Horses in Company E-Book

Lucy Rees

• Herausgeber: J A Allen
• Kategorie: Lebensstil
• Sprache: Englisch
• Veröffentlichungsjahr: 2017

Beschreibung

Horses in Company is an in-depth study of the social interaction and hierarchy of horses in the wild. The book discusses how the characteristic behaviour of the herd has developed in response to the natural selection pressures in their evolution, namely that horses are prey animals and not predators, and how this governs the behaviour of domestic horses living together. Through these studies, the book aims to help the reader understand the difficulties that arise between people and horses due to misunderstanding equine behaviour and motivation. Of great interest to all horse owners and riders and illustrated with 60 colour photographs.

Leseprobe

HORSES

in COMPANY

Lucy Rees

J.A. ALLEN

First published in 2017 by

JA Allen

JA Allen is an imprint of

The Crowood Press Ltd

Ramsbury, Marlborough

Wiltshire SN8 2HR

www.crowood.com

This e-book first published in 2017

© Lucy Rees 2017

All rights reserved. This e-book is copyright material and must not be copied, reproduced, transferred, distributed, leased, licensed or publicly performed or used in any way except as specifically permitted in writing by the publishers, as allowed under the terms and conditions under which it was purchased or as strictly permitted by applicable copyright law. Any unauthorised distribution or use of this text may be a direct infringement of the author’s and publisher’s rights, and those responsible may be liable in law accordingly.

British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library.

ISBN 978 1 90880 957 5

Contents

Introduction

CHAPTER 1Thinking About Behaviour

Reflex and innate behaviour. Early work by Tinbergen and Lorenz. Learning. Tinbergen’s four questions. Motivation. Strategies, costs and benefits. Emotion. Anthropomorphism. Applied ethology.

CHAPTER 2What Horse Evolution Can Tell Us

Natural selection pressures evident in the fossil record. Domestication effects.

CHAPTER 3Equine Ethology Studies

Feral horses: natal bands, stallions and mares. Bachelor bands. Natal dispersal. Multi-stallion bands. My own studies: different populations.

CHAPTER 4Natural Life, With Puma

A stampede. Analysis of defence behaviour. Self-organizing behaviour and the flight algorithm: cohesion, synchrony of velocity and direction, collision avoidance. Initiators of changes.

CHAPTER 5Social Life Revisited

Cohesion and synchrony in everyday life. Leadership but no fixed leaders. Collision avoidance or respect for individual space. Keystone characters? A self-organizing society.

CHAPTER 6Interactions

Band and herds. Stallion tolerance and its influences. Population dynamics of the llanos herd. Survival. Competition for resources? Male competition for mares. Bachelor strategies. Stallion attractiveness to mares. Paternal protection of foals. Two-stallion bands. Social relations and bonds within natal bands. Aggressive actions and their causes. Conclusions.

CHAPTER 7‘Horses Have Strict Dominance Hierarchies’

Definition of dominance and the history of the concept. Measurement. Correlations. Results of horse studies. Problems: unacceptable assumptions, practices and parallels. No single hierarchy. Re-interpreting dominance studies. Feral, free-living and domestic groups. Learned aggression in imposed resource competition. Stress-related aggression.

CHAPTER 8The Dominance Hierarchy Paradigm in the Horse World

Misinterpretations of ethological statements and of behaviour in popular literature and everyday horse-keeping. Bacon’s idols.

CHAPTER 9Changing the Paradigm

New fields of research in equine ethology. Practical considerations regarding attitudes in handling and riding horses.

Notes and References

Index

Introduction

This book presents a radically new view of horses’ social relations and organization, a view that inevitably affects how we interpret our interactions with horses. It proposes that horses’ social life and relations developed in response to the natural selection pressures in their evolution: predators. Horses are prey animals.

In itself, of course, the idea that horses are prey animals is by no means new; we have been paying lip service to it for years. Yet we do not seem to have taken its implications seriously: that it is the focal point of the way horses live together. The current interpretation of their social life is that horses interact according to dominance hierarchies, which have nothing to do with predators.

Throughout many years of working with horses, and especially in resolving problems that arise between horses and people, I had realized that dominance is not a useful concept. Analysing horse-human relations in terms of dominance creates far more problems than it solves. The same has been found to be true of dog-human relations.

However, until a few years ago I had no alternative theory or way of analysing social interactions, although having dispensed with dominance as a factor, others started to become clear and useful in practice: the horse’s acute awareness of our body language, his cooperation, his dislike of physical restriction and, above all, his coordination with others, whether horse or human. But they did not come together as a coherent picture. I lacked a paradigm.

I arrived at the new paradigm through observations of feral horses and especially of their behaviour in the face of predator attack, a curiously neglected field of equine ethology. Gradually I came to see that their whole social organization and relations reflect their adaptation to the ever-present possibility of predator attack. The exact ways in which they behave to escape successfully are reflected in their everyday lives – even those of domestic horses that have no practical experience of predators.

This is an ethological approach: why do animals behave the way they do?

I would like this book to be accessible to those many thoughtful and intelligent horsepeople who have no base in ethology (as well as those who do). I have therefore thought it necessary to start with a brief explanation of some of the ideas that are used later in the book. This is not a potted ethology textbook but a way to avoid breaking a logical flow with explanations of the concepts used.

Behaviour helps an animal stay alive and pass on its genes to the next generation in its own natural environment, in various ways. Some behaviour has a strong genetic basis, and is acted upon by natural selection just as the exact form of an animal’s body is.

Therefore, we must look at the selection pressures revealed in the course of horse evolution. Predators, of course, are one.

The classic and important studies of feral horse behaviour show that they all adopt more or less the same solution to life’s problems, and introduce us to some terms and concepts. These studies help us see what behaviour is constant in all natural contexts. Although I have not wanted to delve deeply into domestic horse behaviour and lose the thread of the story, I hope that thoughtful horse-owners will find much to reflect on.

My own studies, which come next, have not been published before. Since the exact conditions and methods of study can influence results, scientists are careful to detail them, which often makes boring reading to the non-specialist. I’ve tried to convey a more living picture of what it is like to be a field ethologist; the interactions in the herd, the extreme efforts that horses make to avoid competitive conflict, the questions that arise and the ethological approach to answers.

Thus prepared with a general picture, we can examine exactly what happens, and why, when attack comes. My understanding of my observations owes much to an up-and-coming field of research that has not been applied to horses before but is providing great insights into herd, flock and shoal behaviour, using behavioural algorithms for self-organizing group movement.

What then became clear to me is that the factors that govern successful escape also govern social interactions and organization within a band or herd in more peaceful moments, too. We see that social behaviour is adaptive, following a coherent logic given that horses are prey animals. Within feral bands they do not compete against each other but collectively against predators, and maintain social relations that allow them to behave appropriately and instantly in the face of situations, grading from changing their maintenance activity to life-threatening attack.

This hypothesis, or paradigm, since it changes the way we interpret social relations and organization, is based on my own observations, which, for practical reasons, tend to be more qualitative than quantitative. They do, however, suggest a large number of testable questions that, I fervently hope, will be examined more fully in future research.

The paradigm I propose is not the generally promoted and accepted one: that horses interact according to dominance hierarchies. There are serious flaws, including a lack of consistent evidence, in the ethological arguments for dominance hierarchies; there is also a wide gulf between what ethologists mean by dominance and what the general public means by it. A critical examination of these problems therefore follows.

Yet the concept of dominance is so deep-rooted in equestrianism, and indeed seems so self-evident to many horse-owners, that we also need to consider whether there are alternative explanations for the behaviour that we see. We find that there are many reasons why we interpret things in certain ways, even when these are not true. In the same way, we see the sun rise in the east and set in the west; even when we know we are perched on a rolling ball, we find that fact difficult to appreciate.

Finally, we consider some of the implications of this change of paradigm, both ethologically and in our daily interactions with horses.

CHAPTER 1

Thinking About Behaviour

Behaviour is adaptive. It helps an animal survive and leave its genetic mark on the world through its offspring. Cats’ claws, bats’ echolocation, fishes’ fins and horses’ tails, or any other of the myriad physical adaptations that enable an animal to survive in its particular way of life, would be of no advantage if the animal did not use them properly.

A great deal of behaviour is hard-wired, under genetic control. Natural selection operates on this innate behaviour as it does on the genes that govern bodily form, weeding out the animals that do not behave appropriately in their natural environment and leaving the ones that do to pass on their genes to their descendants.

The simplest form of hard-wired behaviour is reflex action. You do not have to think about contracting your iris when emerging from a dark room into bright sunlight, or how hard you need to push against the ground (and with which muscles) to stay upright. On a more complex level, you do not need to think before avoiding a blow coming at your face, if you see it in time. The response is automatic.

Many animals, insects for instance, operate on this automatic level, which is not necessarily simple: their behaviour can reach remarkable complexity, as it does in ants and bees. Karl von Frisch, who with Niko Tinbergen and Konrad Lorenz won the Nobel Prize in 1973, unravelled the secrets of the honeybee’s communication regarding a source of nectar, a remarkably cleverly coded dance on the surface of a comb in the hive. His classic book The Dancing Bees describes the observations and experiments that helped him arrive at his conclusions.

This type of automatic behaviour, involving the whole animal in chunks of behaviour rather than single actions, is what was generally called instinctive behaviour, Lorenz and Tinbergen’s particular field of study.

These two, ethology’s great founder fathers, set out to make a science of ethology, to find the unifying mechanisms underlying instinctive behaviour. They had a long-lasting friendship and collaboration, the immense fruitfulness of which lay partly in their shared interests and partly in their differences in approach. Tinbergen was primarily a naturalist and bird-watcher: he watched animals in their natural habitats, fascinated by the wealth of adaptations that their behaviour showed to different ways of life. Lorenz shared his life and farm with a variety of domestic and tamed animals that he observed and experimented with, delightfully described in King Solomon’s Ring (1949). He had, then, more opportunity to see how instinctive behaviour, adapted to cope with situations occurring in an animal’s natural life, could misfire when the animal found itself in situations outside that lifestyle: in domesticity, for instance.

Working mostly on birds and fish, the two saw that standardized little chunks of behaviour could be triggered or ‘released’ by specific stimuli – a colour, a movement, a sound – that all animals of the same species reacted to in the same way, without having to learn how to do so. Both the recognition of the releasing stimulus and the reaction were inborn, innate, built into the wiring in some kind of way.

Lorenz, eager to give ethology a theoretical base, invented a mechanical model of instinct. He saw that, sometimes, the longer the animal had been without performing a particular little chunk of behaviour, the more easily this was released, and incorporated this feature into his model.

Scientists love models. Models can be formulae, drawings, flow diagrams or, as in Lorenz’s case, a hypothetical structure. Models can be tested by experiment or observation to see if they really work in all cases. Lorenz’s did not. The result was years of investigation and argument about whether all instincts worked the same, about what physical neurological structures could correspond to the various parts of his model, about whether this experiment really tested what it claimed to have done, and the like. Eventually the model and even the term instinct were dropped. ‘Innate’, the term now used, had no historical connotations or pitfalls about assumptions; it correctly implies that this type of behaviour has a hereditary base common to all animals of one particular species.

No one ever said Lorenz did not see what he said he saw: what they debated was his interpretation of his observations. Ethological observations are ‘clean’: they state baldly what the animal did, when and where, excluding interpretations of why. ‘The horse tried to kill the man’ is not an ethological observation. ‘The horse repeatedly struck the fallen man with his forefoot’ is nearer the mark, but is improved by adding the circumstances: ‘the man threw stones violently at a horse trapped in an alley. The man stumbled and fell. The horse repeatedly struck the fallen man with his forefoot.’ Such an observation, like so many, is open to various interpretations.

Lorenz was an ethologist. His observations were valid even though his model failed, and later theories or models had to explain them.

Tinbergen was less inclined towards proposing universal theories, and more to finding out exactly what happened. He set about investigating what were the properties of a releasing stimulus that made it innately recognizable, and how exact they had to be. When parent herring gulls arrive at the nest, the chicks peck at their beaks and the parents regurgitate food for them. Tinbergen found that what stimulated the chicks to peck was not the parent or the food, but a red spot on the bill. He painted a stick white, painted a red spot on it and moved it up and down in front of a nestful of chicks. If the spot was the right colour, in the right place and the stick moved at the right rate and angle, they pecked; if not, they did not. Nothing even faintly resembling a parent bird was necessary.

Fig. 1.1 The new-born foal does not have a concept of searching for milk: he has an innate urge to put his head between two pillars capped by a shadow. Often, as here, this urge brings him no benefit. But when by chance he is rewarded by milk, he soon learns which two legs to aim for.

Foals are born with an innate urge to get to their feet. They then look between two upright pillars capped by dark shadow. They are not looking for anything: if they see the right image, they put their heads there. Sometimes they put their heads between the mare’s forelegs, sometimes between the hind. At some point they hit an udder bursting with milk. Wriggling their lips, they sooner or later find a teat in their mouth, and reflex sucking takes over. This splendid reward shapes the searching behaviour, making clear that some pairs of legs have teats and milk while others do not. After a couple of days they have learned which are valid and make no more mistakes.

The foal’s reaction, then, shows the same pattern as that of the herring gull chicks’ gaping at Tinbergen’s stick with a red dot: both animals react to a drastically simplified but exact symbol, not their entire mothers. Innate releasing stimuli (also called releasing stimuli, trigger stimuli or sign-stimuli) and programmed responses (fixed action patterns and the like) can suffer the same problems as insects’ automatic reactions: the response may not be such a good survival ploy when the animal meets with unusual conditions that fortuitously provide the basic characteristics of a releasing stimulus. Wagtails can spend hours fighting their reflections in car wing mirrors. I once went to sleep in a Colorado forest and was woken by the purr of a humming bird’s wings next to my ear. It was making as if to feed from the red letters on the book I had laid down: humming birds feed from red flowers. A newborn foal I was watching repeatedly put his head between a tree and a gatepost linked by a heavy bar: two uprights capped by a shadow. He ignored his mother.

During the whole magnificent sweep of evolution, of animals, plants, bacteria and viruses inventing new ways to solve the problems of surviving, there have been some consistent trends if we look at newcomers. Evolution is blind: it does not lead anywhere except survival and procreation. There is no endpoint to be arrived at, no apex or pinnacle, but a continuous flux and adjustment to continually changing conditions. Nonetheless, if we consider new life designs in the long history of living beings, there is a consistent trend towards less wastage. Huge numbers of insects die because they cannot modify their innate reactions in inappropriate circumstances, like moths attracted to candle flames. In terms of behaviour, less automatism and more ability to react to a complex of stimuli, not just one outstanding feature, mean fewer suicidal or silly time-wasting mistakes. Decisions can be made as to whether or not to act, or how exactly to do it. In mammals, increasing brain size and ability to analyse and consider situations means that innate reactions are less fixed: not so often is there an automatic reaction to an innate releasing stimulus, as a tendency to find certain stimuli curiously attractive, fumble about a bit and, partly by chance, discover that one way of behaving brings unexpected satisfaction. Next time, the fumbling about diminishes and the animal is more goal-oriented: it now knows there is a goal, instead of being driven by a vague compulsion. Innate reactions provide the conditions for learning.

Fig. 1.2 Many simple behaviour patterns have an innate base but are perfected by experience. Standing with their necks overlapping comes naturally to these foals, but what comes next is not so obvious. With practice they will mutually groom more efficiently. (Photo: Javier Solis)

The formula S → R is a simple way to say that a particular stimulus provokes a particular response. In innate behaviour, the link between the two is, so to speak, ready-made in the animal’s mind. In learned behaviour, the link is forged or changes in response to the animal’s experience of the consequences of its actions.

LEARNING

Learning takes many different forms that, in the field, are often mixed so that what is going on is not clear. For this reason its investigation took place in the laboratory where the conditions could be simplified and controlled. Nevertheless we mostly see the same features in animals’ natural lives once we know what we are looking for. The foal’s learning how to get food is an interaction between innate and learned behaviour and reflex action.

The simplest form of learning, shown even by animals that have no brain at all, is habituation: losing a response to a stimulus that normally provokes it. Horses are particularly prone to fear any moving thing that has not been proven safe, but after a few startle responses that have no consequences except a waste of energy running away, they habituate, or ‘get used to it’. You cannot spend your life running away from butterflies or rabbits, though foals start by doing so. Like most young mammals, foals are helped to distinguish between what is genuinely dangerous and what is not by their mothers’ attitudes.

Fig. 1.3 Investigation or gradual, free-choice habituation: the youngsters lose their fear reaction by exposing themselves to the stimulus.

Investigation is a kind of self-programmed habituation. After removing himself to a safe distance, the horse returns cautiously to the unknown thing, watching and listening, ready to flee again if it reacts adversely. If there are no consequences, he finally gets near enough to examine it with all his other senses – smell, whiskers, lips, teeth, feet – to identify it for further reference.

In associative learning, a new S→R link is forged. Classical conditioning links a new stimulus, previously irrelevant, to an already existing response. Thus horses learn to neigh at the sound of our car. Operant conditioning forms a new response, a new behaviour. Horses also learn to be remarkably adept at opening stable doors. What forge and strengthen the S→R link are reinforcement and repetition.

Reinforcement can be positive or negative. Positive reinforcement is usually called ‘reward’, which makes us think of food. True, food is powerful reinforcer, especially when an animal is hungry. It makes the animal repeat whatever was done before to get it. But any pleasant feeling, like being in good company for a social animal, is reinforcing. So is habit, or repetition.

Negative reinforcement involves an animal finding itself uncomfortable and, as a result of its reactions, becoming comfortable again. The second time round, it is quicker to repeat what it did previously. Thus positive reinforcement involves something pleasant being added, while negative reinforcement involves something unpleasant being taken away. In training horses, negative reinforcement is widely (and by some trainers exclusively) used: pressure and release. A moment’s thought shows that it is not a particularly pleasant way of learning: the horse would prefer not to be made uncomfortable in the first place. He learns just as fast as with positive reinforcement, but is not motivated to ‘go to class’. Rewards, on the other hand, do motivate, for any animal is eager to be rewarded. Once he has learned what to do to get a reward, he keeps performing even when he is not rewarded, although occasional rewards keep both motivation and performance at a high level.

A bunch of feral horses find a distant tree with rich fruit in autumn. After finishing it they wander back to their normal grazing area, to return to the tree (operant conditioning: they have learned the way) occasionally but fruitlessly throughout the year. In autumn they are rewarded again (infrequent rewards keep them performing). In time they may connect the tree’s fruiting with the appearance of blackberries in the area they normally graze (classical conditioning), and cut down on their fruitless visits. As the years pass, the tree is simply where this bunch goes in autumn, a cultural habit.

Punishment, or unpleasant or frightening experience, is capable of temporarily suppressing a reaction, but not of destroying an S→R link: sooner or later the animal will revert to its previous response and keep doing so unless punished again, or rewarded for behaving differently (counter-conditioning). Punishment cannot create a link; it provokes fear of the situation in which it occurred. In horses, punishment provokes avoidance, their usual reaction to a fearful situation. With repeated punishment a horse recognizes, and takes evasive action at, the preliminary signals that the situation will recur: avoidance learning.

To return to the example above: the farmer, enraged, protects his fruit tree with an electric fence. The horses, punished, learn to recognize and avoid it at first sight, though they still return every autumn. Satisfied, the farmer stops electrifying the fence. Sooner or later the horses revert to touching it, find it does not hurt and push through, for which they are rewarded.

Avoidance learning is the secret of success of threat, which is followed by attack (punishment) if ignored. Finally, one animal may avoid the mere approach of another even without threat: passive displacement.

Punishment, like reinforcement, can be positive or negative. Positive punishment is the infliction of pain, discomfort, anything disagreeable. Negative punishment is the removal of something desirable. Since horses desire company, being chased away by others is a form of negative punishment.1

Discrimination learning involves differentiating between a valid stimulus, one that brings reinforcement if you respond, and one that doesn´t. Horses discriminate carefully between hundreds of different types of plant, just by feel. As foals, they nibble tiny samples of plants. Poisonous plants generally taste bad, but the foal does not eat enough to poison himself: he is merely sampling, learning to connect the feel of the plant to his whiskers and lips with a good or bad taste. Later, he will flip aside the bad ones with impressive speed and certainty, using that wonderfully mobile upper lip, as he grazes.

COGNITIVE PROCESSES

Beyond the level of pure connectivism or behaviourism are cognitive processes: the collection and recollection of information, together with its analysis and integration, to form concepts and make decisions. Horses, dogs, chimps and most notably humans are capable of examining situations and considering possible outcomes before acting appropriately rather than simply reacting to cues, whether learned or innate, though we do not always use this ability. It makes us curious. We like information for its own sake, not necessarily with some immediate purpose in mind but because knowing how the world works satisfies an innate urge.

Horses are naturally inquisitive and exploratory. Most domestic horses, though, have little opportunity to investigate, explore, or reach their own conclusions and decisions, for they are too restricted and controlled. Like other animals, horses learn to learn; if they are brought up in dull, unvarying environments, repeating the same mindless exercises, they have little chance to learn and do not strike us as being bright. As the Italian ethologist Francisco de Giorgio insists, limiting our training to behaviourist control techniques annihilates their cognitive abilities and their satisfaction in using them.

One of horses’ particular talents is that of making mental maps of where they have been; it includes the ability to predict how known places or routes link up with each other without ever having explored the link before. This is cognitive learning without reinforcement or punishment. It is seldom allowed to develop, but horsemen who work in difficult terrain – on mountains where thick mist descends suddenly, in dense, monotonous scrub or confusing forests – know and treasure this ability, unconsciously fostering it by allowing the horse more freedom of choice in picking his way than would a dressage rider.

Are horses capable of forming concepts? Experiments show that they learn concepts like always choosing the bigger (or smaller or darker) of two images on a screen, but the best examples are in working horses, cowponies, logging horses, general farmwork horses and the like, who show that they genuinely understand their work despite its huge variability. Their introduction to it is usually via stimulus/response learning, but after a period of confusion and unwillingness, most grasp the idea of what is to be done, do it eagerly and even invent ways of resolving tricky situations without any prompting. A behaviourist might say that this is simply avoidance learning (I can feel he wants me to move so if I do so now I’ll avoid getting kicked), but avoidance learning does not produce eagerness. Many years ago I met a logging horse, quite alone, manoeuvring a huge tree trunk down a steep wooded slope. Whenever it snagged, he made careful tests to feel and calculate angles before throwing his full weight in precisely the right direction to free it. Fascinated, I followed him down; if I offered suggestions he told me to mind my own business, which at that point was realizing how appallingly dull normal training is for them. They like using their cognitive abilities.

Mammals with brains as highly developed as horses have an enormous capacity for learning, which allows them to adjust to the particular conditions they find themselves in. They retain reflexes that deal with simple reactions, automatically controlled patterns like gaits and, especially for the newborn to help them through their first days of life, innate responses of the type described by Tinbergen in birds. But they have shifted the control of most of their behaviour from automatic to formed-by-experience, though what pushed them into the experiences was often an innate urge. Horses, being social animals, seek company, an innate urge that nonetheless does not provide them with the knowledge of how to behave once they find it. In a natural life, a foal learns to distinguish between individuals, to communicate by recognizing and using signals, to respect others’ individual space, to know how to invite another to play and make friends. He sees his father sexually aroused and mating, sees newborn foals, and death, too. All happens so gradually and smoothly that we do not realize what a wealth of learning social life requires and supplies. Domestic horses who have grown up without company are often social inadequates, creating havoc in the company they so yearn for; young mares are often terrified by a stallion’s display, and some are scared of those weird, wobbly, unhorse-like foals they produce. They are ignorant, through no fault of their own.

TINBERGEN’S FOUR TYPES OF QUESTION

When we watch animals going about their everyday lives in their natural habitats, the core business of ethology, we do so with the aim of describing first of all what they do and then finding out how, why and what factors influence it. To find that this behaviour is innate, learned or produced by an interplay of the two is to look at the way it is produced, the how. It does not answer why the animal should behave that way. Tinbergen, who had a particular, fierce clarity about framing questions so that they might have concrete answers, saw that the answers to ‘why?’ questions fell into four groups. This classification has been a lasting help to ethologists in framing their questions.2

1. Proximate Cause

What was it that triggered this behaviour? The trigger might be a stimulus, a situation, a rise of a certain hormone that makes certain stimuli more interesting, or a bodily state, like being cold. The stimulus might be innately recognized or become significant because of experience; the reaction may be innate or learned or, most often, an inextricable mixture of the two. A proximate cause is what provoked the behaviour.

2. Final Cause

How does this specific behaviour affect the animal’s chances of surviving and breeding successfully? What are the long-term effects of such reactions on its life, or the survival of its offspring? Final causes may be stunningly obvious – eating keeps you alive – or may be difficult to tease out. Stallions urinate on their mares’ dung, which might seem to convey a message of ‘she’s spoken for’ to another male finding it, who smells it carefully; he then performs Flehmen, raising his head and rolling his upper lip back over his nostrils to prevent air escaping, pumping it into the vomeronasal organ within the nose. This organ is specialized in the detection of pheromones, smell-messages that change the hormones, and thus the behaviour, of the animal who detects them. An in-season mare’s dung contains such a pheromone that sexually excites a male, who rushes off to find the mare. Kimura (2001) noticed that when, marking his mare’s dung with urine, a stallion does not urinate in the usual jet, which would suffice to tell others of her alliance, but sprinkles it over the dung in a peculiar way. On collecting and analysing samples, Kimura found that the stallion’s urine neutralizes the mare’s pheromone: by sprinkling the dung all over with urine the stallion maximizes its neutralizing effect and fools would-be competitors. Now we know the final cause, even if he does not.

Fig. 1.4 Smelling his mare’s dung, this stallion detects she is in season by doing Flehmen. He then sprinkles the dung with urine, thus neutralizing the telltale pheromone. (Photo: Javier Solis)

3. Development

Some behaviour is appropriate at one stage in life, some at another. Newborn mammals suckle by reflex; adults do not. Young female animals do not produce full sexual behaviour: they do not have the resources to complete their own development into healthy adults and produce offspring at the same time. Animals are by and large parsimonious in the invention of new behaviour, so that what has been an appropriate reaction at one stage may be taken over for another use at a later stage. What has been play becomes courtship; what has been a wolf cub’s begging for its mother to regurgitate food becomes symbolic allegiance to the pack leader. I had a pet starling who, before fledging, gaped vertically when I offered food. At fledging, he gaped horizontally, so I had to drop the food in front of him. Little by little he directed his bill-opening downwards, often missing the food entirely; a couple of weeks later he would stroll around the field, point his bill downwards and open it, the way that adult starlings discover insects hidden among grass. It was not until I saw the full adult behaviour that I realized what the clumsy transition phase was about.

4. Evolution

Selection, whether natural or artificial (i.e. by us) clearly shapes innate, genetically determined behaviour. Sometimes the same behavioural habits are developed as a result of different selection pressures. Foxes do not behave like rabbits, although both dig holes and sleep in them.

Social living has evolved in many different animal groups, but may have different benefits according to the animals’ way of life. Most carnivores are not social. Those that are usually hunt in coordinated packs, like wolves, orca or the little African wild dog Lycaon. Sometimes this enables them to take prey much bigger than they could alone. Many of the animals they prey on, like caribou, seals and zebra, also live in groups, but for wholly different reasons: for them, there is safety in numbers. Others group for other reasons. Red deer form groups in the mating season, with fierce competition between stags for control of the groups of does. Social living can be the answer to a variety of problems. We should not, then, assume that all animal societies are organized along the same lines. Social relations will vary according to the particular benefit deriving from group living.3 By studying the natural selection pressures that operate on the animal, its evolutionary development, we can see how the different patterns of social behaviour evolved to confer different benefits.

In the next chapter we will examine the horse’s evolutionary history to see what selection pressures drove the shape of the equine body and behaviour.

Particularly deep-rooted is an animal’s defence behaviour in the face of life-or-death situations, for there is no time for learning or fumbling about until a satisfactory solution arises. Whatever an animal’s chosen ploy in order to save its life – leap into water, climb a tree, shoot down a burrow or run away – it must follow instantaneously, automatically, when danger threatens. In turn its selected defence reaction shapes other innate behaviour. If water, tree or burrow are essential to the survival plan, avoid straying too far from them; if running away is the plan, better not sleep in enclosed places.

The innateness of patterns of defence behaviour does not mean that animals cannot learn to discriminate between different threatening stimuli and to habituate to some but not others. An animal’s learning ability, too, is shaped by its evolution: not only in terms of whether it can learn at all, but what it can learn readily. With their rich social life, horses are quick to distinguish and remember individuals (as well as plants) and to learn to respond to signals, even ones made or invented by us. They are not quick at learning to recognize triangles or do the Spanish walk. Evolution has shaped the mind so that some S→R connections are easily, almost invariably, made while others take a great deal of effort to connect, if at all. It has also shaped their cognitive processes: a major consideration in horses’ decision-making is safety, whereas a wolf’s is more likely to be affected by the chances of making a kill.

MOTIVATION

Often when we ask ‘why did the animal do that?’ we mean ‘what made it want to?’ or ‘what motivated it?’ This is a tricky question.

What we now call motivation was once called ‘drive’, which perhaps better describes the feeling we have when motivated, as if driven to do something. Enormous efforts were made to produce an overall theory of drive, but it became so convoluted, still without fitting all the facts, that it collapsed, though not before provoking years of argument, definitions and re-definitions. Like the word instinct after the collapse of Lorenz’s model, drive was replaced by a hitherto neutral word without history.

There is no general theory of motivation. Each case is taken on its own merits, though there are groupings, behaviours that seem to work in the same way. Consider the flow diagram, borrowed from systems analysis, which seems to fit many cases of maintenance behaviour well:

Or: an animal gets hungry (is motivated to eat). It goes out looking for food (appetitive behaviour). When it finds some (releasing stimulus), it eats (consummatory behaviour), and no longer feels hungry (negative feedback). If it is very hungry, it may accept food that it would not normally eat (stimulus generalization).

In this case, motivation increases with time. In some cases it may not. A mare comes into season and looks for a stallion, but if she does not find one she does not, as it were, get hungrier and hungrier. After a few days her motivation will disappear anyway, being under hormonal control. Motivation often is, even hunger.