We think of other primates as our closest relatives, and in evolutionary terms they are. However, dogs mirror our social patterns in ways no primate can. We recognize in them elements of ourselves, and this permits a certain level of empathy. Selection for paedomorphism such as small size, outsized eyes, and soft hair has produced dogs having permanent juvenile features, making them reminiscent of pups in arrested development (Fig. 2.1). Like us, dogs are social with senses attuned to voices, scents, postures, and dependent on familiarity and friendship. A dog growls and we know what it signifies. We shout and the dog understands, if not the words. In behavioral terms, “the primary stimuli are so similar in the two species that appropriate and recognizable social behavior is evoked.”

Domestication

In structure and form the dog is the most variable mammal of all, a situation attributable entirely to humans. Genetic mutations are probably too slow to account for the typically rapid alterations in phenotype. A report on 414 purebred dogs of 85 breeds showed that interbreed differences accounted for 27% of the total genetic variation. Western European breed clubs with their attendant standards date from the mid-1800s, and today more than 400 breeds exist. Among the standards adopted was the “breed barrier” rule, which decrees that for a dog to be registered both its parents and grandparents must be registered members of the same breed. The result was to constrict the borders of gene pools through inbreeding, making the physical appearance of each breed reproducible. Inbreeding can sometimes have negative effects including reduced incidence of conception, fewer live births, lowered sperm count, and smaller testis volume.

Members of the genus Canis have reproductive cycles with several unusual features: monogamy, parental care, monestrus, extended proestrus and diestrus stages, a copulatory lock, incorporation of offspring into the social group, social suppression that keeps subordinates from breeding, obligate pseudopregnancy in subordinate females, and helping behavior by group members including males. I define and discuss these concepts here and in the next chapter, demonstrating how their intricate patterns of timing affect societies of wolves and free-ranging dogs.

Aspects of the reproductive cycles themselves differ in degree but not kind. Most changes are endocrinological and therefore invisible, yet they profoundly affect social behaviors and interactions. To ignore their details in favor of strictly observational information puts aside the biological basis of reproductive behavior, revealing only a partial glimpse of its integrated functions. In addition to the expected differences between wolves and dogs are large individual variations in endocrinological profiles throughout the reproductive cycle, and these are routinely evident even in littermates. I start with an explanation of how the endocrinological factors cycle, stitching in brief mention of the external displays we recognize as courtship. Aspects of courtship in owned dogs, wolves, and free-ranging dogs are then discussed specifically.

The zoological order Carnivora includes the canids. When discussing its members the term carnivoran is preferable to carnivore because it excludes unrelated predators. Modern canids appeared about 10 million years before the present (years BP) and diverged into two branches, the dogs and the foxes. Depending on how you divide them, living canids number about 35 species. They have 78 chromosomes, and all are known to admix. The golden jackal (Canis aureus) and gray wolf (Canis lupus) have been considered the domestic dog's possible ancestors, and although all evidence points to the gray wolf, some raise other possibilities, such as an extinct and unknown wolf-like canid. Two of these are the dingo and a hypothetical and now extinct wild dog similar to the dingo. The next closest relatives of gray wolves and domestic dogs are the coyote and Ethiopian wolf (Canis simensis), less accurately called the Simien jackal. As later chapters should help clarify, implications of these relationships reach out from the past, affecting the behavior and social lives of all members of the genus Canis, including the domestic dogs we keep as pets.

Wolves in the beginning

The family Canidae (Canis means dog in Latin) evolved in North America, first appearing in the late Miocene 6 million years BP. When North America and Asia formed a high-latitude connection in the late Cenozoic (3 million years BP) some canids migrated across, where they continued to evolve, and one returned later as the gray wolf. The record infers that North American wolves and the coyote separated about 1–2 million years BP, although genetic findings point to the gray wolf's origin being only 250 000 years BP. According to a slightly different hypothesis, the gray wolf might have evolved in Asia and migrated to North America about 300 000 years BP across the Bering land bridge when sea levels were lower than today.

Our fascination with minutiae is associated with the search for distinctions.Nowhere is this better exemplified than by the recently discovered genetics ofscent reception. Scent-related genes make up the largest family in the mammaliangenome. Visual and auditory stimuli might seem complex, but future research islikely to reveal them as abbreviated and clumsy compared with the subtleemission and reception of odors.

The canid nose is finely tuned to scent reception. However logical it seems thatair-borne compounds in the glandular secretions, urine, and feces of wolves anddogs contain important (and presumably) intraspecific signals, the currentevidence is overwhelmingly observational. Simply watching how a dog behavesafter sniffing urine or feces, for example, does not establish a causal link.Conclusions based solely on behavior are inferential until validatedphysiologically. Between signal and response must be evidence of specificchemical compounds produced by senders that subsequently bind to identifiedreceptors in neurons of receivers.

Odor and pheromone reception

Reception at a molecular level involves olfaction (the detectionof odorant molecules), possible detection of pheromones, and taste (gustatoryreception). Odorants taken in from the external environmentconsist mainly of volatile compounds. Pheromones, which areexcreted into the environment in fluids like urine or retained on the emitter’sbody (e.g. in sweat or saliva), are discrete compounds or blends of compoundsserving as chemical signals among conspecifics to elicit sexual and socialchanges in behavior and physiology. Pheromones are mainly nonvolatile andrequire direct contact with sensory cells to stimulate detection. In otherwords, pheromones are both excreted and taken up as fluids, notaerosols. Their uptake by rodents after arousal is mediated bypumping action of the vomeronasal organ (see below). Because each sensory neuronexpresses just one allele on a single receptor gene, the connection betweenemission and reception becomes fused only when the appropriate molecules bind totheir specific receptors. The result is a signal sent to the central nervoussystem culminating in a sensation of smell (odorants) or some behavioral orphysiological change (pheromones). Olfaction and taste are not easily separable,odorants and pheromones even less so.

Humans name everything, living and dead. We name our pets and the wild animals wecome to recognize while conducting field research. In doing so we bestow on thema human-centered individuality, an epibolic trap from which they can neverescape. Animals, of course, are individuals without us. In the following pages Ihope to shed light on the factors necessary for socialization,the behavioral and ontogenetic progression through which young wolves and dogsbecome integrated into societies of their own kind – or alternatively, tohuman societies. Its rudiments appear early and are reinforced throughoutlife.

Wolves and African wild dogs are the most social of the canids by living in truesocieties. I define a canid society as a group of mostlyrelated conspecifics that occupies a common territory, communicates in complexways, and shares responsibilities, as in a pack. Domestic dogs interact moreloosely, and their groupings do not constitute true societies as just defined.The objective here is to describe the socialization of wolves and offree-ranging dogs. The wolf section includes observations from both captive andwild animals, and the findings do not always apply mutually.

Nearly all domestic animals are social species that form bonds with conspecifics,a quality that makes them more manageable, or tame. Tameness has been inter-preted as “an ontogenetic phenomenon facilitated through artificialselection.” This might explain how it came to be, but not what it is.Tameness, defined earlier as the absence of conflict behavior (Chapter 2), canalso be thought of as the absence of fear, and a tame dog or wolf is one thatdoes not fear humans. Feralization is then equated with the absence or loss oftameness and its replacement by fear.

Philopatry is attachment to place. Wolves and other dogs occupy finite spaces called home ranges and often territories as well to which they are philopatric. Both home range and territory are conceptually straightforward, but often used loosely and sometimes interchangeably. An animal's home range is “that area traversed by the individual in its normal activities of food gathering, mating, and caring for young.” The boundaries of a home range are often vaguely defined and flexible. The caribou (Rangifer tarandus), a circumpolar deer, might migrate 1300 km over the course of a year, the entire distance comprising the home range. Droughts, floods, population fluctuations, human encroachment, intensive predation, excessive self-exploitation of resources, and other factors cause animals to move and establish home ranges elsewhere. Young animals of many species leave the natal home range to establish their own. In practical terms, a home range is where an animal spends its life; that is, where it shelters, mates, rears young, interacts with others, eats and drinks, and eventually dies.

A territory is the part of a home range that an animal defends, ordinarily against conspecifics but sometimes against other species too. Note that the definition of home range says nothing about it being defended. In birds, a territory is often just the nest and immediate surrounding area, although the home range might be much larger.

Behavior, which is a phenotypic trait, can be defined as everything an animal does, but such a bland statement has no functional value. To make the term relevant I shall paraphrase a definition used by John Paul Scott and Emil Fredericson: behavior is the attempt by an organism to adjust to changing conditions. Note that behavior is not a response to an existing condition, but to a change in conditions. Communication, a form of behavior, is an association between the sender's signal and the receiver's behavior as a consequence of the signal. It can therefore be defined as the process of influencing others. I define signal as a specific stimulus used in communication; alternatively, a semiotic sign (see below). The receiver is the individual receiving a signal, the sender (or signaler) the one transmitting it.

Metaphors and semiotics

A term parasitized by many definitions turns stale and ultimately worthless with metaphorical use, becoming a “dead metaphor.” In trying to compare current usage with the original definition, we generally find language and culture to have changed and the relationship no longer relevant. Manufacturing metaphors for scientific purposes is not dishonest, merely hopeless. The conviction that they capture some essential element of nature is “no more important in the interpretation of metaphorical claims than . . . in the interpretation of literal claims.” Their application in scientific description devolves ineluctably into discussions about whose metaphor is the most literal and thus closer to real explanations of nature. Unfortunately, metaphors that are only partly metaphorical are as rare as dogs that are only part mutt.

By now it should be clear that a pack or group of social canids is a collective stomach in search of food. Wolves and free-ranging dogs feed by hunting and scavenging. As defined here, hunting is the act of finding, pursuing, catching, and killing prey, scavenging the act of feeding on carcasses or separating edible components from human refuse and other mostly inedible materials. Both terms are covered by foraging.

Foraging has the obvious benefit of supplying nutrients from which animals gain energy required for growth, reproduction, respiration, generation of body heat, and all other physiological processes driven and controlled by what we call “life.” However, it also has a cost, measured as the energy used to secure food. African wild dogs often chase prey for several kilometers at sustained speeds of 48 kilometers per hour, a tremendous energy expenditure culminating in a burst of strength and effort needed to bring a large ungulate to the ground. Tundra wolves commonly leave the den or rendezvous site (Chapter 8), travel 25–30 km, make a kill, and return with large portions of meat to feed pups and the breeding female all in a matter of hours. Not every hunting trip is successful. If more fail than succeed – that is, if cost exceeds benefit over time – the eventual end is starvation and death. Although wolves can be opportunistic foragers they are primarily hunters of large herbivores. Energetic constraints force them to be. A body mass of 21.5 kg is the point at which carnivorans generally shift from small to large prey, and wolves cross that line. The transition is abrupt: carnivorans smaller than 21.5 kg feed mainly on prey <45% of their own mass, those larger than this select animals >45% of their own mass.

Preface

Behavior and ecology are “two sides of the same coin,” a viewpoint I endorse and use in this brief essay on the social lives of wolves and free-ranging dogs. My goal has been to write a timely, simple, relevant book. Timely with current research findings, simple in minimizing technical information, and relevant by deepening our understanding of wolves and of dogs that run loose in a contentious world of shrinking spaces.

The wolves discussed include gray wolves (Canis lupus) and their admixtures: the Great Lakes wolf (including the Algonquin wolf) and the red wolf, more aptly named red coyote. When referring to first-generation wolf × domestic dog admixtures I use wolfdog; coydog is the term applied when the first-generation offspring are progeny of coyotes (Canis latrans) and domestic dogs. A free-ranging dog is any dog of domestic origin uncontrolled by humans and includes truly feral animals like dingoes, rural and urban strays, and wandering pets. Dingoes that have interbred with domestic dogs are known as wild canids or wild dogs in Australia, but to keep things consistent I refer to their first-generation offspring as dingo-dogs. Although not entirely logical (both are actually domestic dogs), the term has a pleasing sound when spoken aloud.

Spontaneous maternal care is unusual in mammals. The behavior we think of as nurturing requires a flood of priming hormones during pregnancy, compounds produced or regulated by the placenta. “Hence, the onset, maintenance and termination of maternal behaviour are controlled by hormones which, in turn, are released in response to stimuli from the foetus.” Being helpless, altricial young (Chapter 5) require lots of attention. The hormones of pregnancy (see below) induce denning, develop the mammary glands, and ready the brain for maternal care, which to continue through weaning and beyond requires both a fetus and later a neonate. Intensity of maternal behavior becomes associated inversely with development of the young: the mother leaves her pups longer once they can thermoregulate consistently and become less dependent on her warmth; her milk production diminishes concomitantly with decreased nursing.

The reproductive cycle – part 2

The discussion in this section is limited to the dog unless otherwise stated. Diestrus comprises the stage in which corpora lutea are completely functional; that is, the luteal phase of the female reproductive cycle. It follows estrus and extends 55–75 days. Behavioral diestrus is the period starting at the end of estrus when the female refuses to let the male mount. Her restlessness subsides along with the vaginal discharge and swelling of the vulva. External signs as diestrus progresses are those of pregnancy or pseudopregnancy (see below). About 20 days after initiation of diestrus the corpora lutea start breaking down (luteolysis), and the endometrial lining of the uterus begins to regress; later in pregnant females and some that are pseudopregnant, the mammary glands mature and milk-secreting tissues develop.