the neurons’ behaviour is stereotyped, though flexibly so.  And yet that very behaviour is by far the most apparent contributor to the intelligence of brains.  Rival contributors so far identified are minor in comparison, such as hormonal distribution in the body, or falling down when fainting, which maximises the body’s chances of recovery.  What is more, such mechanical controls have even less pretension to intelligence of their own than neurons do.
Repeated observations have shown that most of the activities of individual social insects are stereotyped, with very limited (though vitally important) flexibility.  In fact, even apparently purposive architectural activity such as mound building in termites or comb building in bees or wasps, patently are not understood by the individual participants.  The bees use very simple actions in combination with the geometrical effects of crowding, and the termites use feedback from often random placement of pellets of nest material. 
Granted, it is not all they do; when there is a small breach in the wall, they place their pellets just in the breach, but that activity too, is stereotyped.  The nature of the way in which they cooperate depends in some ways on the pheromones released at the site of the activity. 
Still, it is very important to note that an entomologist knowledgeable about termite nests can recognise a considerable number of species from the morphology of the nest alone, even when the insects themselves look very similar.  The most famous example is the nest of the Australian compass termite, but that is by no means the only one.  For example fungus grower nests such as those of Macrotermes may look less spectacular, but they are more complex internally.  Each species builds nests adapted to its needs, often marvellously so.  Some species build their nests by tunnelling into wood, some below the ground, some in mounds on the surface.  Most are distinguishable by texture, colour, size, shape, orientation and position.  Their designs reflect the available materials, climate, threats, substrates, and resources. 
This is a crucially important point; the nests vary as characteristically as birdsong, as handwriting, and they vary for similar types of reasons.  Though we cannot trace all the details, it is clear that the character of one’s  handwriting stems from one’s mental structure, as reflecting certain control systems in one’s brain, partly innate, partly developmental and partly modified by external stresses and stimuli.  Similarly, the control of the building activity is partly innate, and partly in the systematic informational interaction between the nervous systems of the participants.  It furthermore is modified by environmental stimuli, such as the daily passage of the sun, the depth of the water table, the presence of food, attacks from enemies, and so on. 
One way or another, all the most advanced colonial species rely on their own types of infrastructure, the nest itself, the stores, the labour and security communities, and in each case the form of the infrastructure is characteristic of the particular species as influenced by the local environment. 
It gives one pause to compare the shape of the termite nests with the nests of solitary mud wasps, or the mouth of the male mole cricket’s tunnel, shaped like an exponential horn, the better to propagate his song.  Some solitary species build their own infrastructure, as characteristic as the communal constructions of termites, and sometimes of startling sophistication.  This implies that the solitary organism interacts not only with the environment as it was, but with the products of its own earlier activity and constructions.  Here there is something vaguely reminiscent of the way that a single quantum can interfere with itself in passing through a grating, but more realistically, the creature is responding to the feedback from its own previous labours. 
Some very important work has shown how flexibility in response to feedback can be superimposed on stereotyped repertoires of activity, to achieve building activity.  For example, particular kinds of ants collect scattered refuse such as grains of sand or corpses into patterns.  Such patterns are characteristic of the species.  Similar behaviour determines the placement of nest material in the building of termite mounds. 
Some such patterns are so simple that they can be simulated graphically by fairly elementary computer algorithms, thereby revealing the simple nature of such aspects of the colony intelligence.  However, to find one simple principle is one thing, to assume that it is the whole story would be simplistic.  Note that it is far more difficult to simulate anything like the total building of the nest.  Simple as they presumably are, some aspects of such communal behaviour still have not been worked out.  
Though it is clear that the intelligence of the colony is neither the intelligence of the individual colony members, nor yet the sum of their intelligence, it still looks like a definite intelligence, even if it is rudimentary.  What to compare it with is hard to say.  How to characterise it may be harder.  The beehive might be seen as being like an octopus that locates and retrieves food with its tentacles.  The driver ant colony might seem very like a wandering predator, shedding cells (colony members) in capturing food to grow more cells, and so on.  Some colonies, particularly beehives, learn where there is food and what the food looks and smells like.  Some, like foraging ant columns, learn where food seems to be absent, by marking unproductive paths as such. 
Bee colonies may become inhibited from working when predatory wasps, such as the South African bee pirate, Palarus latifrons, are about.  The wasp lurks outside the nest and captures water carriers or foragers as they emerge to go about their duties.  This suggestively resembles the fear that say, a herbivore might display when predators are about. 
This panicky effect on bee colonies is not unique.  Phorid flies, of the genus Pseudacteon in particular, are being used as biological control agents against the various South American species of fire ants (Solenopsis) that have become pests in the southern United States.  They lay their eggs in worker ants and the larvae eat them from within.  On first thought this seems a little puzzling, because the flies are not particularly fecund and it takes the death of a lot of workers to destroy a fire ant nest.  In fact, estimates are that only a few percent of the ants get infected, far too few to harm the colony much. 
It turns out that the ants detect the presence of the flies and instantly go into defensive mode, turning over to fend them off.  The whole workforce is likely to run underground as soon as they detect the flies.  Just a few flies are enough to waste a whole workday for a colony, and that is more expensive than the loss of a few dozen workers. 
Interestingly, although the reaction of the workers is costly, its very existence, violence, and specificity suggest that the attacks by Pseudacteon have been a feature of the environment of Solenopsis for millions of years.  This also is consistent with the fact that the flies are so very selective in the species of ants that they attack. 
But then why such an expensive strategy that it paralyses the productive activity of the nest?  Atta have a similar problem and their strategy has reduced it to the level of a nuisance.  The most persuasive argument is one of evolutionary opportunism.  Solenopsis does not indulge in any activity in which it would be practical and obvious for one worker to protect another, plus, they do not have multiple worker or soldier castes to take over such duties.  Presumably those facts denied them the opportunity of developing the “shotgun riding” strategy. 
Conversely, as a species they cannot afford to ignore the flies; the current rate of loss is trivial, but if they did nothing to frustrate the flies, they would increase indefinitely and might wipe out the ants.  If this were not so, there would not have been any selective pressure for them to develop the behavioural patterns for avoiding the flies. 
To return to the honeybees, other species such as Apis cerana in Japan, will ball around certain types of enemy wasps till they kill them with heat, most famously the giant hornet, Vespa mandarinia.  This suggests a tempting analogy to metazoan fevers in reaction to certain diseases, particularly viruses such as  influenza. 
Studies on beehives have shown that there are particular stimuli, smells, movements, colours, textures and so on, that are likeliest to provoke any hive to attack.  None the less, some hives are more aggressive than others and breeders successfully select for docile colonies.  Different strains have different personalities.  South African beekeepers returning from exhibitions overseas often are dumbfounded to see the casual behaviour of European or American colleagues.  In South Africa foreign beekeepers would change their modus operandi pretty smartly, especially in dealing with Apis mellifera dorsata, which has a notoriously low threshold of intolerance. 
Another observation is that beekeepers dealing with the more irascible breeds of bees find that hives differ in their preferred target.  Some go for the eyes, others for the buttocks.  I am not aware of formal work on this matter, but certainly in South Africa this observation is a commonplace among long-suffering practitioners.  Whatever the mechanisms behind these patterns of behaviour, they are inescapably analogous to temperament in individuals in say, humans and dogs.  Here too, temperament is affected by both genetics and environment. 
What is more, it cuts no ice to claim that differences between the hives are a trivial matter, being due to the relative sensitivities of the individual members.  A perfectly valid counter observation is that slight differences in the receptor and neuronal sensitivities of persons and animals lead to drastic differences in temperament and behaviour.  So does the history of the individuals and the community; a colony that has lived undisturbed for longer than the life of any of its workers is likely to be more tolerant than one that has frequently been provoked. 
And yet, we are left with a serious question.  In analogy to an individual animal, does the colony learn to recognise particular patterns?  Actually they do.  An example is the appearance of its own nest opening.  This is so specific that if one moves a hive a few metres, the effect on the colony is disastrous.  The workers returning from foraging for food or water will go to where the nest had been the previous day.  Conversely there is no problem to moving the nest several kilometres to outside the previous foraging range of the workers, because then the workers do not recognise the new territory and the first thing they do is to familiarise themselves with the new position of the hive. 
Some tricks do help for moving the hive over short distances.  One can persuade the colony by moving it a few centimetres at a time, say less than the width of the hive per day.  This is a homing task that the bees can adjust to.  Or one can decorate the hive with large projections, or paint the front of the hive a vivid colour combination in a bold pattern that dominates the territory for a few metres around.  This trick also is particularly useful when one keeps many hives in close proximity.  It reduces the tendency for workers to enter neighbouring hives by mistake. 
Another trick is to put a sheet of clear glass just outside the hive exit each time the hive has been moved the previous night.  The emerging bees bump their heads on their way out, and that concentrates their attention marvellously.  They then fly around the hive to confirm its precise location.  A couple of days later one can repeat the trick, till the hive reaches its new site. 
These observations are in no way trivial.  All these cases present examples of learning by individuals, in such a way as to amount to learning by the colony. 
But can the bees learn recognition in the same way as the most intelligent vertebrates?  Can they learn to recognise a face?  I am not aware of any such effect in insects.  Worker bees certainly have been trained to collect nectar where there is a picture of a particular face, and no doubt some of the workers that they have recruited learn to recognise the same picture.  Personally I regard this exercise as just a demonstration of the complexity of patterns that the bees can learn to recognise.  As far as I know there is no evidence that a swarm can recognise the face when a person passing, nor that they recognise different people from their faces. 
It also is not clear say, that bees learn to know any of their nest mates as individuals.  I already have mentioned workers’ cliques within hives, but I know of no evidence that such cliques are based on much more than shared smells and shared preferences for particular zones in the hive.  It is in any case questionable how important recognition of individuals might be for insects that live only a few months. 
There is however an important question concerning bees’ recognition of relatedness.  I mentioned the subject of conflict of interest in hives, arising from genetic differences between hive members.  One basis for cliques might be recognition of shared genes.  Olfactory and visual clues that support clique formation, mate choice, or family attachment have been documented in mammals, including humans.  Some such clues are genetically determined.  I do not know of similar research in bees, but I may be behind the times.  If such effects can in fact be confirmed, it would go far to support the idea that workers’ and queens’ relatedness could underlie various patterns of bias in their respective support of reproductive activity. 
Such sub-populations within a colony could have their own “personalities” or “intelligence”.  Like regions in a human brain, they could contribute to the intelligence of the entire colony. 
One way or another, it is not safe to be too dismissive.  Bee colonies certainly do react to certain stimuli such as particular threats in the region to which they are adapted.  In Japan Apis cerana react to the giant hornet.  In Africa they react to the wasps called pirate bees.  Presumably such recognition is innate, but it still amounts to a component in the intelligence of the community.   
Conversely, colonies of sociable birds such as jackdaws certainly can learn to recognise individuals.  Lorenz describes how, once one flock member has been sufficiently alarmed by a given human or large animal, it can pass on its alarm to others by example, and soon that individual is persona non grata to that flock in perpetuity, or even all the flocks in the neighbourhood.  If several individuals achieve that status, then soon all members of that species are automatically regarded as enemies.  In such a situation, the intelligence of the individuals is the basis of the intelligence of the flock. 
There are similar effects in human societies.  Among criminal gangs in various countries there are variously named conventions by which an individual can be named as fair game to the first person who gets a favourable chance to kill him. 
In South Africa at least, it is recognised that such an sentence cannot be cancelled; not because of any fixity of purpose, but because, while it only takes a few words to start such an alert, one could no more recall it with any confidence, than recall a rumour.  In the USA there is the Hell’s Angels cliché that “The Angels is like an elephant.  It never forgets.”  It is no surprise that such memories and alerts are in fact highly unreliable and frequently lead to mistakes and failures, but at the same time they are dauntingly effective and justly dreaded. 
In their inefficient, unreliable, but effective and flexible way, the behaviour of individuals who are party to such sentences strikingly resemble the behaviour of neurons in mental systems.  They really are poorly predictable in their individual behaviour, in the pulses they emit, and the combinations of stimuli in response to which they do emit pulses.  And yet the systems not only work, but are amazingly effective overall. 
Such effects also worryingly recall inter-group prejudice in human society as well as among other animals. 
The “Mongol Horde” principle of attacking an objective is pervasive among large populations, from virus to human, from evolution to military assault.  There are various aspects that vary in relative importance according to the population in question.  The first is that if there are enough attacking agents, one is likely to be lucky and strike a vulnerable point, however obscure that point might be.  This is the kernel of truth in the designer’s traditional lament: “It is impossible to make anything foolproof, because fools are too ingenious.” 
Another strategy for employing hordes of fools relies on the fact that if there are enough participants, some tasks become worth while, even if they are inefficiently approached.  A simple example is the way that ants carry loads too large for just one.  Instead of the conscious sophistication of human stevedores in cooperation, the ants tug at the load from all sides, at considerable cost in labour.  It works well enough to have pre-empted costly development of more efficient algorithms that would have required more elaborate communication and planning. 
One item of sophistication in the algorithm is that it can redirect the ants to divide food items that are too large.  If a group of them cannot lift the prize, they are stimulated to begun the cutting process. 
Note that none of the evolution of the behaviour patterns entailed a high selective cost; their adaptation required very little that was not already present in primitive ants. 
Another strength of the mass attack is that if there is to be any organised resistance to the assault, then the sheer numbers of attackers are likely to disorganise resistance.  For one thing, the defender cannot always know which attackers are ineffectual and may be ignored.  Among honeybees for example, it is well known that they lose the sting after inserting it.  This ignores two salient points to the principle.  Firstly, the sting goes on pumping venom into the victim, behaving like an independent attacker.  Incidentally, it is not just a passive little bag of venom, but a complex device that is continuing with the attack, so the advice to avoid squeezing it, but to get it out with the back of a blunt knife, is an old wives’ tale.  The thing to do is get it out fast to reduce the amount of venom injected; squeezing makes no significant contribution to the injection and may even destroy the mechanism. 
While the sting still is dedicatedly pumping, the now stingless bees do not immediately drop dead or go away.  In pathetic futility they go on trying to sting.  Only it is not as futile as lit looks.  The victim cannot easily tell which bees still have stings, so the bee that at first had been just one assailant, became two: one envenomator and one distracter and intimidator.  This is not a trivial point; it is an example of a sophisticated application of hive (or horde!) tactics and it is innate to the species.   
Apart from foraging and fighting, there are other aspects to the ways in which colonies employ their population resources.  In fact, the sheer variety of colony dynamics in nature almost leaves one at a loss to think of plausible principles that have never been exploited.  Some kinds of colonies are small, just a few dozen members to a mature nest.  Ants of that type usually are predatory and a single good kill is likely to feed a large part of the nest for quite a while.  One also gets moderate-sized colonies of several hundred or thousand members.  Their size limitations vary according to seasonal constraints, available food, nest space, and other considerations.  For instance, a hollow root or thorn might not accommodate a nest for a large population, not even of ants. 
Either way, modest colony sizes offer certain options, but large colonies offer different options.  Large colonies can use strategies of defence, foraging, food culture, reproduction, and building, that simply are not viable for modest-sized colonies.  Driver ant colonies simply have to be large, or they could not accept the attrition of open-air bivouacking and attacking of a wide range of prey sizes.  Termite nests of certain kinds simply have to be large, which means large populations.  It takes thousands of foraging trips for honeybees to make the merest spoonful of honey.  In turn, it takes a great deal of even the best honey to make up a little wax.  Only a large community can afford such costly materials, especially in large quantities. 
In short the use of population numbers is another option that may be used by either the colony or the species in meeting environmental challenges.  The strategies not only vary, but vary in several dimensions.  For example, one way is in the growth of the colony size.  Most ants, termites, social bees, and wasps propagate by sending out young reproductive caste individuals to establish their new nests alone.  This includes some of the species that build the very largest colonies.  In turn it implies that they must start out with the smallest colonies possible and pass through all the stages up to the largest. 
At each stage a successful colony must employ a strategy that suits its size and other circumstances, or it will not survive to reach the next stage, or if it does succeed, then at the least, its reproductive success will be lower than that of a colony that adapted more effectively. 
The question of size seems at first to be a matter of a few trivial details, but several of the principles are fundamental.  Size is a major factor in evolution, in interspecific and in intraspecific competition, in the relationship between the creature and its physical environment, its ability to hide or to survive exposure, in its dependence on its food supply, in general its place and strategy in its ecology. 
There definitely are differences between the relevance of size to individual animals and to colonies, but actually the analogies are much more striking than the differences.  All the variables that affect large animals reappear in related forms in colonies.  The individual colony members may be able to fit into smaller dens than lions, but a colony of a million members will not fit into just any cleft in the rocks, nor can it feed as cheaply as a colony of a few hundred members. 
From the point of view of this essay the details are less important than the fact that the species must have a strategy for exploiting the powers and dealing with the penalties that its particular size entails.  As I have explained, a strategy, evolutionary, communal, or conscious, implies information processing in some sense that we still have not learned to distinguish definitively from what we call intelligence. 
Then again consider interspecific inter-colonial associations of types that are analogous to interactions between individuals.  One gets several kinds.  Simple coexistence occurs between some species of ants, where they share trails and nesting cavities without aggression.  This is startling when one considers how viciously most ants fight even foreign colonies of their own species, never mind alien species.  This mutual tolerance is rather like herds of antelope and zebra keeping company, except that it is not clear in either case, how much of the coexistence is simply for convenience rather than active mutual benefit. 
Such interactions shade into various degrees of mild to severe parasitism.  Some just make mild inroads into the food supply of the host, such as Megalomyrmex inhabiting nests of Sericomyrmex and eating their fungal food without obvious opposition, and apparently without causing much harm, rather like an infestation of rats or pets in a human home.  In contrast, Solenopsis fugax, a small relative of the fire ant, builds its tiny tunnels to interconnect with those of larger species of ants, and robs their larvae and eggs.  There are also many species that undertake “slave raids”, robbing related species of their larvae, eating some, and in some cases even relying on the survivors to act as the worker caste in the nest. 
Predation is another type of colonial interaction.  I have already mentioned giant hornet colonies.  If they are not stopped by beekeepers or by the strategy of Apis cerana, they wipe out the bee colony by killing the adults and carrying off the larvae.  Such conflicts are very suggestive of colonial intelligence, much like the intelligence of bee colonies in scouting for food sources or new sites for hives.  The hornet scouts scent-mark the site when they discover a hive.  Conversely Apis cerana workers react to the scent by instantly attacking the scout and balling round it to kill it by heat.  The behaviour is stereotyped and innate, but the inter-colonial behaviour might be interpreted as quite complex, though still stereotyped. 
There is no doubt in my mind that such effects vary quantitatively and qualitatively in their underlying mechanisms, in the type, speed and persistence of the communications between members of the colony, in the contributions and intelligence of particular members of the colony, and in the genetics responsible for particular types of behaviour. 
Be that as it may certain fundamental mechanisms in colonial behaviour seem to be directly analogous to those within the metazoan body.  They include:
Communication by pheromones that are analogous to hormonal control within the body
Communication by contact and sensory signalling that are analogous to neuronal and synaptic signalling in the bodies of creatures with elaborate central nervous systems. 
Elements of information processing within the communication process are temptingly analogous to both synaptic amplification and inhibition in neuronal signalling. 
There are similarities between some aspects of memory in social and synaptic systems.  
Conversely, we see repeatedly that there are certain differences.  True, our insects (and shrimps and mole rats, and even humans) have certain stereotyped reactions that they are poorly aware of, if at all, but we also see repeatedly that they have some flexibility of action that is vital to the effective activity of the colony.  They have certain capacities for information cultures of various types, that do not closely resemble cultures in the cellular behaviour within our tissues. 

Culture and intelligence:
Elephants, apes, orchestras and Selenites

They had no vision amazing
Of the goodly house they are raising.
They had no divine foreshowing
Of the land to which they are going;
But on one man’s soul it hath broken,
A light that doth not depart,
And his look, or a word he hath spoken,
Wrought flame in another man’s heart.
         Arthur O’Shaughnessy.  Ode.  The music makers
We are left with the question of the role of intelligence in directing communities, as opposed to intelligence of communities as such.  There are no impressive eusocial examples on this planet; probably the closest we come to it is in the naked mole rats.  This is not very satisfactory.  Cerebrally a mole rat is more impressively endowed than a bee say, but it is not clear that this plays much more role in their eusociality than equipping them for socialisation and domination of subordinates.  Among insects the nearest equivalent might be the power struggles among the females of some paper wasp species.  There is little sign of the development of a body of learnt experience being preserved and passed on among such colonies. 
Among the more highly intelligent sociable mammals things may be different.  Experience in culling animals such as elephants has shown that the social structure of herds and the experience of the elders are vital to the well-being of the herd for generations to come.  Senior members not only are the repositories of skills and wisdom, but of security.  They are the community’s repository of experience.  They know where to get water in dry seasons, what to eat, where to trek to when seasons change, where there is danger, and what their social relationship is to neighbouring communities. 
They also socialise the young, punishing harmful behaviour.  Orphans without such guidance tend to grow into delinquents.  If communities that orphans establish survive at all, they are likely to take a generation or two before they recover and develop advantageous cultures from scratch. 
Interestingly, though not surprisingly, experiments in pigtail macaque communities have shown that removing the top-ranked males from a troop has unfortunate effects on troop behaviour.  It turns out that the bosses suppress bullying and disruptive behaviour among hoi polloi.  Furthermore the effect has material significance.  In the absence of the dominant leaders, there is less beneficial social behaviour such as grooming, playing, and sitting together; there is more violence, and less social stability, with more clique formation and re-formation.  The absence of authority radically reduces the quality of life in the troop.  The resemblance to elephant and human sociology is striking.  It is a safe bet that many more such relationships will be discovered as studies advance. 
Communities of sociable primates are similarly dependent on the control and leadership of the more experienced members.  The best-known examples are chimpanzees and gorillas, but there are similar effects among all sizeable primate communities so far studied, including various species of baboons and sociable monkeys.  A species of Japanese macaque was the source of the notorious “hundredth monkey” inanity, in which pseudoscientific Western reports claimed that Japanese researchers had said that when enough monkeys had learnt a new technique, say of washing food or floating grain in seawater, then the technique mysteriously spread through the islands without interpersonal contact and teaching. 
On inquiry the Japanese researchers scouted this claim as nonsense.  They had reported nothing of the kind.  What was far more important and interesting was the fact that the techniques did in fact spread, and that they did so neither mysteriously nor infallibly.  They spread as culture does, by contact, by example, and by fits and starts, not every monkey understanding and adopting the new technology.  As a rule the youngsters were the most receptive learners and exponents. 
Tool use among chimpanzees, such as catching ants with straws, or cracking nuts with branches or stones, spreads similarly, with variations of technique between troops, and with the dimmer members of the troop failing ever to master some of the techniques.  This suggests that there could be fairly strong selection for intelligence whenever it offers an advantage in finding or utilising food sources.  Another point is that among at least the chimps, tool usage and hunting techniques, neither of which is innate, both of which are components of the cultural heritage of the troops, get passed on so stably that ethologists familiar with the subject can identify a troop by the range of techniques they use at a given period.  Of course, such technological accomplishments are not fixed, because occasionally some member of a troop learns something new and may pass it on. 
Chimpanzees, and to a lesser degree of sophistication, baboons also have their political structures within troops, and as a rule the larger the troops, the more complex the structure.  In fact, because alliances and parent-child bonds are such an important factor in their politics, one actually can see rudiments of hereditary rule. 
Social insects have nothing of that type.  The closest any of them get to it is a simple version of pecking order that depends on the ability to recognise individual community mates and remember who is superior and who subordinate.  Perhaps some kinds of paper wasp communities have something like this.  Mole rat social structures are far more sophisticated in this respect. 
Many sociable birds and mammals that form flocks, herds and other communities become far more sophisticated yet.  Within communities they form social sub-structures based on kinship and alliance.  Each form of community control is effective only within constraints of size and environment.  Depending on the nature of their foraging, very small groups may be able to survive better in times of famine, so that is when large communities split up, perhaps only until times improve, perhaps permanently. 
Conversely, in times of plenty large groups, especially of carnivores, can evict or dominate smaller groups.  However, if a community grows beyond the size that the political structure can maintain, then sooner or later the community splits or becomes depleted, often by really serious fighting. 
Interestingly, it is mainly among chimpanzees that politics can become so dominant an activity and so sophisticated, that pernicious violence can break out even in manageably small troops.  Whether this suggests anything about human politics, I leave to the ideological tastes or personal interpretation of the reader. 
Culture and politics of various degrees of complexity occur in wide ranges of animals, and many more examples will emerge as more kinds of community are studied.  Cetacean ethology has shown considerable social complexity, and it also is probable that sociable species of pigs, such as the peccaries, have social relationships far more complex than just a dominance order. 
It is very important to notice that in those species that develop any form of cultural infrastructure, whether innate or by experience, conservation of the information resource and its processing and application become vitally important.  In sociable mammals and birds the most flexible and rapidly developing of the information resources are those in the minds and the educational procedures of the communities.  In insects the innate, stereotyped cultural infrastructures are more important.  However, the situation is not pure; insect control behaviour is not strictly rigid, and many communication behaviour patterns and signalling equipment in troops of mammals and birds, involve genetically determined gestures and physical features.  They might be landing or take-off patterns in territorial birds, ear coloration to highlight a threat in cats, or large fangs, or the colours of gums or eyelids in monkeys. 
There are thousands of such examples, and for the most part the animals themselves patently are not consciously aware of their own signals.  Humans are in no position to sneer; we have a fairly elaborate repertoire of body and voice language.  Part of it is culturally determined, and part innate, but most of us are poor at reading it consciously.  However, within our respective cultures most of us signal and react to signals far more reliably than we realise.  Most humans also are poor at consciously simulating misleading messages. 
Interestingly, a fair number of mammals are fairly good at dissimulation.  So are mating cuttlefish. 
Such interpersonal signalling within a community might be seen as analogous to inter-neuronal signalling within a brain.  Within a species its development obviously is vital. For one thing it is easy to see that anything that reduces waste of resources or risk of injury, and increases one’s dominance in a community, is likely to increase one’s fitness.  For another, in many species the sheer ubiquity and specificity of the selection for signalling and reception of signals, conspicuously or subtly, is incredible.  Nothing of the type could have stood the costs of such specific selection pressure if it were less than evolutionarily necessary.  Even within most solitary species it is important, but within social species, the number and elaborateness of the signalling system increase accordingly. 
It does not follow that the signalling becomes more spectacular; in social insects one does not find much equivalent to anything in the line of antlers or the startlingly extended eyes of the stalk-eyed flies (Diopsidae) or peacock plumage.  Their signals tend to be for specificity of information on a number of subjects, rather than for competitive exaggeration of mainly one point.  Where eusocial insects do signal for spectacular effect, it tends to be behavioural rather than anatomical.  One obvious example is the way bees, even of inoffensive strains, will buzz threateningly around the face and eyes of an intruder.  If intimidation succeeds well enough to render physical attack unnecessary, that is a lot cheaper than stinging.  
Political structures are of particular importance in the present context.  They are an example of where individual intelligence and whatever passes for the intelligence of the community affect each other directly.  Techniques such as washing food or cracking nuts might be important; fortuitously they might even be components of the culture that turn out to be vital to the survival of the community, but political control is where the mind of the member directs the action of the community. 
Direction of the action of the community by leaders sounds impressive, but not everyone is convinced of its importance.  Some historians claim that nations such as the Roman Empire had successions of rulers ranging from something like secular saints to the most appalling despots and parasites, without detectably affecting daily life and commerce of the bulk of the citizens.  Personally I regard this view with suspicion.  Granted that the effect of the ruler gets filtered and moderated through the administration, as brilliantly satirised by say, the “Yes Minister” series, the view of the historian is necessarily in many ways the view of the outsider, at once simplistic, selective, and superimposing his own interpretations with limited scope for falsification.  One is reminded of the closing paragraph of H. G. Wells’ short story “The Star”, in which an event that nearly destroyed humanity looked trivial to Martian astronomers. 
For counterexamples even in pre-modern times, there were the Chinese periods of imperialism, followed by the inward-turning of their policies.  Both affected every aspect of their people’s lives.  Genghis Khan, Alexander and a number of major autocratic leaders were of crucial importance in the lives of their nations and of their people.  In modern times with the more advanced technology permitting more elaborately controlled infrastructure, we have seen a whole series of despots, from the last of the Tsars to the first of the so-called communists and nationalists.  There has not been a time in the last few centuries when there have not been peoples who lived and died under the control of pernicious despots, often genocidal despots.  Their control might not have covered every detail, but it was enough to affect daily life in every respect, whatever things seemed like to outsiders at a safe distance. 
Be that as it may, the point is that here too, we see the interaction between the character of the community and of the individual brain. 
Mode of rule is only one aspect of such interaction.  Culture is another.  So is infrastructure.  Both are affected by the inventions and procedures instituted by individuals.  This is not an effect that we can see in the eusocial insects; palaeontology is not good at preserving the deeds that gave rise to selection.  This is a field where the Just-So Story and the argument from analogy come into their own, not to prove the detail of history, but rather to show that a particular type of course of events is a reasonable conjecture, consistent with the realities of biology as we observe them today, and a basis for research on principle and mechanism. 
At least one experiment has been tried at having an amateur orchestra play without a conductor.  I understand that the orchestra acquitted itself very creditably, but it was a tour de force rather than an influential development.  In fact, historically the original function of the conductor was far less formal than we take for granted today, and the development of the role was suggestive from the point of view of this essay.  One could say that the conductorless orchestra is rather reminiscent of real life eusocial communities.  There is a score (analogous to the innate behaviour of the individuals), and a beat and pitch (analogous to the environmental releasers of behaviour). 
However, once the role of the brain and creativity of the conductor became established in musical practice, the interpretive enrichment and flexibility of the orchestral behavioural repertoire attained new dimensions of complexity and size.  Conductorless performance was effectively relegated to small groups of performers such as in chamber music.  Again, the analogy of the ruler of the state becomes suggestive.  Certainly the conductor of a large orchestra cannot control the character of his performers in detail, nor can he control the character of the group in detail, but he certainly can affect them profoundly in many ways, including in the speed with which his central authority can bring about changes in performance. 
It is interesting to note that there has been no eusocial species whose individuals have been of a high order of intelligence relative to say apes and pigs.  Probably naked mole rats come the closest, but to argue that they represent highly intelligent eusocial individuals, involves slippery lines of reasoning that threaten to end in simplistic arguments. 
It is arguable that such a thing, if possible in principle, is implausible in practice. 
However, H. G. Wells, the greatest science fiction genius in history, head and shoulders above any rival so far, supplied us with a thought experiment that was characteristically ahead of his time.  His Selenites with their ruler the, “Grand Lunar”, in “The First Men in the Moon” were exactly that, highly intelligent eusocial individuals, and his short story “Empire of the ants” refers chillingly to related concepts.   

Evolution of intelligence

There are no whole truths: all truths are half-truths. 
It is trying to treat them as whole truths that plays the devil.
                Alfred North Whitehead

A few generalisations on this aspect of the subject may deserve consideration.  Viewed simplistically, evolution of intelligence is fairly simple in that it looks like any other adaptive evolution.  There is a genetically determined set of variables in a given species in a given environment, and where some particular values of those variables directly or indirectly affect reproductive success in a population of significant size, adaptive selection follows as a matter of course.  Many such genetically determined variables affect behaviour patterns, including flexible behaviour in response to external stimuli. 
Experiments, both deliberate and incidental, in artificial selection for behaviour patterns, have been very revealing.  By artificial selection breeds of dogs have had their natures changed within decades.  Herding behaviour has been selected for in sheepdogs, coach escorting behaviour in Dalmatians, and large breeds have successively been bred for viciousness and gentleness.  Within about a dozen generations of concentrated selection of guinea pigs, hamsters and other small pets for either docility or aggression, breeders succeeded either in producing almost paralytic docility or practically pathological viciousness.  And in Europe selection of the honeybee for docility has produced colonies with behaviour almost unbelievable to the African beekeeper who must deal with more recently domesticated lines or subspecies. 
Selection for intelligence in the weak sense of behaviour adapted to their environment has affected all fairly complex animals and colonies and has produced results in various degrees of complexity and sophistication.  The complexity of colonial behaviour (as opposed to the behaviour of the members of the colonies) could match the complexity of the behaviour of individual animals practically step for step, and it proves to be roughly as modifiable through selection. 
However, one thing that I cannot find with any degree of confidence, is an example of colonies that have generalised intelligence that permits flexible response to almost arbitrary new challenges, such as individual specimens of primates, pigs, some carnivores, and some birds exhibit.  Instead, colonies invade adventitious niches where their resilience, small size, and inexhaustible opportunism are advantageous.  Examples include the Argentine ant (Iridomyrmex humilis), Pheidole megacephala, and the pandemic Formosan termite. 
However, a few species of non-social cockroaches spread just as successfully.  Clearly there is no general way to predict when social intelligence will be responsible for the success of a species, much diagnose that it amounts to intelligent response to complex challenges. 
The evolution of colonial behaviour however, is impressively supported by observation of extant species.  Unfortunately that sort of evidence necessarily reduces to the classic type of “Just-So Story”, but as long as one is not arguing with radical fundamentalists or naïve positivists, such reasoning is enough to illustrate how innate intelligent behaviour can be adapted to the challenges and opportunities of the environment.  We may not be able to say just how the species involved in fact, but often we can point to other species that demonstrate that particular hypothesised intermediate developments are indeed viable. 
In particular, in principle we also can see how the primitive foraging and parental behavioural components of solitary insects could have become elaborated progressively to amazing levels of sophistication.  As things stand extant species illustrate behaviour corresponding to putative stages in the evolution even of the dances of the honeybee. 
One remark about evolution of intelligence in insects (in particular in social insects) is that they have been extremely conservative in their modes of communication.  They use few chemicals for many purposes, and say, the honeybee uses similar dances for indicating sources of water, nectar, and new nesting sites. 
I have severe reservations about this remark. 
The observation is true enough as far as it goes, except that recent work has greatly multiplied the range of compounds that insects are known to use for signalling.  In guiding mating activity, some wasps use at least seven successive compounds for control of the seven successive stages of the mating act.  I already have remarked on the startling degree to which some orchids have adapted to relying on those wasps for pollination.  The key point however, is that the very fact that the orchids had time to evolve such an elaborate system does imply that the wasps had been using the same system for a long time, probably many millions of years. 
Anyway, biological systems in general, not just insect signalling systems, are very conservative when a change to a working system is difficult to bring about without teleology.  This is particularly so when multiple complementary components of a complex system must evolve in synchrony if the system is to survive at all.  Again, my attached essay on pheromonal evolution discusses some aspects of this. 
Superficially and qualitatively, there are few truly incomprehensible mysteries specific to the evolution of intelligence as such, whether colonial or individual.  When genetic information supports the basis for nervous control, adaptation in response to selective pressures seems to be quite as effective as adaptation of genetically determined somatic features. 
Just as the dinosaurs and whales produced the largest bodies, complete with the necessary adaptations of support and circulatory systems, so Homo has produced the largest increase in abstract, flexible intelligence so far.  It remains to be seen whether we can thereby match the duration of the dominance of dinosaurs, which (as an enormously assorted lot with a vast range of internal succession), lasted for well over 100000000 years. 
As for the idea of a subjective colonial intelligence, I find it hard to imagine one in the colonial organisms we know, if one omits the view of metazoans as colonies of cells.  However it remains difficult to justify my failure of imagination as a valid argument, seeing that we have not yet learned even how to measure subjective intelligence in ourselves, let alone in alien communities. 
There has however been a major evolutionary leap in our intelligence.  It is not at all clear that any other species ever has developed non-trivial levels of inductive or deductive thought, or imagination, either individually or in the form of community thinking. 
Still, anecdotal evidence suggests that something like imagination or deduction might exist in non-primates.  I have seen a zoo hyena lying immobile till it saw me trip over a raw chicken leg camouflaged with dust.  The morsel no doubt had dropped from the feeding trolley some hours before.  Even as I stooped to pick it up without so much as a betraying glance at the hyena, it charged the fence, correctly anticipating my throwing it into the enclosure.  The leg barely had time to reach the ground.  To me this suggested awareness that the food had fallen there, and that a human might pick it up and oblige.  It also suggested a keen personal interest in the outcome. 
Again, a hand-reared black rhino has been known to open its enclosure, and release the dogs to find its human foster mother, who had gone out onto the reserve, leaving the animals behind.  Pigeons have been experimentally shown to be able to learn specific techniques to open food containers by watching other, pigeons experienced in manipulating the apparatus. 
I am not aware of anything of such a type in insects, much less in eusocial communities.  In eusocial species?  Do symbolic language, complex nests, abandonment of nests threatened by ants or fire, mean anything?  There is a long road before we can formulate such questions usefully.  The day that we succeed, we will have made giant strides in the comprehension of intelligence in any form, and that would make the study of communal intelligence worth while whether it were interesting in its own right or not. 
Where we go from here, in what form, and how fast, is unclear.  The fact that we are monkeys rather than termites precludes human society in its present form from undertaking really worthwhile, large, long-term projects.  We operate on conflicting interests that bedevil our individual tribes, let alone our planet.  Until, as a species, we master the virtues of the insect colony, our long term future as a species is discouraging. 

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