10
Animal Agents in the Human Environment
Steven Ammerman
Abstract
Humans’ increasingly close relationship to animals constitutes one of the most important cultural, social, and economic developments of the past 10,000 years of our history. Animals have been and continue to be important sources of food, labor, and secondary products in many societies. As relationships with animals intensify, processes such as domestication ensure that humans are potentially able to control the behavior and deployment of large numbers of animals, altering ecosystems and creating an anthropogenic landscape. However, these types of relationships are heavily structured by the innate attributes of the animals involved: pre-evolved characteristics create the set of possibilities on which human agents can act, and actions undertaken by animals have major impacts on human as well as conspecific behavior. The agentive actions of wild and commensal animals are evident in trajectories of domestication and are further brought to the fore through the process of ferality, which is an agentive re-imagination of the domesticated niche and not always controllable by human actions.
All animals interact with other animals that share their environment, not only through predator/prey relationships but also through a variety of biological symbioses. One type of symbiosis is a mutualism in which individuals of both species benefit from the interaction, such as cleaner wrasse eating the parasites of larger fish: the larger fish benefits from a reduced parasite load, and the wrasse benefits by having an easily procured meal. Relationships in which individuals of one species benefit from the relationship while individuals of the other are neither helped nor harmed are known as commensalisms. Examples of this include cases where an animal benefits from the environment created by another, such as birds exploiting the disturbance caused by cattle to improve their success at hunting for insects (figure 10.1). A relationship in which individuals of one species benefit to the detriment of individuals of another is known as parasitism; this includes relationships in which members of one species feed off hosts of another species (such as ticks or lice feeding off mammalian hosts), but it can also encompass more complex interactions, as in the brood parasitism of a parasite that tricks or coerces the host into rearing its young. An amensal relationship is the opposite of a commensal one: members of one species are neither helped nor harmed by the interaction, while members of the other are harmed. An example of this could be the relationship between a swarm of locusts and a prairie dog colony. When the swarm of locusts denudes the vegetation from the colony’s location, the prairie dogs are left with nothing to eat (causing harm) but the locusts are indifferent to their presence or needs. A final type of symbiosis is neutralism, in which species share the same landscape but don’t directly benefit from one another’s presence, as in mixed herds of zebra and wildebeest.
Humans likewise have always maintained various types of relationships with the animals that shared their environment. Early hominins were poorly equipped to defend themselves from large predators, so a feature of this interaction would have been avoidance of becoming prey. Early hominids such as Australopithecus afarensis were frequently preyed upon by large cats, dogs, hyenas, bears, and even raptors. The relationship of humans as prey is not only represented in our early evolutionary history but continues today (albeit much less frequently; Hart and Sussman 2018). Our human ancestors also took advantage of changes to the environment created by other animals, such as scavenging the kills of other predators (Shipman 2010) and exploiting trails made by large animals (Haynes 2006). The development of tools by human ancestors was motivated at least in part by the need to process animal carcasses and to aid in the capture of animals for consumption (Shipman 2010). The importance of animals to early humans is shown by the preponderance of animal imagery in early figurative art (for more on the symbolic importance of animals, see Bishop, chapter 8, and Tomášková, chapter 11, both this volume). As human relationships with their environment became more complex, animals were relied on as sentries, as walking meat larders (Clutton-Brock 2012:35), and as living tools, as well as a source for power (Shipman 2010). While we tend to conceptualize these interactions as driven primarily by choices made by humans, the agency of animals also plays a large role in defining these relationships, particularly with reference to domestication as a clear symbiotic partnership.
Starting at least 11,000 years ago (Zeder 2012a), humans began experimenting with a strategy that involved closer relationships with animals, and some species of animals started to take advantage of closer relationships with humans. We now recognize this as the early stages of the process of domestication. Over many generations, animals that found reproductive success through these relationships manifested the evolutionary effects of this changed strategy via the physiological, behavioral, and genetic changes we associate with domestic animals (the so-called domestication syndrome; Larson and Fuller 2014; for the limitations of this concept, see Lord et al. 2020). Of the multitude of animal species co-occupying environments with humans, only a few went on to become domesticates. Is this small sample size the result of unwillingness on the part of humans to domesticate other species (lack of need) or the result of other species not having a suitable temperament for domestication?
In the process of domestication, the agency of animals to enter into a social contract with humans (giving subservience and docility in exchange for food and protection; figure 10.2) drastically altered survival strategies. For many domesticated species, this new survival strategy has resulted in much greater reproductive success compared to their wild progenitors. The domestic cow is one of the most plentiful large animals on earth, while its ancestor, the wild aurochs, is extinct (Felius et al. 2014). The same is true of the domestic horse; its wild ancestor, the tarpan, is extinct. The wild progenitors of sheep and goats, the Asiatic mouflon and Asiatic bezoar, are both listed as vulnerable by the International Union for Conservation of Nature (IUCN 2019).
Animal Agency
Agency relates to the ability of an organism to impact its environment by selecting among options for how to fulfill basic needs of food, shelter, and reproduction. The fields of cognitive psychology and philosophy provide a framework for thinking about animals, which can move and act autonomously, as agents. Shaun Gallagher’s (2000) attempts to reconcile philosophy and cognitive science lead him to divide the self into two levels, the more basic of which is a “minimal self.” The “minimal self” is “a consciousness of oneself as an immediate subject of experience . . . One does not have to know or be aware of this to have an experience that still counts as a self-experience” (15). The minimal self is the part of the self that understands that something is happening to you. The concept of the minimal self allows the ideas of a “sense of agency” and a “sense of ownership.” In this definition, agency is the sense of being the causative initiator of an action, whether a movement or a thought. Agency in this context is a causative phenomenon rather than a manifestation of volition, or as Stephanie Spengler and colleagues (2009:290) put it, “was it me or was it you?” Agency as a causative phenomenon is different from the sense of ownership, which refers to the sense of participation in an experience, this experience being voluntary or involuntary (sense of ownership applies when someone else moves my hand—I know it is my hand that is moving, but I did not cause the motion). In applying Gallagher’s concept to the “was it me or was it you” definition of agency, most animals are capable of meeting his definition of “minimal self” and being self-aware.
Being self-aware is one condition for animal agency, but it is not the only requirement. The philosopher Helen Steward (2009) developed a four-point definition of agency, in which three of the points are manifestations of Gallagher’s minimal self. She suggests that animal thought may not consist of circumscribed, causation-based units of ideation in the way human thought does, but this “human” way of thinking is not necessary for the animal to possess agency. Her four-point definition of agency as it specifically applies to non-human animals is:
- 1. An agent must be capable of independent movement. If something is unable to move, it cannot act upon other components of its environment.
- 2. An agent must be “a centre of some form of subjectivity” (Steward 2009:226) while being self-aware as an entity—aware of a difference between itself and its environment. Without self-awareness, there can be no purpose to any movement: it is either purely reactive or random.
- 3. An agent has at least some basic intentional state (e.g., in the shape of wanting, trying, awareness of its surroundings). In other words, the action must be toward some purpose.
- 4. An agent has some kind of directed control over the actions it takes to cause outcomes; the motion the agent undergoes to fulfill a need is the result of internal impulse, not external stimuli or triggers—in other words, “it was me.” While this criterion is similar to (1), which only requires the agent to be able to move independently, it also requires that movement to be (a) controlled by the agent and (b) aimed at achieving a purpose.
Importantly, this definition of agency allows room for instinct to influence the animal’s needs and desires. As long as the animal selects from a number of possible actions to fulfill these instinctual imperatives, it is acting as an agent. This definition excludes actions over which no control is exerted by the animal (i.e., reflex), but it does attribute agency to some of the actions taken by wild, commensal, domesticated, and feral animals.
Below, I make use of concepts that provide an explanation of the self that does not rely on a socially constructed understanding of selfhood or free will but focuses instead on individual action, in which the agency of animals leads them to alter the landscape where they live through a process known as niche construction (cf. Lewontin 1983; Quintus et al., chapter 9, this volume). Perhaps the most striking example is the dams beavers build to create a water impoundment that supplies them with food and protection from predators (figure 10.3), although all animals engage in niche construction to some extent. Whether it is the burrow of a gopher, the nest of a bird, or the wallow of a bison, the presence of animals has an effect on their surroundings. Some effects are due to intentional action, but in other cases the actions of animals have unintended and sometimes even detrimental impacts on their environment. When a herd of elephants forages through a forest region, it creates a new environment for grasses to grow and at least temporarily replace the woody plants of the understory. This new growth is not a preferred food for elephants, instead competing with elephants’ favored shrubberies.
Another manifestation of animals’ agency is seen in the defensive mechanisms they adopt to avoid becoming prey (whether and how to flee, hide, or fight). Tactics that may have worked well as defense against four-legged predators were not always successful when facing hominids that had a different hunting strategy and a different set of weapons that replaced tooth and claw. Whether behaviors to counteract these human threats were individually learned or the result of evolutionary pressure is unknowable, but the outcome is the same—individuals and species with better defense strategies fared better than those with insufficient ones. As humans intensified their hunting behaviors over the past million years or so, prey animals’ agency led them to new defensive techniques that had a feedback loop with human hunters. Animals’ attempts to evade human hunters led to changes in hominid hunting techniques, and the natural migration of animals (both large-scale and local) forced hominid hunters to follow.
Wild Animals
Regardless of whether humans are present, wild animals’ agency has an effect on an individual member of a species, on their group, on the other species with which they interact, and on their environment. An animal’s behavior has an impact on everything around it: when a herbivore eats, the nutrition it receives from that particular source has a primary effect on the individual; mate selection has an effect on the future of the species through the combination of genetics imparted to the offspring produced; a carnivore’s agentive selection of prey also has an effect on the prey species; and even something as simple as moving along a habitual pathway has an effect on the environment by creating a trail. To show animal agency in the natural environment, consider that any animal’s desire for self-preservation will motivate it to escape from its predators. However, for many smaller prey animals, escape can be accomplished in a variety of ways; and a selection is made among the options of hiding, fleeing, or fighting. In the predator/prey interaction between a coyote and a hare from the perspective of the hare: (1) the hare is obviously capable of motion; (2) the hare is aware that the coyote is a threat to its safety; (3) the hare’s desire to escape predation is to avoid being caught; and (4) the hare has control over which actions to take to effect that escape. The hare is therefore exhibiting the four hallmarks of agency outlined by Steward (2009). The action the hare will take depends on its awareness of the environment. If it is close to a haven (such as a pile of boulders), it will likely choose to run to that haven. If it is in an area with sufficient cover, it may remain motionless in an attempt to keep the coyote from detecting it. If fleeing or hiding are not options or are not successful, the hare can forcefully defend itself in an attempt to convince the coyote that other prey may be less trouble.
Within a population of animals, some individuals will be more prone to one strategy than another (for avoiding predation, finding food, or exploring their environment) as a function of the individual animal’s temperament. Field observations of wild animals have shown that the personality traits that can be observed range from passivity to curiosity, skittishness to boldness, and submissiveness to aggression. Acting based on these personality traits can be beneficial or detrimental, depending on the circumstance, in ways that can impact the reproductive fitness of wild animals. For example, Phylis C. Lee and Cynthia J. Moss (2012) suggest that differences in temperament are a deciding factor in social leadership among elephants. Peter A. Biro and Judy A. Stamps (2008) have shown that behavioral tendencies or personality traits such as boldness and aggression have a positive correlation with life history successes (including increased growth rate, increased adult size, and fecundity) in a wide variety of animal species, from insects to mammals. Because some elements of temperament are heritable (van Oers and Sinn 2013), when one temperament leads to enhanced fitness, the population will be skewed toward that temperament (McDougall et al. 2006). Within captive populations, tests have been developed to measure various aspects of temperament and demonstrate the variability that exists among individuals. Russell Greenberg and Claudia Mettke-Hoffman (2001) discuss methods for assessing neophobia in birds, such as placing a new object adjacent to a food source and measuring the time it takes for the birds to come to the food. A study on the behavior of captive spotted hyenas showed the reliability of five general personality traits (assertiveness, excitability, human-directed agreeableness, sociability, and curiosity) in quantifying the differences in behavior among individuals (Gosling 1998).
Animals in a Commensal Relationship with Humans
Animals in a commensal relationship with humans are those that take advantage of human-modified environments and live in close proximity to humans. Although humans are not necessary for animals to exhibit agency, their presence provides new opportunities for animals that are willing and able to exploit them. The relationship between commensal animals and humans may be one of true commensalism, mutualism, or parasitism. Evidence of animals exploiting new niches created by increased human sedentism dates back at least to the time of the Natufians (~15,000 BP), one of the earliest known sedentary populations. Here, the common mouse (Mus musculus) was able to create its own niche within this environment and out-compete the Macedonian mouse (Mus macedonicus), another small rodent native to the Near East that was less adept at exploiting the new conditions created by human niche construction (Weissbrod et al. 2017). The relationship continues today, and the common mouse has exploited the relationship so well that it is now a globally distributed species that lives everywhere humans do (even in outer space; Neff 2017).
As human populations have grown and their niches have diversified, so too have the number of commensal species that take advantage of human niche construction. Cities, as heavily constructed environments with high concentrations of people and a wide variety of microhabitats to be exploited by commensal animals, represent the ultimate arena for human-commensal animal interactions; animals that are successful in urban environments must be well adapted to tolerating the presence of humans. However, even in non-urban environments, a commensal strategy that makes use of human-created niches is adaptive for a wide variety of species. A good example can be found in modern exurbs with relatively low housing density: in these regions, there is ample space for animals as well as a plentiful supply of food in the form of enhanced foliage, garbage, food placed out for domestic animals, and purposeful feeding of wildlife (e.g., birdfeeders). In North America, white-tailed deer, raccoons, and opossums (among others) live in proximity to humans and take advantage of these resources. The commensal coyote takes this niche adaptation one step further by directly consuming some human-provided resources and by preying on the other species drawn into the niche—all the while benefiting from the human removal of the large predators such as wolves, mountain lions, and bears that previously provided competition.
Commensal animals are frequently so successful in human environments that their increased population sizes lead them to become nuisances. One example is the rhesus macaque in India, which, according to Charles H. Southwick and M. Farooq Siddiqi (1994:223), “is probably the world’s best example of primate commensalism.” Commensal rhesus macaques flourish in disturbed habitats and in close contact with humans in locations such as cities and villages (including temples and railway stations) and in agricultural zones, adeptly taking advantage of human niche construction. These commensal macaques depend on humans for food provision by raiding crops, causing significant economic damage (Rattan 2011, cited in Southwick and Siddiqi 2011; also personal observation, Talapada, India). This problem is “highly seasonal and location specific” (283), as it is related to the abundance or lack of foods depending on seasonal agricultural cycles. In urban areas, macaques often harm property in their pursuit of food (Saraswat et al. 2015) and have been known to resort to direct contact and antagonistic behavior, such as biting people or snatching mobile phones, glasses, or bags and returning the objects only when they are threatened or offered food (Chauhan and Pirta 2010). Moreover, temple visitors and tourists voluntarily feed the macaques, exacerbating the animals’ reliance on human-derived foodways (Rattan 2011, cited in Southwick and Siddiqi 2011).
Domesticated Animals
Domestication is a process involving groups and occurring over multiple generations. The actions of a single animal (or a single human) will not lead to a domesticated species; rather, the collective action of groups of animals and people brings about this result through interactions over a prolonged period of time. Melinda A. Zeder (2012b) describes three pathways to domestication: commensal, prey, and directed. Each of these different pathways can be understood in terms of the agency animals exerted in their development.
Commensal Domesticates
Commensal domesticates are animals that became domesticated through a process of habituation, such as the dog, cat, pig, Guinea pig, golden hamster, chicken, duck, and turkey. These species were able to successfully exploit the niches created by human activity and became associated with humans in the initial stages of their domestication. In the commensal pathway to domestication, animal agency is obvious: the animals chose to be in close contact with humans and chose to remain with the humans; no enclosures or capture devices were used. There were, however, at most a few locations where this kind of domestication took place, compared to all the possible locations in which the wild progenitors of commensal domesticates were in the vicinity of humans. Was there something unique about the temperament of the individual animals within the particular groups that continued down the commensal pathway to domestication, or was there something unique about the group of people that took advantage of the animals’ agentive choice to stay nearby?
The story of pig domestication provides a distinct example of mutualistic commensal accommodation. Wild boars were widely distributed throughout Eurasia and northern Africa during the Mesolithic and early Neolithic periods. The process through which these wild boars became the domestic pig took place over a long period of time and probably in more than one location. Early evidence of domestication comes from eastern Anatolia, where pig remains from the sites of Hallan Çemi and Çayönü Tepesi show signs of domestication (e.g., morphological changes and changes in kill-off patterns) from ~10,000 BP (Hallan Çemi) and ~9,000 BP (Çayönü Tepesi) (Rosenberg et al. 1998; Ervynck et al. 2001). Studies of ancient mitochondrial DNA by Greger Larson and colleagues (2007) suggest that early domestic pigs came from the Near East, but other domestication events could have taken place in Europe and did take place in East Asia. Analyses of 2,000 years of pig remains at Çayönü Tepesi suggest a long process of domestication (Ervynck et al. 2001), and analyses of pig remains from a variety of sites in southern Anatolia and the northern Fertile Crescent support a long-term process of morphological change following the initial period of domestication (Price and Evin 2019).
Wild boar are omnivorous; therefore, this process could have begun with boar exhibiting agency and coming to early sedentary habitation sites to feed on human refuse or early agricultural products—a truly commensal relationship with humans. The close proximity to humans and the change in diet could have led to the initial stages of domestication, both behaviorally and morphologically; but for the relationship to progress to full domestication, there had to have been a benefit to the humans from having the boar in close proximity (a mutualistic symbiosis). The exact details of early pig husbandry are not clear from the archaeological record, but a modern analog in New Guinea may provide one possible process. In this case, domestic females are kept but no breeding males; instead, the females mate with wild males. The piglets are born in the village and are coddled within households, so they imprint on their human owners. All of the male piglets are castrated, so humans only control the female half of the breeding pair. When the piglets are 4–5 months old. they are allowed to forage throughout the village for food (Rowley-Conwy et al. 2012). There are no physical barriers keeping the pigs in the village, and only their own agentive actions—taken because of the benefits they obtain from close interaction with people—keep them where they are beneficial to humans.
Modern zooarchaeological techniques have been applied to faunal collections in an effort to clarify the transition from commensalism to domestication in ancient pig populations. For example, examination of stable isotope ratios in purported early animal domesticates can provide insights into dietary differences from their wild progenitors. Thomas Cucchi and colleagues (2016) used this method to determine that early domestic pigs in China were foddered on cultivated millet grain, which was absent from the diet of wild boar. The ready supply of the millet may have provided the incentive for the pigs to remain in association with humans in the earliest part of the domestication record, similar to New Guinean practices in more recent times.
Prey Domesticates
Human practices such as selective hunting and game management constituted a process toward domestication for cattle, sheep, and goats in the Old World and for llamas and alpacas in the New World. As an example, bezoars (wild goats) were widely distributed throughout the Near East during the Mesolithic and early Neolithic periods (Makarewicz and Tuross 2012). Demographic profiling shows selective harvesting of wild goats in southeastern Turkey and northwestern Iraq, which may have begun as early as 12,000 to 13,000 BP (Zeder 2012b) with the selective harvest of large adult males. This game management demographic pattern is different from that shown in early domestic herd management, which favors the killing of young sub-adult males, as documented in the Zagros Mountains of Iran at about 10,000 BP (Zeder and Hesse 2000). Mitochondrial DNA analyses of goats and wild bezoars indicate that the domestication process probably consisted of at least two independent events, one in eastern Anatolia and the other in the southern Zagros Mountains/Central Iranian Plateau. Most modern domestic goats are descendants of those domesticated in eastern Anatolia (Naderi et al. 2008).
Using stable isotope analysis, Cheryl Makarewicz and Noreen Tuross (2012) showed an intermediate step between game management hunting strategies and herd management domestic strategies in the Southern Levant about 10,000 BCE. In this scenario, humans selectively foddered goats and influenced their pasture sites to promote the health of goat populations as a beginning of a mutualistic symbiotic relationship. The selective foddering was shown by differences in the isotopic mixture of the goats’ diet compared with that of gazelles, whereas in the wild these two species would have had diets with similar isotopic mixtures. The influence on pasture location was evidenced by differences in hydrological regimes used by manipulated goats versus unmanipulated gazelles. Whether the goats studied were fully domestic or at an intermediate stage in the domestication process is not clear, but what is clear is that humans were attempting to make use of the agency of goats by providing them with an alternative to wild foods and directing their grazing locations.
A similar process of transitioning between wild herd management and domestication was used for reindeer, a process that can still be seen in the management style of Sami herders in the Kola Peninsula of northwest Russia. Herders only occasionally round up strays and move herds to fresh pastures, but at certain times of year tighter control is exercised, such as during the calving season (Baskin 2000). Within this loose-control strategy, there is ample opportunity for interbreeding between wild and domestic populations of reindeer. Knut H. Røed and colleagues (2008) determined that in some locations, there is little difference between the gene pools of wild and domestic herds. Part of the reason for this is likely because reindeer are a very recent domesticate (probably the most recent prey domesticate), having only entered into a mutualistic relationship with humans 2,000 to 3,000 years ago (Zeder 2012b). Unlike other prey domesticates, the wild population of reindeer is still extensive (see Tomášková, chapter 11, this volume).
While the directed action of animals plays less of an obvious role in the prey pathway to domestication than in the commensal pathway, animal agency still contributes to the way the domestication process unfolds. The agency displayed by animals and the differences in temperament and behavior between individuals and among species influenced which of the many prey species exploited by human hunters successfully made the transition into fully domestic animals.
Directed Domesticates
Once humans had domesticated animals such as goats, resulting in a template for how to obtain benefit from animals without having to hunt or trap them, other animals became domesticated. Directed domesticates include the horse, donkey, Old World camelids, buffalo (American bison), ferret, mink, silver fox, chinchilla, emu, ostrich, and fish. These species were domesticated not only for food but also for other benefits they could supply, such as transportation or tractive power (horses, donkeys, dromedaries, yaks), furs (mink, chinchillas), and fibers (silkworm) (Larson and Fuller 2014). The decision to domesticate animals that were not entirely intended for use as food stemmed from the utilization of domesticates for these purposes in the early stages of the secondary products revolution (Sherratt 1981). For example, the use of cattle (originally a prey domesticate) for new activities such as traction or milk production could have been the impetus for the domestication of the dromedary and the yak, as these species thrive in arid and high-altitude environments unsuited to cattle.
Implications of Domestication: An Animal Perspective
Considering all the wild animals that could provide benefit and the relatively small number of species that have been domesticated over time, the choice of domesticates was certainly not entirely up to the human domesticators. Further, domestication does not diminish animal agency; anybody who has a pet dog or cat realizes the degree of control they exert over their owners and the ways they rely on a strategy of human interaction as part of their problem-solving strategies that also have measurable physical consequences (e.g., Kaminski et al. 2019). Bradley Smith and Carla A. Litchfield (2010) report on experiments comparing dingo and dog problem solving in which the two groups have a very different approach to solving the problem placed in front of them. In the experiment, a V-shaped fence was constructed; the subject was placed either inside or outside the V, and food was placed on the other side. To get to the food, the subject had to go around the end of the fence. While dingoes would tend to go around the fence to retrieve the food prize, domestic dogs attempted to look to the experimenters for guidance rather than independently completing the task. While this may appear to suggest that dingoes are quantitatively more intelligent than fully domestic dogs, I would suggest instead that this demonstrates a response evolved by domestic dogs to choose collaboration with humans for the purpose of problem solving. Notably, both the dog and the dingo performed actions in response to a challenge, demonstrating that both retain agency. Every individual possesses a suite of possible reactions to a problem, from which a response is selected. In other words, the domestic dog exercises its agency to select an approach that relies on its evolved symbiosis with humans.
In general, domestic animals have developed a great deal of reliance on humans to fulfill their survival needs, and if humans had failed to supply those needs during the domestication process, the pathway would have hit a dead end. Even in cases where the domestication pathway is long and well established, however, there are instances where an animal may elect or be forced to reduce its reliance on human interaction, an agentive process that leads to feralization.
Feral Animals—a Different Kind of Agency?
Feral populations arise from the reproduction of domestic animals that have abandoned a human-based survival strategy (animals that are in the first generation to leave human control are strays, while their descendants are feral; cf. Slater 2001). While many feral populations continue to live in commensal relationships with humans, they no longer have an intimate social connection with them. Feralization, therefore, is an evolutionary trajectory that comes as a direct result of an application of agency by domestic animals: feralization is an additional stage of the process of evolution, not an undoing of domestication.
Domestic pigs, camels, and goats are all good examples of species that have sometimes successfully abandoned their close ties with humans and developed large feral populations, mostly in regions where the domestic population’s wild precursors are absent and to which a domestic population has been introduced. An excellent example of this is the feral dromedaries of the Australian Outback. Dromedaries were initially domesticated in Arabia around 3,000 years ago (Almathen et al. 2016) and were introduced into Australia in the mid-nineteenth century for use as draft animals. The climate of the interior of Australia is well suited to camels, and approximately 20,000 of them were imported (Saalfeld and Edwards 2008). By the mid-twentieth century, motorized vehicles had made the use of camels obsolete, and many camels were released into the wild. From this initial stock of probably fewer than 10,000 individuals, the population of feral dromedaries in Australia had expanded to nearly 1 million animals by the early twenty-first century (Saalfeld and Edwards 2008). Clearly, the domestication process did not impact the dromedaries’ ability to survive without human intervention. The success of the feral camels has established them as a pest species over much of their range, where they are outcompeting native animals for food and water.
The survival strategy to abandon close contact with humans may not be the preferred option for domestic animals, but it is one they are not always evolutionarily excluded from using. If this abandonment is not a successful strategy, the population of strays will not reproduce and rear young with enough success to become a self-sustaining feral population (this failure to create a self-sustaining feral population is frequent, for example, in populations of stray and feral dogs; Zeder 2012b), so the fact that there are sustained feral populations of some species indicates at least a moderate level of success and a forward movement along the evolutionary pathway.
Conclusion
Acknowledging the importance of animal agency reorients our understanding of human cultural strategies as part of broader ecological and environmental systems. Animal agency has had an influence on many, if not all, human-animal interactions and therefore on the development of all the systems in modern human society that are reliant on those interactions. The ramifications of this agency become even clearer when exhibited by the domestic animals that share the most intimate relationship with humans. The agency exerted by domestic animals in any given situation depends not only on the human-created environment but also on the temperament of the individual animals (and eventually the predominant temperament selected for the population by the domestication process) when exposed or introduced to that environment. Some species of animals have ranges of temperament that better position them to enter into relationships with humans, and for the most part, these are the species that have continued down the various domestication pathways to the present.
The domestic animals we have today are as much a result of the agency of animals as they are of human action. While many animals could be useful as domesticates, the number of domesticated species is relatively small. The animals that have entered into a social contract with humans through the long-term evolutionary process of domestication have certainly had meaningful impacts on every facet of human life: from the nutritional impacts of having readily available sources of meat to the economic advantages of harnessing the strength of animals to providing us with clothing materials (leather, wool, silk, felt, furs) and providing companionship. While domesticated animals have thus contributed much to economic and physical health through their engagement with human-created ecological niches, they have also contributed to negative impacts when serving as vectors for zoonotic disease transmission (see Juengst et al., chapter 6, this volume).
An interrogation of the actual ramifications of these relationships reveals a much more complex ecological tapestry in which animals contribute to and shape the direction of human niche construction and human culture. Perhaps the term symbiotic more accurately reflects the nature of these relationships, suggesting a broad spectrum of possible relationships between humans and animals that for either party can be harmful, beneficial, or neutral. Aspects of the symbiosis between humans and animals can vary over time and space, even for a single individual. For animals that live in environments heavily shaped by human activity, some aspects of human niche construction may provide benefits, such as a reduction of predators or greater access to food, while other aspects simultaneously introduce new dangers, such as automobiles, pollution, and slaughter. If the net benefit of these relationships to humans and animals outweighs the harm, these relationships persist or even intensify. If the net harm to both humans and animals is greater than the benefit, these relationships diminish or end. The overall effect is generally perceived at the population level, but the benefit or harm plays out on an individual basis where agency influences the reactions taken. For individuals, behaviors vary temporally on a daily basis (night versus day), on a seasonal basis (migration or crop availability), and on the scale of a lifetime (long-term resource depletion or accretion). These same dynamics play out for domestic animals, which for certain parts of the day, year, or lifetime may have varying degrees of interaction with humans and may play a greater or lesser part in human lives. Because of this, one animal may exist in a variety of niches and experience a variety of different types of symbiosis with humans over the course of its life. The agency of individual animals is a key factor in establishing these changing relationships.
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