9Pathways to Animal Domestication
Melinda A. Zeder
Jack Harlan was a polymath. His life-long study of crop evolution combined plant
sciences, archaeology, systematics, genetics, and conservation, leaving a legacy of
ﬁve decades of inﬂuential publications that explored all aspects of crop plants –
their origins, their dispersal, and their continued and future role in supporting the
Earth’s burgeoning populations. To Harlan, agriculture was not an invention or
the product of a single big idea. Instead, he saw agricultural origins in terms of a
long co-evolutionary process involving humans and plants that grew out of “many
independent tentatives in many locations that fused over time to produce effective
food production systems” (Harlan 1995). Harlan’s remarkable body of published
work contains only one short encyclopedia entry on the subject of animal domesti-
cation (Harlan 1994). He was, in fact, somewhat dismissive of the contribution of
animal domesticates to humankind’s food supply stating that “(a)nimals are not
essential, plants supply over 90% of the food consumed by humans” (Harlan
1995). Jack Harlan would likely agree, however, that understanding livestock
evolution requires the same breadth of focus that he brought to the study of crop
evolution. Here I follow Harlan’s example in a consideration of domestic animals,
bringing together information from animal sciences, genetics, and archaeology to
explore the multiple pathways leading to animal domestication and the implica-
tions of these pathways for current and future relationships between humans and
their animal partners.
1 Domestication as a process
All considerations of domestication, whether focusing on crops or livestock,
acknowledge that it involves a two-way relationship between humans and target
plant or animal populations. There is less unanimity in different conceptual
approaches to domestication on whether emphasis should be placed on the
human or the plant/animal side of the equation (see Zeder 2006a). Some cast
humans as the dominant partner in a relationship in which humans consciously
Biodiversity in Agriculture: Domestication, Evolution, and Sustainability,editedbyP.Gepts,T.R.Famula,
R.L. Bettinger et al. Published by Cambridge University Press. #Cambridge University Press 2012.
and with deliberate intent assume “mastery” over all aspects of the production,
movement, feeding, and protection of the domesticate (Hale 1969, Ducos 1978,
¨nyi 1989, Clutton-Brock 1994). Others see domestication as a form of
biological mutualism in which both partners (humans and domesticate) reap
beneﬁts (O’Connor 1997). Some even contend that domesticates manipulated
unwitting humans into relationships that gave the domesticate great evolutionary
advantage at the expense of human ﬁtness (Rindos 1984, Budiansky 1992, Morey
I take a more centrist position. I acknowledge that both partners in the
domestic relationship reap beneﬁts through their increasing reliance on each
other. In this way domestication is indeed quite similar to mutualistic relation-
ships in the natural world like those between farmer ants and “domesticated”
fungi or between other ant species and their aphid “herds”, the most commonly
cited analogies with human plant and animal domestication (Rindos 1984).
But the mutualism that lies at the heart of the domestication process differs
from these convergent forms of biological mutualism in one important and
uniquely human way (Schultz et al. 2005, Zeder 2006a, 2009). Mutualistic
relationships in nature are the product of extended evolutionary processes
driven by selection operating on mutation-induced variation in behavior and
morphology in both partners, with adaptive changes in behavior and body form
passed on to future generations through genetic transmission in the course of
sexual reproduction. The co-evolutionary relationships between humans and
target domesticates, on the other hand, are largely driven by the human ability
to spontaneously invent new behaviors that maximize the return of a desired
plant or animal resource and, most importantly, to pass on behaviors that best
meet these goals to their offspring and to others through social learning. This
capacity for the cultural transmission of learned behavior ramps up the mutu-
alism between humans and emerging domesticates and transforms it beyond
anything seen in nature. Both partners still derive mutual beneﬁt, plants and
animal partners vastly enhance their reproductive ﬁtness and humans gain a
predictable and secure resource base. But the human capacity for social learn-
ing puts humans in a dominant role in an increasingly asymmetrical mutualism
that moves at a vastly accelerated pace and carries a much broader impact than
any such relationship in nature.
The process of domestication unfolds across multiple axes on both the plant/
animal and human sides of the equation (Figure 9.1). A primary axis on the plant/
animal side involves the phenotypic expression of genetic changes that transforms
the plant or animal from its wild phenotype to its domestic phenotype. Progress
along this axis is driven by a number of selective and random processes that
operate either sequentially or in tandem depending on the domesticate and the
nature of its relationship to its human partners (see Price 1984, 1999, 2002).
Human-orchestrated directed, or artiﬁcial, selection for desired traits is only one
of the forces that shape a domesticate’s trajectory along this axis. Other forces
may play an even more important role in shaping the domestic phenotype,
228 Melinda A. Zeder
especially early on in the domestication process. These include both the relaxation
of natural selection once the plant or animal comes under human control and the
adaptation to the new selective pressures on the plant or animal as it enters a
human environment. Random forces also play a role when, often through human-
mitigated movement, small populations of plants and animals are isolated from
broader breeding pools, creating “founder” populations that carry a small, more
or less random selection of the much broader range of genetic variation of the
progenitor population. Another axis on the plant/animal side of the partnership
involves their increasing dependence of the domesticate on the relationship with
humans. Movement along this axis ranges from free-living populations, to man-
aged ones which can still revert to a wild state, to plants and animals unable to
survive outside of the domestic partnership.
The degree of human investment in the plant or animal species forms an
important axis on the human side of the domestic partnership. This axis moves
from no investment (either because humans ignore the plant or animal entirely or
do not engage in any effort to manipulate its availability), to a wide range of
increasingly intensive activities aimed at encouraging the supply of a desired
plant or animal resource. These behaviors may involve altering the plant or
animal’s environment, providing nourishment and protection from predation,
Evolving Mutualism Between
Dependence on Domestic Partnership
Genetic Change & Phenotypic Expression
& Genetic Drift
Plants: tilling, watering, land clearance, selective harvesting, sowing and transplanting
Animals: taming, protecting, selective culling, selective breeding
Human Groups & Plant/Animal Populations
Dependence on Domestic Partnership
Low Level Food Production:
Mix of wild and
Low Level Food Production:
Increasing reliance on
No Investment High
Figure 9.1. Multiple axes of domestication.
229Pathways to Animal Domestication
or intervening into its reproductive cycle (Harris 1996, Smith 2007a, b). Increas-
ing human dependence on the plant or animal forms another axis on this side of
the relationship. In this case, humans move from complete dependence on free-
living populations, through various levels of what has been termed “low-level
food production” based on a mix of managed and free-living resources, to an
agricultural economy in which domesticates make up 40%–60% of human
caloric intake (Smith 2001).
All these different axes operate simultaneously during the domestication pro-
cess. And while every instance of plant or animal domestication involves move-
ment along these axes, not all domesticates travel across these axes at the same
pace or direction. Pathways to domestication vary depending on a range of
morphological, physiological, and behavioral constraints in the target domesti-
cate, the intensity of human investment, the importance of the resource in the
human subsistence economy, and the overall environmental context within which
the relationship unfolds. The challenge to those studying domestication, whether
in plants or in animals, is to identify ways to trace the variable pathways to
domestication and identify the forces that direct humans and domestic partners
along these pathways, in the past and the present and into the future.
2 Basics of animal domestication
Although similar co-evolutionary processes drive the domestication of both plants
and animals, there are fundamental differences between plants and animals that
determine which taxa enter into domestic partnerships with humans and how they
respond to the domestication process once underway. Characteristics that make
certain plants attractive candidates for domestication center on morphological
attributes (i.e., the possession of edible fruits, seeds, or underground storage
organs) or growing habits (i.e., a generalized ability to colonize and adapt to
open, disturbed habitats). Responses to the ongoing domestication process in
plants may take the form of alterations in germination or dispersal mechanisms
as plants adapt to the human-mitigated ecosystems, loss of various defenses
against herbivory due to the relaxation of selective pressures for defense once
humans start tending plants, or changes in fruit size, starch content, and sugar
content that may arise as the result of deliberate human selection for desired traits.
In animals, on the other hand, candidacy for domestication, the targets of
selection under domestication, and the responses to domestication center almost
exclusively on behavioral characteristics (Hale 1969, Clutton-Brock 1981, Price
1984, 2002). Physiological and morphological responses to these pressures in
animal domesticates are often secondary artifacts of the intense selection on
animal behavior (Zeder 2006b). Identifying the behaviors selected for under
domestication and the impact of this selection on domestic animals is, then,
essential for understanding animal domestication.
230 Melinda A. Zeder
2.1 Behavioral characteristics in domestic animals
Attributes thought to be “pre-adaptive” to domestication in animals can be
grouped under ﬁve general categories of behaviors that affect (1) Group structure,
(2) Sexual behavior, (3) Parent–young interactions, (4) Responses to humans, and
(5) Flexibility (Figure 9.2), (Hale 1969, Price 1984, 2002). Many of these behaviors
make it possible for humans to insert themselves in the animal community – either
co-opting leadership of group structure, determining breeding partners and repro-
ductive timing, or assuming a parenting role over young animals soon after birth.
Behaviors that affect an animal’s response to humans are also critical, with those
that determine ﬂight distances and reactivity to external stimuli especially import-
ant in this regard. Other pre-adaptive attributes include behaviors that afford the
animal more ﬂexibility in meeting dietary and other environmental requirements
for survival. In general, the degree to which a species is pre-adapted to domesti-
cation is positively correlated with the degree to which its behavior in its natural
environment resembles its behavior in its captive environment. Species with
the fewest behavioral pre-adaptations to domestication are either never considered
as potential domestic partners or, when domesticated, experience the most exten-
sive changes in response to the selective pressures of the domestication process
Once animals embark on the pathway to a domestic partnership with humans,
the primary target of the new selective pressures introduced by this developing
relationship are those that determine the animal’s response to humans and the
1. Social Structure
a) Large gregatrious social groups
b) Hierarchical group structure
c) Males affiliated with social group
2. Sexual Behavior
a) Promiscuous mating system
b) Males dominant over females
c) Sexual signals provided by movement or posture
3. Parent-Young Interactions.
a) Social bonds created through imprinting
b) Female accepts young soon after parturition or
c) Precocial young
4. Responses to Humans.
a) Short flight distance away from humans
b) Low reactivity to humans or sudden changes in
c) May solicit attention
d) Readily habituated
5. Feeding Behavior & Habitat Choice
a) Generalist feeder or omnivorous
b) Wide environmental tolerance
c) Non-shelter seeking
1. Social Structure
a) Family groupings
b) Territorial structure
c) Males in separate groups
2. Sexual Behavior
a) Monogamous mating system
b) Females dominate males/males appease females
c) Sexual signals provided by markings or morphology
3. Parent-Young Interactions
a) Social bonds created on basis of species characteristics
b) Female accepts young on basis of species
c) Altricial young
4. Responses to Humans
a) Extreme wariness and long flight distance
b) Easily disturbed by humans or sudden changes in
c) Independent/avoids attention
d) Difficult to habituate
5. Feeding Behavior & Habitat Choice
a) Specialized dietary preferences or requirements
b) Narrow environmental tolerance
c) Shelter seeking
Figure 9.2. Pre-adaptive behavioral characteristics in animal domestication. From Hale 1969,
Price 1984, 2002.
231Pathways to Animal Domestication
human environment. In all domesticated animals, the single most important
behavioral response to domestication is reduced wariness and low reactivity to
external stimuli (Price 1998:51–2, 2002:18). This is true of all orders of domesti-
cated mammals, including carnivores (Trut 1999, Coppinger and Coppinger
2001), herbivores (Tennessen and Hudson 1981), and rodents (Murphy 1985), as
well as domestic birds (Andersson et al. 2001) and ﬁsh (Waples 1991), and even
domesticated invertebrate species (Marliave et al. 1993, Price 2002:27–9). And it is
the selection for reduced wariness and low reactivity that has the most profound
and most universal impact on domestic animals.
2.2 Brains and behavior
The most signiﬁcant impact of this intense selection for reduced wariness and
low reactivity to external stimuli is seen in the size, organization, and function
of the brains of domesticated animals. Numerous studies have noted a system-
atic reduction in the overall size of brains in domestic animals compared with
their wild progenitors (Figure 9.3), (Kruska 1988, 1996, Plogmann and Kruska
1990, Ebinger 1995, Ebinger and Ro
¨hrs 1995). Within broad classes of domes-
ticated mammals, there is a positive correlation between the degree of ence-
phalization (brain mass above that related to an animal’s body mass) and
brain size reduction. Mammals with larger brains seem to have experienced the
greatest degree of brain size reduction, whereas smaller-brained mammals may
experience little or no overall reduction in brain size with domestication (Kruska
1988:217, Figure 9.3). Pigs (Sus scrofa) seem to have undergone the greatest
degree of brain size reduction of any domesticate (33.6%), followed by various
domesticated carnivores in which brain size reduction varies between 20% and
30%. Brain size reduction in domesticated ungulate species ranges from 14% to
24%, while domesticated rodents show the smallest degree of brain size reduc-
tion. This same relationship generally holds true among domesticated birds
¨hrs 1985, Ebinger 1995), with the exception of domestic turkeys (Meleagris
gallopavo) which show a large, almost 30%, reduction in brain size compared
with their relatively small-brained wild counterpart – although this large differ-
ence may result from the comparison of a possibly non-ancestral wild subspecies
with a highly improved modern breed of turkey (Ebinger and Ro
¨hrs 1995). Even
captive-reared ﬁsh like rainbow trout (Oncorhynchus mykiss) show signiﬁcant
reduction in brain size compared with wild trout, especially in the areas of
the brain linked to aggression, feeding behavior, and reproduction (Marchetti
and Nevitt 2003).
The degree of brain size reduction does not seem to be positively correlated with
the length of time since original domestication. Sheep (Ovis aries), domesticated
more than 10,000 years ago, display a 24% reduction in brain size compared with
ancestral species (Ovis orientalis), whereas ferrets (Mustela furo), domesticated
for only 2,500 years, show a 30% brain size reduction compared with wild polecats
232 Melinda A. Zeder
(M. putorius). Cage-reared ranch mink (M. vison) have experienced a 20% reduc-
tion in brain size since they were brought under domestication a little over 100
years ago (Kruska 1996). Silver foxes (Urocyon cinereoargenteus) selectively bred
for tameness experienced a reduction in cranial height and width, and by inference
in brain size, after only 40 years of intensive breeding (Trut 1999). It seems likely,
then, that reduction of brain size in animals undergoing domestication may have
occurred relatively quickly during the early phases of the domestication process
and is directly linked to selection for reduced wariness and low reactivity to
Not all parts of the brain are equally affected by the selective pressures intro-
duced by domestication, and there are species-speciﬁc differences in the degree
of size reduction in different parts of the brain in both domestic mammals
(Figures 9.4 and 9.5; Kruska 1988, 1996, Plogmann and Kruska 1990) and in
domestic birds (Ebinger 1995, Ebinger and Ro
¨hrs 1995). In pigs, for example,
brain parts involved in the processing of auditory and olfactory stimuli are less
reduced than visual structures, leading Plogmann and Kruska (1990) to conclude
that structures controlling critical functions in the ancestral species may be less
affected by domestication-induced brain size reduction than less critical functions.
Although the telencephalon, which controls higher thought processes and sensory
perception, is the region of the brain most profoundly reduced in most domestic
mammals, in domestic mink the mesencephalon and the cerebellum, which control
Carnivores Ungulates Rodents BirdsOmnivores
Duck & Geese –16%
Figure 9.3. Reduction in brain size in different groups of domestic animals. Shown as the
percentage of brain mass (corrected for body mass) loss compared to wild ancestral species
at comparable body mass. From Kruska 1988, 1996, Ebinger 1995, Ebinger and Ro
¨mper et al. 2008.
233Pathways to Animal Domestication
body movements, show the greatest degree of size reduction – an adaptation, it is
proposed, to the spatial restrictions of cage rearing in this highly active mustelid
species (Kruska 1996). It is interesting to note in this regard that ranch mink kept
in cages have experienced a much more signiﬁcant overall reduction in brain size
(20%) than mink raised in open-air enclosures, in which brain size is only 11%
smaller than in wild mink (Kruska 1988, 1996, Price 2002:87). Not all brain size
changes in domestic animals are in the direction of smaller size, however.
Although net brain volume is reduced in domestic pigeons (Columba livia) com-
pared to wild rock doves, the hippocampus, important in memory and learning,
is larger, especially among racing and homing pigeons – a likely functional
adaptation to homing that requires spatial cognition and sensory integration
¨mper et al. 2008). As a general rule, phylogenetically younger parts of
the brain are more profoundly affected than are “older” structures (Kruska
1988:219). This general pattern is seen by some animal scientists as evidence of
“regressive evolution” in domestic animals (Ro
¨hrs 1985:547). Others, however,
suggest that these structures change more because they are more plastic than more
basal brain structures and therefore more responsive to the relaxed need for
higher-level brain functioning once humans become buffers between the animal
and its environment (Price 2002:89).
Mesencephalon Total Brain Weight
Total Brain Weight
Total Brain Weight
Total Brain Weight
Total Brain Weight
Total Brain Weight
Figure 9.4. Reduction in total brain mass and size of fundamental brain structures in
several species of domestic animals. Shown as the percentage of brain mass (corrected
for body mass) loss compared to wild ancestral species at comparable body mass.
From Kruska 1988.
234 Melinda A. Zeder
The proposed correlation between the degree of reduction and phylogenetic age
of brain structures is undercut somewhat by the fact that the region of the
telencephalon most profoundly affected by domestication in highly domesticated
animals, such as dogs, sheep, and pigs, is the complex set of structures that belong
to the limbic system, which are embryologically, and likely phylogenetically, some
of the oldest parts of the brain (Figure 9.5). Comprising the hippocampus, the
hypothalamus, the pituitary gland, and the amygdala, the limbic system is respon-
sible for controlling emotionally guided behaviors and memory. It operates by
inﬂuencing the endocrine and the automatic nervous systems that directly control
an animal’s responses to threats and other external stimuli. The profound reduc-
tion in the size of structures within the limbic system in domestic animals can,
then, be directly tied to raising the behavioral thresholds for the display of such
behaviors as aggression, fear, and ﬂight resulting in an overall reduction of
emotional reactivity that is the keystone behavioral attribute of domestic
animals (Kruska 1988:221, Price 2002:89).
RatPig Dog Sheep
Total Brain Weight
Total Brain Weight
Total Brain Weight
Total Brain Weight
Figure 9.5. Reduction in the volume of brain structures in different functional systems
in several species of domestic animals. Shown as the percentage of brain mass (corrected
for body mass) loss compared to wild ancestral species at comparable body mass. From
235Pathways to Animal Domestication
The linkage between behavior, endocrine function, and domestication is power-
fully demonstrated by the work of Ku
¨nzl and colleagues, which compares behav-
ior and endocrine function in domestic guinea pigs (Cavia porcellus) and wild
cavies (Cavia aperea) (Ku
¨nzl and Sachser 1999, 2000, Ku
¨nzl et al. 2003). When
compared with wild-trapped cavies and their ﬁrst-generation offspring, domestic
guinea pigs show signiﬁcantly less aggressive behaviors and more socio-positive
behaviors toward conspeciﬁcs. They are also less attentive to their surroundings,
but are more likely to engage in courtship and sexual behaviors. The endocrine
responses of wild and domestic cavies are also signiﬁcantly different. Wild cavies
subjected to stressful situations (handling and placement in an unfamiliar cage)
register much higher responses in both the pituitary–adrenocortical (PAC) and the
sympathetic–adrenomedullary (SAM) systems, major endocrine stress axes that
under conditions of long-term hyper-activation can contribute to an animal’s
injury or death. At the same time, serum testosterone levels in domestic males
are signiﬁcantly higher than in their wild counterparts, a factor no doubt contrib-
uting to the greater degree of courtship behavior seen among domestic males.
The correlation between higher stress and lower courtship thresholds in domestic
guinea pigs reinforces the notion that, rather than having undergone regressive
evolution, domestic animals have developed highly successful adaptations to
captive environments. Interestingly, captive wild cavies reared in captivity for
30 generations without selective breeding for tameness were found to exhibit
the same behavioral and endocrine responses as wild-trapped cavies, suggesting
that it takes more than simple captivity to bring about the attenuation of
emotional reactivity found in domestic animals (Ku
¨nzl et al. 2003). This ﬁnding
is reinforced by studies that report similar differences in brain chemistry in silver
foxes bred for tameness (Popova et al. 1991a) and in Norway rats (Rattus
norvegicus) selected for reduced aggression to humans (Naumenko et al. 1989,
Popova et al.1991b).
2.3 Pleiotropic effects of selection for behavioral attributes
Selection for reduced wariness and low reactivity may be pleiotropically linked to
other behavioral, physiological, and morphological features commonly found in
domestic animals (Price 2002:79). Among these traits are those that relate to
alterations in developmental events, or heterochrony, especially a reduction in
the rate of change in development known as paedomorphosis. One form of
paedomorphosis commonly found in domestic animals is neoteny, in which an
animal passes through fewer developmental stages before it reaches adulthood so
that as an adult the animal resembles a juvenile stage of its ancestor (Goodwin
et al. 1997). Neoteny may be manifested in the early onset of sexual maturity or in
the retention of both juvenile behaviors and morphology, especially the retention
of juvenile cranial morphology. The classic example of domesticates thought to
display all these neotenized features is the dog (Coppinger et al. 1987, Coppinger
and Schneider 1995, Fox 1978, Goodwin et al. 1997, Morey 1994). Other
236 Melinda A. Zeder
pleiotropically linked effects of selection for reduced aggression may be found in
coat color, especially the manifestation of white markings or piebald coloration,
with a connection drawn between the melanins involved in coat coloration and the
biochemical pathways traveled by neurotransmitters like dopamine that play a
role in shaping behavior and cognition (Hemmer 1990:121–30, see also Keeler
et al. 1968). Features like lop ears and shortened or curled tails, which arise
relatively quickly in foxes bred for tameness, may also be part of a linked complex
of domestic traits (Trut 1999).
Recent work on gene expression provides a possible explanation for the pleiotropic
linkage of seeming disparate behavioral, physiological, and morphological traits in
domestic animals. Qualitative-trait-locus (QTL) analysis of second-generation
crosses between jungle fowl (Gallus gallus) and white leghorn chickens (Gallus
domesticus) successfully identiﬁed loci containing alleles that differentially affect
the expression of the phenotypic traits (i.e., egg production, growth rates, plumage
coloration, fearfulness, and aggression) (Jensen 2006). Speciﬁcally, several of the
QTLs responsible for various productive traits were closely linked with QTLs for
behavior. Extrapolating from these results, the author of this study suggests that
the pleiotropic cascade of traits observed in domestic animals may be caused by
mutations in regulatory genes responsible for the orchestration of gene expression
during development. Under such a scenario only a handful of mutations in
regulatory genes are needed to account for major and rapid evolutionary changes
that separate wild from domestic animals. If so, the intense selection for certain
behavioral attributes of animals embarking on a domestic partnership with
humans could be responsible for a suite of other behavioral, physiological, and
morphological changes in domestic animals.
2.4 The imprint of domestication
The lasting impact of the changes associated with domestication in animals can be
clearly seen in feral animals that have left the domestic relationship and have
reverted to living in a wild state. Sometimes referred to as a process of domesti-
cation in reverse, feralization is often looked to as a model for understanding the
nature and permanence of the changes associated with domestication (Letts 1964,
Hale 1969, Brisbin 1974, Price 1984, 1999, 2002).
Domestication-induced changes in brain size and function may well be irrevers-
ible. Feral dogs, cats, goats, donkeys, and ferrets that have lived outside of a direct
association with humans for many generations show no sign of regaining the brain
mass (Herre and Ro
¨hrs 1990, Birks and Kitchener 1999). Wild mouﬂon (Ovis
orientalis musimon) on Mediterranean islands that a combination of morpho-
logical, cytological and genetic studies conﬁrm are the feralized descendents of
the domestic stock of Neolithic colonizers (Nadler et al. 1973, Poplin et al. 1986,
Bruford and Townsend 2006) retain the smaller brain size of their domestic
ancestors even though they look in every other regard like wild sheep (Groves
237Pathways to Animal Domestication
1989). Dingos (Canis familiaris dingo) in Australia and New Guinea, which have
been living outside of a domestic relationship for thousands of years, have the
same brain size as domestic dogs (Schultz 1969).
A study of feral pigs in the Galapagos found that although there was some
evidence of a reversal of the effects of domestication in certain attributes, other
attributes remained unchanged (Kruska and Ro
¨hrs 1974). Over the approximately
150 years since these animals were introduced onto the islands they have regained
some of the body structure of European wild boar (i.e., longer legs and snouts),
but still retain the coloration of domestic pigs. An examination of the brains of
four of these feral animals found some increase in size of structures related to
olfaction, and the size of the hippocampus may have increased slightly, although it
still remained at least 30% smaller than in wild boar. The brains of feral Galapa-
gos pigs also exhibited a greater degree of variability in the size of the limbic
center. A similar degree of variability in the size of this region of the brain is also
seen in wild pigs, but not in domestic swine. Kruska and Ro
¨hrs suggest that the
greater degree of variability in the size of limbic structures in the Galapagos pigs
might be linked to an increase in aggression and reactivity among these animals,
which, though they lack natural predators on the Galapogos, have been inten-
sively hunted by humans. However, these feral pigs showed no signs of overall
brain size increase, nor was there any detectable increase in the size of the
telencephalon, the region of the brain generally most signiﬁcantly affected by
domestication-induced brain size decrease.
The domestic imprint is also quite evident in the behaviors of feral animals.
Feeding habits of feral domestic cats (Felis catus) in Hungary are quite generalized
and include a relatively wide range of small animals, especially terrestrial
´et al. 2005). Hungarian European wild cats (Felis silvestris), on
the other hand, specialize to a greater degree on particular small mammal species.
They also consume a much higher proportion of arboreal prey that includes both
small perching birds and some larger birds like pheasant and woodcock, suggest-
ing that wild cats had more hunting prowess than feral cats. Studies of feral and
wild cats in Scotland (Corbett 1979) and Iberia (Gil-Sa
´nchez et al. 1999) lend
support to this conclusion. Where rabbit (Oryctolagus cuniculus) populations are
large, wild cats predate them heavily. Feral cats, in contrast, only occasionally
consume rabbits in quantity and when they do they focus mainly on juvenile or
sick individuals. Moreover, although the Hungarian feral cats are much less likely
to consume household foods than scavenging domestic cats, they nevertheless
derive one-ﬁfth of their prey from human settlements (Bı
´et al. 2005). Wild cats
in this study took no prey from human settlements. Free-ranging cats and dogs in
southeastern Brazil are found in signiﬁcantly higher density in suburban areas
than in rural ones, suggesting that these animals retain a strong connection to the
human environment and are not exploiting more open niches in rural areas to the
same extent (Campos et al. 2007).
In a broad-ranging synthesis of the literature on the social ecology of feral dogs
(Canis familiaris), Boitani and Cuicci (1995) conclude that even fully feralized
238 Melinda A. Zeder
animals, which receive no food or shelter from humans and show a strong
avoidance of human contact, are still dependent on the human niche for survival
and have not regained the self-sustaining behaviors found in wolves (Canis lupus).
Whereas wolves live in single family packs with established hierarchical social
structure, feral dogs live in ﬂuid groups made up of breeding pairs. The size of
these groups is limited by the lack of social structure and other social bonds that
keep wolf packs operating as functional units. Although feral dogs generally have
larger litters and breed more frequently than wolves, they lack the parenting skills
of wolves, often leaving very young pups unattended, resulting in high juvenile
mortality. Older feral pups also suffer higher mortality rates than wolf pups of
comparable age when they leave the den to explore and forage on their own or
when their mother enters a new oestrous cycle and loses interest in her most recent
litter. As a result, groups of feral dogs are not self-sustaining and can only be
maintained through recruitment of new members from populations of stray dogs.
Feral dogs do not hunt in packs like wolves and, again unlike wolves, feral pups
are not taught to hunt by adult animals. As a result these dogs have a highly
diversiﬁed diet comprising primarily smaller, easier to catch prey. Although they
may not receive food directly from humans, they still concentrate on human-
mitigated landscapes when foraging for food.
Thus the imprint of domestication on animals is profound. Domestication-
induced changes in brain size and function are deep-rooted and perhaps irrevers-
ible. Behavioral attributes selected for during domestication make it difﬁcult, if
not impossible, to recapture the behaviors and social ecology that sustained their
progenitors in the wild. Feral animals attempting to divorce themselves from the
domestic partnership with humans seem only partly successful in doing so and in
many ways remain strongly tied to humans, even if indirectly.
3 Pathways to domestication
There seem, then, to be universal attributes found in almost all domestic animals
in the behaviors selected for under domestication and the impact of this selection
on brain form and function. But did all domestic animals acquire these attributes
in the same way? Did all animal domesticates travel the same general pathway that
took them from free-living wild animals to animals forever tethered to a partner-
ship with humans, even when they attempt to revert to the wild state? I think that
there is ample evidence that this is not the case and that even though there are
universal attributes found in all animal domesticates, the pathways to animal
domestication were highly variable and contingent on broadly deﬁned biological
and cultural parameters, as well as a range of highly localized factors that shaped
the trajectories of individual cases of animal domestication (Zeder 2009). These
varied pathways can, however, be grouped into three general domestication
scenarios that seem to account for the full spectrum of animal domesticates – a
commensal pathway, a prey pathway, and a directed pathway (Figure 9.6).
239Pathways to Animal Domestication
3.1 Commensal pathway
The ﬁrst of these pathways, the commensal pathway, is most frequently traveled
by animals that come into initial contact with humans to feed on refuse or to
prey on other animals attracted to human settlements. At some point in their
association with humans and human habitats, these animals developed closer
social or economic bonds with their human hosts than did other commensals
inhabiting this niche. These bonds brought them, eventually, into a domestic
partnership with humans. The classic example of an animal that likely traveled
this pathway to domestication is the dog, whose domestication is thought to have
begun when less wary wolves were drawn to human encampments to scavenge on
human refuse (Coppinger and Coppinger 2001, Morey 1994). In a comparative
study of the skeletal morphology of wolves and dogs, Morey (1992) concludes
that the principal difference between the dog and its wild progenitor lies in the
juvenilization of the adult dog’s cranial morphology, which resembles that of a
juvenile wolf in its shorter face, steeped forehead, and wider cranial dimensions.
Morey suggests that these differences in cranial morphology are artifacts of
neotenization arising out of a general process of paedomorphism in develop-
mental rates (see also Trut 1999, but see Wayne 1986, who contends that dog
and wolf skulls differ primarily in cranial width, which is not a paedomorphic
trait). The mandibles of dogs are shorter than those of wolves and there is
Dogs (Canis familiaris)
Cats (Felis catus)
Pig (Sus scrofa)?
Guinea pig (Cavia porcellus)
Golden hamster (Mesocricetus auratus)?
Chicken (Gallus domesticus)
Muscovy duck (Cairina moschata)
Turkey (Meleagris gallopavo)
Commensal Domesticates Prey Domesticates Directed Domesticates
Goat (Capra hircus)
Sheep (Ovis aries)
Cattle (Bos taurus)
Zebu cattle (Bos indicus)
Pig (Sus scrofa)?
Water buffalo (Bubalus bubalis)
Mithan (Bos frontalis)?
Bali cattle (Bos javanicus)?
Yak (Bos grunniens)
Llama (Lama glama)
Alpaca (Lama pacos)
Reindeer (Rangifer tarandus)
Horse (Equus caballus)
Donkey (Equus asinus)
Dromedary (Camelus dromedarius)
Bactrian camel (Camelus bactianus)
Buffalo (Bison bison)
Ferret (Mustela furo)
Mink (Mustela vison)
Silver fox (Urocyon cinereoargenteus)
Chinchilla (Chinchilla lanigera)
Emu (Dromaius novaehollandiae)
Ostrich (Struthio camelus)
Recent aquatic domesticates
Asian elephants (Elephus maximus)
African elephants (Loxodonta africana)
Cheetah (Acinonyx jubatus)
Peregrine falcon (Falco peregrinus)
European buzzard (Buteo buteo)
Tam e d Ca pti ves
Red deer (Cervus elaphus)
Père David’s deer (Elaphurus davidianus)
Fallow deer (Dama dama)
Blackbuck (Antilope cervicapra)
Eland (Taurotragus oryx)
Musk ox (Ovibos moschatus)
Barbary sheep (Ammotragus lervia)
Experiments in Domestication
Figure 9.6. Possible pathways to domestication of animal species mentioned in the text.
240 Melinda A. Zeder
considerable reduction in the length of their molars (Wayne 1986). These features
are found in even the earliest examples of domestic dogs like the 13,000-year-old
mandible recovered from Palegawra Cave in northeastern Iraq, which shows clear
evidence of tooth size reduction and crowding in a shortened jaw (Turnbull and
Reed 1974). The dogs buried with humans at the roughly contemporary site of Ain
Mallaha in the southern Levant also display these traits, and their occurrence in
burial contexts speaks to the strong social bonds that had been forged between
dogs and humans at this early date (Davis and Valla 1978, Tchernov and Valla
1997, Morey 2005). This same site also contains some of the earliest remains of
commensal animals like the house mouse (Mus musculus), the spiny rat (Echimys
chrysurus), and the house sparrow (Passer domesticus) (Tchernov 1991), species
that traveled a commensal pathway into close association with humans but that,
unlike dogs, did not complete the journey to domestication. Recent genetic data
support archaeological evidence that suggests initial dog domestication took place
in the Near East (vonHoldt et al. 2009).
A surprising number of common domestic animals may have traveled this same
pathway to domestication. Archaeological evidence pushes back the date of cat
domestication to at least 8,500 years ago (Vigne et al. 2004), and new genetic
evidence points to a Near Eastern origin for initial cat domestication (Driscoll
et al. 2007). It is likely, then, that cats, like dogs, were drawn into initial contact
with humans when humans began to live in more permanent settlements and these
obligate carnivores were attracted to human habitats to prey on other small
commensal species occupying this niche. Interestingly, however, cats did not
experience similar changes in cranial morphology, nor does the social ecology of
these more solitary animals seem as profoundly altered by domestication.
Chickens, which genetic evidence suggests were domesticated multiple times in
southeast Asia, China, and perhaps India (Liu et al. 2006, Kanginakudru et al.
2008), may also have entered the domestic relationship with humans through a
commensal pathway as wild jungle fowl sought out human dump heaps for easy
sources of grain. A similar pathway might be suggested for the domestication of
the turkey in the southwestern US (Munro 2011). Muscovy ducks (Cairina
moschata) in Amazonia have been argued to played an important role in reducing
insect populations in human settlements (Angulo 1998), an outgrowth, perhaps of
a similar initial commensal introduction into the human sphere. Another likely
commensal domesticate is the guinea pig (Cavia porcellus), ﬁrst domesticated in the
highland Andes around 7,000 years ago (Spotorno et al. 2006). Golden hamsters
(Mesocricetus auratus), which are indigenous to Syria, Israel, and eastern Turkey and
known for the ease with which they can be handled in the wild, are another candidate
for a commensal domesticate – although the hamsters kept so widely today as pets
and laboratory animals are thought to be descendants of a quite recent and deliber-
ate domestication of hamsters from dwindling wild populations (Murphy 1985).
It is also possible that pigs, a major livestock species, entered into domestication
through a commensal pathway. Archaeological evidence from the site of Hallan
C¸ emi in southeastern Anatolia suggests that, as with dogs and cats, a special
241Pathways to Animal Domestication
relationship between pigs and humans began as early as about 12,000 calendar
years ago, soon after humans began living in more established year-round settle-
ments (Redding 2005). There are multiple lines of evidence for the intensiﬁcation
of this relationship over the course of nearly 3,000 years of occupation (from
10,500 to 8,300 cal BP) at the near by site of C¸ ayo
¨nu (Ervynck et al. 2001, Hongo
et al. 2002). As with dogs, the leading edge indicator of pig domestication is a
gradual reduction in molar length, a marker of domestication in pigs thought to
result from the neotenization of skull morphology (Flannery 1983). These gradual
changes in tooth morphology are ﬁrst detected several hundred to a thousand
years prior to the appearance of other markers of pig domestication at this site.
Ervynck et al. (2001) see this long lead-up period as evidence of an evolving
mutualism between pigs and humans initiated by wild pigs originally drawn to
human settlements to scavenge off refuse dumps.
3.2 Prey pathway
Most major livestock species, however, entered into domestication through what
might be called a prey pathway. Rather than initiating the relationship, these
animals were primary prey species that humans had hunted for their meat and
hides for thousands of years. The prey pathway likely began when, perhaps as a
response to depletion of local stocks of these prey animals, humans developed
hunting strategies designed to increase prey availability. Over time and under
certain circumstances, these game management strategies developed into actual
herd management and, eventually, the controlled breeding of managed animals.
Archaeological evidence from the Near East suggests that sheep (Ovis aries),
goats (Capra hircus), and cattle (Bos taurus) all followed this pathway to domesti-
cation, with the transition from generalized hunting to specialized hunting and
then herd management taking place within the natural habitats of wild progenitor
species (Zeder 2008a, 2009, 2011). This process seems to have unfolded over many
hundreds, if not thousands, of years without any clear-cut, archaeologically
detectable morphological changes in the animals traveling down this pathway to
domestication. These early stages of the transition from hunting to initial man-
agement may only be detectable in the demographic proﬁles of the animals
harvested by humans, especially in the separate harvest patterns of male and
female animals. Constructed using the fusion patterns of post-cranial skeletal
elements, these sex-speciﬁc harvest proﬁles are capable of distinguishing between
the prey strategies of hunters that seek to maximize immediate meat return (often
reﬂected by an emphasis on large adult males) from those of herders directed at
promoting herd growth (most commonly met by the early harvest of all but a few
males and the delayed cull of older females past peak reproductive years) (Zeder
2006b, 2008b). This distinctive herd management harvest proﬁle is ﬁrst detected in
goat assemblages from the archaeological site of Ganj Dareh in the Central
Zagros Mountains of modern-day Iran at about 10,000 calendar years ago (Zeder
and Hesse 2000, Zeder 2006b, 2008b). Lower-resolution harvest data collected by
242 Melinda A. Zeder
using other methods, however, suggest that the management of both sheep and
goats began perhaps 500 to 1,000 years earlier in the highland regions of the
eastern Taurus and northwestern Zagros Mountains (Peters et al. 2005, Zeder
2008a, 2009, 2011). The initial phases of the transition from hunting to herding
in this region may also reach back to about 12,000 to 13,000 calendar years ago.
New demographic data from southeastern Turkey and northwestern Iraq point to
the development of hunting strategies, which may have helped restock local herds
of wild sheep depleted by people living in increasingly sedentary settlements
(Redding 2005, Zeder 2008b, 2009). Demographic data for cattle suggest that a
similar process was underway in the upper Euphrates Valley by about 10,500 to
10,000 years ago (Helmer et al. 2005).
A recent genetic analysis by Naderi et al. (2008) has identiﬁed all six domestic
goat lineages among modern wild bezoar goats from eastern Turkey and western
Iran. The authors of the study argue that the presence of these domestic hap-
logroups among wild goats is not an artifact of recent introgression between
domestic and wild goats. They maintain instead that genetic signatures of popula-
tion growth and geographic translocation represent residual evidence of initial
human domestication of these different lineages and the human-mitigated move-
ment of managed animals within and out of this heartland region of initial
domestication. This exciting new study suggests that although the prolonged
periods of human management of goats within their natural habitat had no
detectable morphological impact on these animals, it nevertheless left a genetic
imprint observable even today among the descendant populations of wild goats
from which domestic lineages were originally drawn.
Domestication-induced morphological change in animals traveling along this
prey pathway may only be detectable once humans took managed herds out of the
natural habitat of their wild progenitors where factors like genetic drift and
adaptation to new environments came into play. The movement of managed herds
outside of the range of wild populations also eliminated the chance of introgres-
sion between wild and managed animals or the possibility of restocking managed
herds with wild animals – both probably quite common occurrences in the initial
phases of herd management. Once this link was cut in managed sheep and goats,
we begin to see distinctive changes in horn size and shape like those evidenced at
about 9,500 to 9,000 cal. BP among the remains of goats recovered from the
archaeological site of Ali Kosh in lowland Iran (Zeder 2006b). Changes in the size
and shape of horns of domesticated ungulates like sheep, goat, and cattle likely
arose from a combination of factors including: (1) the relaxation of selective
pressures for large horns previously used to both attract and compete for females,
(2) the expression of random mutations previously selected against when horns
were used in mate competition, (3) the impact of new selective pressures against
energetically expensive and no-longer-needed horn architecture, and (4) directed
human selection for more tractable males.
These same factors likely also played a role in changes in body size in these early
domestic livestock species. However, rather than an overall reduction in the body
243Pathways to Animal Domestication
size of initial domesticates as was once thought to be the case (Uerpmann 1979,
Meadow 1989), it now seems more likely that domestication-induced body size
changes in early livestock species took the form of a reduction in the degree of
sexual dimorphism, especially a shortening in the length of the legs of males
(Zohary et al. 1998, Zeder 2001, 2006b, Helmer et al. 2005). Smaller body size in
archaeological populations of managed sheep and goats is not seen until sometime
after 9,000 cal BP, and it is not clear whether the smaller size of these animals is an
artifact of domestication, of climate change, of the introduction of smaller-bodied
domestic stock from different regions, or of a general process of body size reduc-
tion that began with the end of the last Ice Age and has affected domestic and wild
ungulates alike (Zeder 2006b, 2008b).
In addition to these core Near Eastern livestock species, it is likely that other
common animal domesticates followed this prey pathway to domestication. This
includes East Asian sheep, which genetic data suggest were independently brought
under domestication (Guo et al. 2005) and perhaps the pigs independently domes-
ticated in East Asia and in Europe (Larson et al. 2005, 2007). It is hard to say in
the case of pigs, however, whether these separate domestication events followed a
commensal path, a prey path, or in the case of the European wild boar the ﬁnal
directed pathway discussed below. Other likely prey pathway domesticates are the
humped zebu cattle (Bos indicus) and the water buffalo (Bubalus bubalis) domesti-
cated in South Asia (Fuller 2006, Kumar et al. 2007). The yak (Bos grunniens)is
another early domesticate that may have been brought under domestication in this
way in the Himalayas (Olsen 1990, Guo et al. 2006). The mithan (Bos frontalis)of
South Asia and the Bali cattle (Bos javanicus) of island Indonesia, whose origins
are poorly understood (Clutton-Brock 1981:137–8), may represent additional
examples of prey pathway domesticates. The increasingly well-resolved record of
the domestication of South American camelids clearly points to a prey pathway
along which the heavy predation of the guanaco (Lama guanaco) and the vicun
(Vicugna vicugna) developed into initial management and then full domestication
of the llama (Lama glama) and the alpaca (Lama pacos) (see Mengoni-Gon
and Yacobaccio 2006, Wheeler et al. 2006).
Reindeer (Rangifer tarandus) may be the most recent, and perhaps last, species
to follow a prey pathway to domestication. In many ways, reindeer herding serves
as a good model for the initial stages of domestication of other prey pathway
domesticates like sheep and goats. The only successfully domesticated cervid
species, these cold-adapted gregarious herd animals have been heavily predated
by humans since the last Ice Age (Speiss 1989). The close association between
hunters and reindeer in the northern Eurasian Holarctic stretches back thousands
of years with the loose domestic partnership between humans and reindeer
thought to have developed sometime in the past 2,000 to 3,000 years (Mirov
1945, Gordon 2003). Baskin (1974) sees reindeer herding as the product of
sophisticated hunting methods in which reindeer hunters, familiar with the migra-
tory routes of wild reindeer herds, drove reindeer into stone traps or human
settlements where they could be harvested at will. Over time northern peoples
244 Melinda A. Zeder
developed a number of different reindeer herding strategies including a “close”
herding system that involves following large demographically diverse herds of
migrating animals, a “free-camp” system in which smaller herds are kept within
the vicinity of human settlements, and a “loose system” in which free-ranging
animals are periodically gathered and moved to different pastures (Baskin 2000).
Managed reindeer are exploited for their meat, hides, and antlers, for their use as
draft animals and for riding and traction, and, to a lesser extent, for milk.
Reindeer herding takes place alongside active hunting of wild reindeer, with
domestic females sometimes used in the past as “bait” to attract wild males
(Manker 1963:16). As appears to have been the case with other prey domesticates
like sheep and goats during initial stages of their management, there are no major
morphological differences between wild and domesticated reindeer that would be
detectable archaeologically (Clutton-Brock 1981:134).
Recent genetic analysis of modern wild and domestic reindeer from localities
across Eurasia (Røed et al. 2008) extends the homology between reindeer herding
and the initial stages of caprine domestication even further. As with sheep and
goat, there is evidence for multiple independent reindeer domestication events
within the natural habitat of the wild reindeer. One such “event” was apparently
localized in the western part of their range in Fennoscandia, with perhaps two
additional events occurring in western and eastern Russia. The high level of
genetic diversity in domestic reindeer herds is seen as an artifact of the frequent
augmentation of domestic herds with local wild reindeer. The authors of this study
also found evidence for the frequent introgression of domestic haplotypes into
wild herds. Some wild populations in Finland and Norway and a population in
southeastern Russia, however, seem to have contributed no genetic material to
domestic stock. Røed et al. interpret the different contribution of various wild
populations to domestic populations as evidence of the differential domestication
potential of wild reindeer. The more gregarious populations residing in open
tundra habitats, they argue, were more attractive candidates for domestication.
Forest-dwelling populations that may have been less well pre-adapted to domesti-
cation, however, seem not to have played a role in this process. This system has
many parallels with that documented in the Naderi et al. (2008) study of goats
in the Zagros, providing a living model of a management system which, though
it leaves no mark in the morphology of the managed animals, has a lasting
genetic imprint on both managed animals and the wild populations from which
they were drawn.
3.3 Directed pathway
The prey pathway was likely driven by the goal of securing a predictable source of
protein in the form of animal ﬂesh. But it did not take long before people started
to exploit other, largely regenerative secondary animal resources. Recent analysis
of lipid residues found in pottery from sites in Turkey and the Levant indicates
that dairying may have been well established, especially in northwestern Anatolia,
245Pathways to Animal Domestication
by about 8,500 calendar years ago (Evershed et al. 2008). A ﬁgurine of a wooly
sheep from the site of Sarab in the highlands of western Iran has been interpreted
as evidence that changes in coat composition needed for wool production were
in place by about 7,500 calendar years ago (Bo
¨nyi 1977). Finding direct
evidence for the use of animals like cattle for traction is difﬁcult, but the discovery
of a ceramic bull with a churn on its back in the southern Levant suggests that
the use of cattle for both dairy products and labor had been established by
6,000 years ago (Ussishkin 1980). The precedent set by the domestication of
former prey species and the broadening of the range of resources extracted from
them paved the way for the ﬁnal category of animal domestication – the directed
pathway. This fast-track to domestication begins when humans use knowledge
gained from the management of already domesticated animals to domesticate a
wild species that possesses a resource or a set of resources that humans see
This is likely the pathway followed in the domestication of the horse (Equus
caballus), which both archaeological and genetic evidence suggests was domesti-
cated, perhaps multiple times, across the steppe regions of central Eurasia (Levine
1999, Olsen 2006, Vila
`et al. 2006). Possibly originally domesticated to help in the
hunting of wild horses (Olsen 2006), domestic horses also provided people with a
wide array of primary and secondary resources, including meat, hides, milk, draft,
traction, and transport. It is interesting to note that none of the traditional
markers used to track domestication in animals that followed the commensal or
prey pathways is useful in documenting horse domestication. There are no appar-
ent morphological markers that can be used to discriminate domestic horses from
wild horses (E. ferus), nor are demographic proﬁles much use in distinguishing
management strategies from prey strategies. Instead, archaeologists employ mul-
tiple lines of circumstantial evidence to monitor this process, including butchery
practices, evidence of corrals, the presence of quantities of manure signaling corral
cleaning, or the use of manure as building materials, and changes in long-distance
transport of lithic resources (Olsen 2006). The most recent, and perhaps most
compelling, evidence of horse domestication is provided by the successful retrieval
of equine milk lipids in 5,500-year-old pottery from northern Kazakhstan
(Outram et al. 2009).
Donkeys (Equus asinus) are another animal that likely entered into the domestic
partnership with humans through this route. Genetic evidence puts the initial
domestication of two populations of wild Nubian ass (E. a. africanus) in northern
and northeastern Africa (Kimura et al. 2010), with recent thinking crediting their
domestication to pastoral people who about 6,000 years ago began to use these
desert-adapted animals to carry heavy loads across arid lands (Rossel et al. 2008).
As with the horse, however, traditional archaeological markers of animal domesti-
cation have been of little utility in tracing the process of donkey domestication.
Rossel et al. (2008) provide compelling evidence for the use of donkeys as beasts of
burden in their analysis of ten complete ass skeletons recovered from an early
pharaonic mortuary context at Abydos in Middle Egypt. All of these ritually
246 Melinda A. Zeder
slaughtered animals show unambiguous signs of, often quite advanced, spondylo-
arthopathies, vertebral pathologies consistent with the exertion of considerable
pressure on the spine. Appendicular skeletal elements of these animals also display
considerable compression-induced pathologies, leading to the unmistakable con-
clusion that these were fully domesticated animals used to carry heavy loads.
Morphometric analysis, however, found that the metapodials of these domestic
donkeys (the bone thought most likely to respond to any changes in body size
or life habits in these animals), closely resemble the metapodials of wild asses
(especially the Nubian wild ass) and, in most respects, are quite distinct from the
metapodials of modern domestic donkeys. Even though the ass was likely brought
under domestication at least 1,000 years before the burial of these ten sacriﬁcial
animals, the only hint of domestication-induced morphological change in the
Abydos donkeys is a slight modiﬁcation in metapodial mid-shaft depth and distal
Other likely instances of directed domestication are provided by Old World
camels, both the one-humped dromedary (Camelus dromedarius) of the Arabian
Peninsula and the two-humped Bactrian camel (C. bactrianus) of Central Asia.
Once again there is little direct evidence for camel domestication. There are no
archaeologically detectable morphological differences between the domestic and
wild two-humped camels (Peters and von den Driesch 1997). Moreover, although
there are no longer any wild one-humped camels to compare with the domestic
dromedary, there is little evidence of morphological differences in the skeletons of
camels from likely pre- and post-domestication contexts in the Arabian Peninsula
(Clutton-Brock 1981:124–6). In the absence of distinctive morphological change,
the presence of camel dung and hair at the site of Shahri-Sokhta in far eastern Iran
and the recovery of ﬁgurines of camels attached to clay carts from archaeological
sites in Turkmenistan dated to between 3,000 and 2,500 BC have been interpreted
as circumstantial evidence that two-humped camels had been domesticated by the
third millennium BC (Masson and Sarianidi 1972, Compagnoni and Tosi 1978,
but see Peters and von den Driesch 1997). A case of the domestication of the one-
humped camel in the Arabian Peninsula is based on the abundance of camel
remains from third millennium sites in Oman and associated mortality patterns
indicating a shift from a prime adult harvest strategy to an emphasis on the
slaughter of juvenile animals (Hoch 1979, but see Uerpmann and Uerpmann
2002). If indeed both these species were brought under domestication during
the third millennium BC, the primary target of these two instances of directed
domestication may well have been their utility in carrying people and goods across
vast arid regions in both Central Asia and in the Arabian Peninsula and the
important role these animals might have played in the active global trade networks
that developed during this time (Zeder 2006c, Zeder et al. 2006).
Elephants, both Asian and African (Elephas maximus and Loxodonta africana),
also represent animals brought under human control for a directed purpose –
either for carrying large loads or for heavy labor (a purpose for which Asian
elephants are still used today), or for use in hunting, warfare, or in public
247Pathways to Animal Domestication
spectacles (as was the case for both African and Asian elephants in antiquity and
today with circus performers) (Clutton-Brock 1981). Given the long life span of
elephants, their slow maturation rate, the difﬁculty in getting captive elephants to
breed, and the relative ease with which captive young adults can be tamed and
trained to perform desired tasks, elephants used for these purposes are generally
not bred in captivity (Baker and Manwell 1982). Instead, domestication begins
anew with each young animal that is captured and tamed. Another captive animal
that has been tamed and used for a directed purpose is the cheetah (Acinonyx
jubatus) that Ancient Egyptians, Assyrians, Mogul emperors, and Medieval
European elites kept as pets and hunting companions (Clutton-Brock 1981).
Falcons and other trained birds of prey (e.g., Falco peregrinus and Buteo buteo)
used in hunting are also generally not bred in captivity, another example of a
captive animal brought under human control for a speciﬁc purpose that did not
follow the subsequent pathway to full-ﬂedged domestication.
Recent examples of directed domestication include the various carnivore and
rodent species, like mink (Mustela vison) and chinchilla (Chinchilla lanigera) that
have been selectively bred for coat quality over the past 100 – 200 years. Other
examples of animals following this pathway to domestication also include even
more recent domesticates, like buffalo (Bison bison), emu (Dromaius
novaehollandiae), and ostrich (Struthio camelus) bred for their meat and hides.
Experiments in domestication are currently underway, with mixed success, with a
number of terrestrial mammals including red deer (Cervus elaphus), Pe
deer (Elaphurus davidianus), fallow deer (Dama dama), blackbuck (Antilope
cervicapra), eland (Taurotragus oryx), musk ox (Ovibos moschatus), and Barbary
sheep (Ammotragus lervia) (Clutton-Brock 1981:177–87, Hemmer 1990:161–77).
The number of freshwater and marine species (both vertebrates and invertebrates)
brought under human management has increased markedly over the past 100 years.
Ninety-seven percent of the 430 currently managed aquatic species were brought
under human control during the twentieth century, 100 of these species in just the
past 10 years (Duarte et al. 2007). The staggering explosion of aquaculture as a
major world-wide industry has momentous implications for the human food
supply, for biodiversity, and for the environment.
3.4 A domestication road atlas
There are, then, multiple pathways to animal domestication, which vary in
length, direction, and travel time (Figure 9.7). The progress of individual animal
domesticates and their human partners down these different paths is highly
variable and shaped by the combination of constraints and opportunities,
biological and cultural, that these fellow travelers face while they make the
It probably took a long time for animals traveling the commensal pathway to
move from being simply habituated to humans and human habitats to developing
an active partnership with humans (Figure 9.7a). The timing and the nature of the
248 Melinda A. Zeder
forces that propelled the next and ﬁnal stage of the journey – human-directed
breeding and more or less complete subjugation to human control – likely varied
in different commensal domesticates. And some animals entering into domestic
partnership through this route – like the cat – have arguably never reached this
While animals traveling the commensal pathway began the journey on their
own initiative, animals entering into a domestic partnership with humans though
the prey pathway were likely less willing fellow travelers. As with the commensal
route, however, progress down the prey pathway was also likely quite slow and
possibly circuitous, as generalized hunting strategies evolved into game manage-
ment strategies aimed at promoting availability of prey species, which, in turn,
morphed into the selective harvest of managed animals, followed, at some
perhaps quite distant part of the road, by directed breeding (Figure 9.7b). But
not all animals traveled this route in the same way, nor did all animals
embarking on this pathway reach its conclusion. A strong case can now be made
that sheep, goats, and cattle all traveled a quite direct, if lengthy and slow, prey
pathway to domestication in central and eastern portions of the Fertile Crescent
arc that stretches from southern Iran, across northwestern eastern Iraq and
southeastern Turkey, into Lebanon, Israel, and eastern Jordan. Pigs in the
central Fertile Crescent, in contrast, may have wandered between prey and
commensal pathways at different points of the journey. Moreover while humans
and these four future livestock species were beginning down pathways to domes-
tication, in the western arm of this Fertile Crescent region people and gazelle
(Gazella subgutturosa and G. gazella), the primary prey species in the northern
and southern Levant, may also have been taking the ﬁrst tentative steps down a
prey pathway to domestication. Gazelle hunting strategies seem to intensify in
the Levant at about 12,000 calendar years ago (Munro 2004) and there is some
Habituation Partnership Directed Breeding
Game Management Herd Management Directed Breeding
Competitor Directed Breeding
Figure 9.7. Pathways to domestication. (a) Commensal pathway, (b) prey pathway,
(c) directed pathway.
249Pathways to Animal Domestication
indication that humans were altering prey strategies to promote availability of
this key resource in ways that were having some impact on the demographic
structure of gazelle populations (Davis 1983, Henry 1989, Cope 1991, but see
Sapir-Hen et al. 2009). Parallels can be found between the corrals and traps used
much later by Eurasian Arctic peoples early on in the process of reindeer
domestication (Baskin 1974) and the stone kites and possible corrals found
throughout the Levant that are thought to have been used to capture migrating
gazelle (Legge and Rowley-Conwy 1987, Betts and Yagodin 2000, Bar-Oz et al.
2011). Redding (2005) has recently suggested these structures may also have
served as holding areas for animals that could be consumed as needed, clearly
a step toward animal management. Gazelle, however, are behaviorally less well
suited than sheep and goat to domestication. They have very strongly developed
ﬂight reﬂexes and an aversion to penning, and are highly territorial and unlikely
to breed well in captivity (Clutton-Brock 1981:172). These behaviors are thought
to have made it impossible for gazelle to travel much further down the prey
pathway than the early stages of game management.
The ﬁnal directed pathway is a much shorter and speedier route to animal
domestication. Animals traveling down this path may have begun as human prey
or competitors for prey, or may have had little or nothing to do with humans.
However, once embarked on this pathway they took an immediate and abrupt
departure from a free-living state to one in which they were under tight human
control that often involved intensive and deliberate breeding to enhance targeted
resources. Animals selected for the directed pathway may have possessed few of
the pre-adaptive behaviors that qualiﬁed other animal domesticates for a trip
down either the commensal or prey pathways, and their domestication likely
required intensive efforts to overcome behavioral and biological barriers to
domestication. Today this route has become a kind of domestication super-
highway as animals that previously would never have been considered candidates
for domestication are brought under human control through the application of
increasingly sophisticated technology for animal breeding and care, and the
enhanced understanding of animal behavior, reproduction, and biological require-
ments coming out of the animal sciences (Price 2002:22).
4 Questions for future research
Categorization of the various pathways to animal domestication in this way raises
a number of questions that point to productive areas for future research. First, it
would be interesting to know whether there are differences in the timing and the
nature of the behavioral, physiological, and morphological responses animals
make to the selective pressures they experience along these different pathways to
domestication. A related question asks whether these selective pressures leave
distinctive genetic, morphological, or other archaeological markers that can be
used to detect the various paths taken to domestication. Are animals that enter
250 Melinda A. Zeder
domestication through a commensal route, with its long “getting-to-know-you”
phase of habituation to humans and human environments, tamer and more
integrated into human society than animals that traveled the prey pathway? Are
the changes in cranial form seen as an early marker of domestication in dogs and
pigs, but not as apparent in animals such as sheep, goats, or horses, an artifact of a
more prolonged and perhaps more intense selection for human habitation? If so,
why is it that other animals that may have taken this same commensal route, such
as the cat, do not display similar cranial morphology? Why is there often little
evidence of morphological change in animals brought into the domestic relation-
ship by way of the directed route? Are there different genetic signatures that can be
used to trace the behavioral adaptations that grew out of the commensal relation-
ship and differentiate them from those that arose through the prey pathway? Does
the intensive, focused selection for speciﬁc traits under directed domestication
leave a distinctive genetic signature distinguishable from the genetic signatures left
by the broader play of selective factors and random events that shaped both the
commensal and prey routes?
The different capacities for feralization of different domesticates and the
residual imprint of domestication seen on these animals raises another set of
questions. Among them, are commensal domesticates more successful feral
animals because they can revert to the commensal behaviors that brought them
into the relationship in the ﬁrst place? How do feral commensal domesticates vary
in behavior, physiology, and morphology from commensal species that never
traveled any further down this path?
Looking forward, this discussion raises a series of questions about ongoing
processes of domestication and breed improvement. Are there lessons that might
be drawn from the different pathways humans and animals followed to domesti-
cation in ancient times that might be applied to current-day breed improvement
programs? Do ancient efforts at game management and initial herd management
have bearing on current-day ranching practices directed at animals like the eland
and the fallow deer? Can this perspective help animal scientists better balance the
dual goals of enhancing both animal productivity and animal welfare (Grandin
and Dessing 1998, Price 2002:204–29, Keeling and Jensen 2002)? What are the
environmental and biodiversity impacts of the massive wave of recent directed
domestications, especially those involving aquatic species, and how can a broader
understanding of the multiple pathways to domestication help mitigate these
Finally, recognition of the multiple pathways to domestication and their impact
on domestic animals has a bearing on a range of issues involving the care and
welfare of captive animals, as well as conservation efforts directed at endangered
species (O’Regan and Kitchener 2005). Can these different models of domesti-
cation contribute to a better understanding of the impact of captivity on wild
animals kept in zoos or in captive breeding programs? Are there parallels to be
drawn between feralized domesticates and the reintroduction of captive animals
into the wild?
251Pathways to Animal Domestication
Answering these questions requires drawing broadly from genetics, animal
sciences, and archaeology. As this review has shown, researchers based in each of
these general disciplinary areas are actively contributing to a large and growing body
of knowledge on animal domestication and the potential for cross-illumination
between these different perspectives is only just beginning to be realized. Clearly
the interdisciplinary model set by Jack Harlan in his career-long study of crop
evolution holds much promise for the study of animal domestication. And it is
rewarding to see researchers from all of these different disciplinary backgrounds
included in the second Harlan International Symposium and the publication of the
proceedings of that stimulating meeting. Hopefully, contributors to the third Harlan
International Symposium will be able to report considerable progress in addressing
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259Pathways to Animal Domestication