Niche Construction and the Behavioral Context
of Plant and Animal Domestication
BRUCE D. SMITH
Many animal species attempt to enhance their environments through niche con-
struction or environmental engineering. Such efforts at environmental modification
are proposed to play an important and underappreciated role in shaping biotic
communities and evolutionary processes.1,2 Homo sapiens is acknowledged as
the ultimate niche constructing species in terms of our rich repertoire of ecosys-
tem engineering skills and the magnitude of their impact. We have been trying to
make the world a better place?for ourselves?for tens of thousands of years. I
argue here that it is within this general context of niche-construction behavior that
our distant ancestors initially domesticated plants and animals and, in the pro-
cess, first gained the ability to significantly alter the world?s environments.
The general concept of niche construction also provides the logical link between
current efforts to understand domestication being conducted at two disconnected
scales of analysis. At the level of individual plant and animal species, on one hand,
there recently have been significant advances in our knowledge of the what, when,
and where of domestication of an ever-increasing number of species worldwide.3
At the same time, large-scale regional or universal developmental models of the
transition to food production continue to be formulated. These incorporate a vari-
ety of ??macro-evolutionary?? causal variables that may account for why human
societies first domesticated plants and animals.4,5 This essay employs the general
concept of niche construction to address the intervening question of how, and to
connect these two scales of analysis by identifying the general behavioral context
within which human societies responded to ??macroevolutionary?? causal variables
and forged new human plant or animal relationships of domestication.
REGIONAL AND SPECIES-LEVEL
APPROACHES TO UNDERSTAND-
ING DOMESTICATION
The initial domestication of plants
and animals and the subsequent de-
velopment of agricultural economies
marks a major escalation in the niche-
construction potential of human soci-
eties. Domesticates provided humans
with a significant new means of re-
shaping biotic communities and natu-
ral landscapes more to their liking.
Eight to ten centers of domestication
are now recognized worldwide.3?10
These independent centers of domes-
tication exhibit considerable variation
in terms of climate, physical environ-
ment, biotic community composition,
and human developmental history. As
a result, they comprise a significant
comparative data set of independent,
parallel, regional-scale case-study sit-
uations with which not only to search
for similarities, but to consider alter-
native explanatory frameworks for
domestication. At the same time, dif-
ferent plants and animals were inde-
pendently domesticated in these world
regions, and when taken together,
provide an equally diverse set of com-
parative case-study opportunities with
which to consider domestication at
the species level.
Efforts to explain and understand
domestication carried out at these
two different scales of analysis, re-
gional and species-specific, also address
different aspects of what is increas-
ingly recognized as a long, complex,
and regionally quite variable co-evo-
lutionary process. At the regional
level and above, analyses focus on
characterization and consideration
of the relative importance of a vari-
ety of ??macroevolutionary?? variables,
among them climate change, popula-
tion growth, landscape packing and
hardening of between-group bounda-
ries, and intra and intergroup com-
petition for resources and social sta-
tus. Identified as potentially causal in
nature, these variables are cast in
roles of varying importance in frame-
works of explanation that address
the central question of why human
societies initially established and
subsequently sustained relationships
of domestication with some species.
At the species level of analysis, in
contrast, research focuses on docu-
menting a variety of different aspects
of where and when domestication of
different species occurred, and spe-
ISSUES
Bruce D. Smith is an archeologist inter-
ested in the initial domestication of plants
and animals worldwide and the associ-
ated transition from hunting and gathering
to food-production economies. He has
recently authored or coauthored articles
on this topic in Science, Proceedings of
the National Academy of Sciences,
Trends in Genetics, and Cell, and is co-
editor of the recent volume Documenting
Domestication: New Genetic and Ar-
chaeological Paradigms, University of
California Press (2006). A new edition of
Smith?s essays on early food production
economies in eastern North America, Riv-
ers of Change, is being published by the
University of Alabama Press (2006).
Key words: archeology; agricultural origins; food
production; ecosystem engineering; resource
management
VC 2007 Wiley-Liss, Inc.
DOI 10.1002/evan.20135
Published online in Wiley InterScience
(www.interscience.wiley.com).
Evolutionary Anthropology 16:188?199 (2007)
cifically what genetic and morpholog-
ical changes distinguish the newly
created domesticates from their wild
ancestors.
DOCUMENTING THE WHAT,
WHEN, AND WHERE OF
DOMESTICATION AT THE
SPECIES LEVEL
Identification of the likely wild
ancestors of domesticated species
has been accomplished by compar-
ing the genetic profile of domesti-
cates with that of the modern de-
scendant populations of their sus-
pected progenitors throughout the
present-day range of a species, re-
sulting in the identification of the
wild populations that provide the
closest genetic match.9?14 The pres-
ent-day distributions of such ??best-
match?? wild populations often can be
employed to infer the specific envi-
ronmental and spatial contexts in
which domestication of the species
in question occurred. In a growing
number of cases, the earliest directly
dated archeological evidence for do-
mestication of different species has
been recovered from sites located rel-
atively close to where the modern
best-match wild progenitor popula-
tions still exist, providing additional
evidence of both the spatial and tem-
poral context of initial domestica-
tion.15 Not all domesticates, of course,
are developed from single progenitor
populations. Efforts to unravel the
complex hybridiztion histories for a
range of domesticated taxa constitute
a rapidly expanding area of inquiry.16
The morphological changes that
show evidence of domestication in a
particular species can also provide
insights regarding the specific human
behavior patterns that produced them.
In seed-bearing plants, for example,
a number of distinct morphological
markers of domestication, such as
seed retention, uniform seed matura-
tion, terminal seed clusters, loss of
germination dormancy, and increased
seed size, have been identified as
automatically resulting from a spe-
cific set of human behaviors involved
in the intervention in the life cycle of
the plant populations in question?
the deliberate and sustained human
planting of stored seed stock.17
Any such morphological changes
associated with domestication are, of
course, the result of genetic changes
in the domesticates. Considerable
progress has been made in recent
years in gaining a better understand-
ing of such changes at the molecular
level. More than half a dozen ??do-
mestication genes?? have now been
identified that control many of the
major morphological and other changes
associated with domestication in
seed-bearing plants. These changes
include, for example, seed retention
in rice, seed compaction in wheat,
loss of branching and germination
dormancy in maize, and an increase
in fruit size in tomatoes.10,18 Only a
few of these genes that have been
linked to specific attributes associ-
ated with domestication have so far
been studied in archeological speci-
mens.19,20
Like the morphological changes I
have mentioned, genetic changes
associated with domestication in a
particular species are in turn the
result and evidence of a specific set of
human behaviors that comprise the
established and sustained relation-
ships of domestication, such as, for
seed plants, the sustained planting
and harvesting of stored seed stock.
Along with morphological and genetic
changes associated with domestica-
tion, evidence of the human behavior
patterns that comprise a relationship
of domestication with a particular
species are also sometimes directly
observable in the archeological re-
cord. Shifts in the age and sex compo-
sition of faunal assemblages, for
example, can provide a clear signature
of initial human herd management
and domestication.21 Pollen and phyto-
lith evidence of forest clearing, struc-
tural evidence of corrals or water man-
agement projects such as canals or
check dams, and changes in material
culture assemblages have also been
proposed as potential markers of the
existence of relationships of domestica-
tion between human societies and a
range of different species.17
Paralleling this progress in docu-
menting the details of domestication
at the species level, general explana-
tory approaches focusing on the pos-
sible reasons why human societies
first domesticated plants and animals
also continue to be developed at the
regional level or higher.
REGIONAL SCALE
MACROEVOLUTIONARY CAUSAL
EXPLANATIONS FOR WHY HUMAN
SOCIETIES DOMESTICATED
PLANTS AND ANIMALS
In the seven decades since V. Gor-
don Childe?s22 pioneering effort to
explain the ??Neolithic Revolution??
within a framework of human response
to a changing world, researchers
have continued to develop a wide
range of different overarching ex-
planatory frameworks for why humans
initially brought a variety of different
species under domestication.23 Some
of these explanations remain rela-
tively simple and straightforward,
and are proposed as being univer-
sally applicable. In looking outward
beyond the actual human behavior
patterns that comprised the newly
formed relationships of domestica-
tion in search of larger-scale causal
understanding, for example, recent
studies have outlined a universal ex-
planation for the transition from
hunting and gathering to food pro-
duction in terms of end-Pleistocene
climatic shifts. It has been argued
Along with
morphological and
genetic changes
associated with
domestication, evidence
of the human behavior
patterns that comprise a
relationship of
domestication with a
particular species are
also sometimes directly
observable in the
archeological record.
ISSUES Behavioral Context of Plant and Animal Domestication 189
that human domestication of plants
and animals was not possible before
the end of the Pleistocene at about
12,000 B.P. due to several key
aspects of Pleistocene climates. Low
rainfall over large areas of the globe,
as well as low CO2 levels, it is pro-
posed, restricted plant growth during
the late Pleistocene. Climates are
characterized as having been turbu-
lent, with high-amplitude fluctua-
tions on time scales of less than a
decade to a millennium.24,25 With
the onset of warmer, wetter climates
in the Holocene, however, accompa-
nied by an increase in CO2 levels,
and most importantly, the establish-
ment of much more stable and quies-
cent weather patterns, constraints
were lifted on human development
of viable, sustainable relationships
of domestication and agricultural
economies.
Few researchers would argue with
the general point that Pleistocene
climates may well have constrained
the development of relationships
of domestication. Considerably less
plausible, however, is the corollary
argument that early Holocene climates
were the central, universal, and imme-
diate cause of plant and animal
domestication: that the onset of stable
Holocene climates was the ??trigger??
that made subsequent domestication
and agriculture not only possible but
compulsory, with no alternatives or
other options.24,25 Improving Holocene
climates, it is proposed, produced
greater abundance of plant and animal
resources and an associated increase
in carrying capacity. Human popula-
tion levels, in turn, steadily increased
and the ensuing resource competition
between neighboring groups generated
a ??competitive ratchet?? favoring the or-
igin and diffusion of agriculture.25
At the opposite end of the spectrum
of proposed universal explanations,
Brian Hayden?s26 ??food fight?? theory of
domestication assigns a central causal
role not to an external environmental
variable such as Holocene climatic
amelioration, but to an internal social
motivating force. According to Hayden,
the worldwide domestication of all
plant and animal species can be
explained in terms of the increasing
demand for rare delicacies to enhance
competitive feasts hosted by different
kin-units within societies in an effort to
increase their relative social standing.
In stark contrast to such explana-
tory frameworks based on ??universal
inevitability,?? a rich variety of other
far more fine-grained and more
empirically based macro-evolution-
ary approaches have been proposed
at a regional scale of consideration.
These regional perspectives incorpo-
rate a range of different theoretical
perspectives, including human be-
havioral ecology, macroevolutionary
theory, and several processual
approaches.4,5,27?30 These regional-
scale models are also region-specific;
they focus on the interplay of cli-
matic, environmental, and social var-
iables within specific natural and cul-
tural-developmental contexts.
In a landmark study, for example,
Flannery31 offers a detailed and well-
supported model of cost-based shift-
ing resource use in the valley of
Oaxaca leading up to the first domes-
tication of a crop plant (Cucurbita
pepo) anywhere in the Americas.
Similarly, Piperno32 addresses the
initial domestication of crop plants
in the tropical deciduous forests of
the Neotropics within an exhaus-
tively documented context of post-
Pleistocene environmental change
and associated shifts in demography
and resource selection, arguing that
people began to cultivate some
plants as soon as the net return from
subsistence strategies involving plant
propagation exceeded those resulting
from full-time foraging. In a compa-
rable manner, a developmental sce-
nario has been proposed for initial
plant domestication in eastern North
America that considers the specific
attributes of the species in question,
as well as their economic roles and
habitat preferences in the lead-up to
their domestication.33
Various other similar explanatory
frameworks, each specifically tai-
lored to a particular world region,
have been developed in the last dec-
ade to address the question of why
human societies first domesticated
plants and animals, with the Near
East drawing the most interest
because it provides the most detailed
archeological record of the develop-
mental transition from hunting and
gathering to food production.34?37
In a recent comprehensive overview
of alternative macro-evolutionary
approaches for the Near East, Zeder5
argues for a nuanced fine-grained
framework of explanation incorpo-
rating macroevolutionary and proc-
essual approaches that also draws
on concepts of human agency, his-
torical contingency, and directed
variation.
Although these explanatory frame-
works for why human societies
established and maintained relation-
ships of domestication differ widely
in terms of the temporal precedence,
causal importance, and interrelated-
ness assigned to potential causal var-
iables, they all share a common em-
phasis on a region-specific approach
and the integration of relevant infor-
mation of many different kinds.
Rather than assessing the relative
strengths of these causal explana-
tions or adding to them, however, I
am interested here in considering
how they can be better linked to
available evidence regarding the spe-
cifics of the what, when, and where
of domestication.
NICHE CONSTRUCTION: THE
GENERAL INTERFACE BETWEEN
REGIONAL-SCALE
EXPLANATIONS AND
SPECIES-LEVEL STUDIES
Regional-scale explanatory frame-
works that consider the why of
domestication continue to be devel-
oped and refined for different world
areas. Substantial progress continues
to be made in documenting the what,
when, and where of domestication at
the species level. Relatively little
attention has been focused, however,
on the nature of the interface
between these two scales of analysis.
How do changes in the various re-
gional-scale environmental and cul-
tural variables actually influence
societies to domesticate some spe-
cies? How are such larger-scale ex-
planatory frameworks linked, at the
species level, to the actual initiation
of relationships of domestication by
humans? What is the nature of the
intersecting general behavioral con-
text within which human societies
190 Smith ISSUES
establish and sustain relationships of
domestication? The term ??niche con-
struction?? provides a recently well-
characterized and much-discussed
heading for consideration of this
larger behavioral domain within
which human domestication of
plants and animals emerged.
Niche construction is defined as
??organism-driven environmental mod-
ification?? and ??the activities of organ-
isms that bring about changes in
environment.??1 Not surprisingly, Homo
sapiens is acknowledged as being the
species with the most impressive re-
cord of niche construction. Odling-
Smee, Laland, and Feldman1 identify
humans as ??the ultimate niche con-
structors.?? They also provide abun-
dant examples of niche construction
by a wide range of other taxa, pro-
posing that niche construction plays
an important and almost ubiquitous,
if underappreciated, general role in
shaping environments and evolution-
ary processes worldwide. Through
niche construction or ecosystem en-
gineering many organisms reshape
both their own environments and
those of other organisms. By forming
complex ??engineering webs?? of inter-
action, they modify the natural selec-
tion pressures acting on a range of
other components of the biotic
community.2 To have any sort of ev-
olutionary effect, however, such
niche-construction efforts by humans
or other species must persist across
generations, either through genes or
??ecological inheritance?? or, in the
case of humans, by cultural trans-
mission. Human dependence on cul-
tural processes, however, does not
make human niche construction
unique: ??niche construction is a gen-
eral process exhibited by all organ-
isms??.1
This recognition of the existence of
niche construction by other species
and across all of the earth?s ecosys-
tems and biotic communities pro-
vides an important general founda-
tion and broad context for under-
standing niche construction as
carried out by those human popula-
tions that initially brought plants
and animals under domestication.
Much of human niche construction
can be included within the general
category of ecosystem engineering
activities carried out by nonhumans;
such activities include tool making,
selection and consumption of resour-
ces, storage of food and water, farm-
ing of food resources, production
of detritus, import and export of
nutrients, modification of chemical
environments, creation of paths, and
transport of other organisms.1 Given
that niche construction is not a
uniquely human attribute, but widely
documented among other species,
and that most if not all human niche
construction efforts can be grouped
under general niche construction cat-
egories developed for the animal
kingdom, it is reasonable to conclude
that the human societies involved in
the initial domestication of plants
and animals worldwide were actively
engineering their ecosystems in a va-
riety of different ways. Other than
cultural inheritance, what qualita-
tively distinguishes humans from
other species in terms of niche-con-
struction efforts is, of course, the di-
versity of the human repertoire of
ecosystem engineering and the far
greater potential impact of human
niche-construction efforts.
HUMAN NICHE CONSTRUCTION
AND DOMESTICATION
Against this general backdrop of
niche construction as an activity
exhibited by all organisms, human
efforts to restructure local biotic com-
munities have been documented
across a wide range of diverse envi-
ronmental zones worldwide. Over
the years, human efforts at shaping
their natural landscapes have been
classed under a number of terms,
including environmental manipula-
tion,38 indigenous management,39
aboriginal agronomy and domestica-
tion of environment,40 domesticated
landscapes,41,42 indigenous resource
management,43 and traditional resource
management.44 In addition, general
classification schemes for human eco-
system engineering have been devel-
oped.45,46 All of these roughly synony-
mous terms and characterizations fall
comfortably under the now far more
commonly used and more general
heading of niche construction.
Both ethnohistorical and present-
day descriptions of human niche
construction have usually focused on
particular types of human resource
management influencing wild resour-
ces or efforts focused on individual
species or species groups of nondo-
mesticates. Often, such descriptions
are embedded within the context of
general descriptions of overall sub-
sistence economies of both hunting
and gathering and food-producing
societies.
In contrast to studies that are
focused on particular human man-
agement activities such as burning,
or human intervention in the life
cycle of certain wild species, Kat
Anderson44 provides comprehensive
documentation of the full spectrum
of different land management prac-
tices employed by Indian societies in
what is now California over the past
10,000 years. She includes, for exam-
ple, discussions of the following cate-
gories of ecosystem engineering that
targeted plant populations: burning,
coppicing, pruning, sowing, weeding,
tilling, selective harvesting, trans-
planting, irrigating, harrowing, seed
scattering, and digging. As Anderson
underscores, all of these different
activities comprised an integrated
and coordinated traditional resource
management strategy of direct and
sustained manipulation of a broad
array of culturally significant popula-
tions of plants and animals and their
habitats in order to maintain and
heighten their abundance, productiv-
ity, and diversity.44 Such broad-spec-
trum strategies of human niche con-
struction are not unique to Califor-
nia. They were and, in some places,
still are an integral aspect of the
human condition, involving keen ob-
servation, patience, experimentation,
and long-term relationships with a
broad spectrum of plants and ani-
mals, and have been adjusted and
refined over many generations of ex-
perience and contact with the envi-
ronment.43,44 Anderson44 defines the
traditional resource management
strategies of Californian Indian pop-
ulations as a ??culturally mediated
relationship with the natural world
in which humans intervene in the
life cycle of native plants and ani-
mals to direct their growth and
ISSUES Behavioral Context of Plant and Animal Domestication 191
reproduction.?? This general defini-
tion of human resource management
or niche construction, not surpris-
ingly, comfortably encompasses most
definitions of domestication and sup-
ports the recognition of domestica-
tion as a particular form of niche
construction.17
In contrast to ethnohistorical and
present-day research on human re-
source management activities, ??the
empirical search for signatures of
past human cultural niche construc-
tion??1 remains a difficult challenge.47
Relatively few archeological indica-
tions of human management of non-
domesticated plants or animals have
been documented to date, and these
provide only temporally and geo-
graphically scattered clues regarding
the time depth and developmental
history of integrated systems of
human environmental management.
Occasionally, however, the archeo-
logical evidence of landscape man-
agement by human societies can be
impressive. In the Phoenix, Tonto,
and Tucson Basins of the Southwest
United States, for example, more
than 550 individual locations where
agave plants were transplanted by
Hohokam societies at ca. A.D. 600?
1350 have been documented.48?50
Individual agave plants were taken
from naturally occurring populations
and transplanted to locations near
stream or river-valley dwellings and
village sites, or into larger fields lo-
cated on suboptimal bajadas some
distance from dwellings (Fig. 1). Water
manipulation features, specifically
rainfall runoff collection features
and rock-pile complexes around indi-
vidual plants to increase soil mois-
ture, as well as discarded processing
tools, roasting pits, and relict agave
populations mark the location of
these cultivation sites.
In the same general time frame
(A.D. 600?1000) human societies in
the Owens Valley of eastern Califor-
nia were enriching and expanding
water-meadow habitats of various
wild bulbous hydrophytic plant spe-
cies through labor-intensive con-
struction of diversion dams and
feeder ditches.51,52 These archeologi-
cally visible niche-construction activ-
ities were sustained into the historic
period, when they were described in
some detail.
The archeological record of eastern
North America provides several less
compelling but still plausible examples
of human management of wild resour-
ces. A shift in the relative frequency of
gray squirrels versus fox squirrels in
Late Archaic period faunal assemb-
lages, for example, has been proposed
as evidence of the selective cutting of
trees in favor of nut- and mast-bearing
species.53 In addition, evidence of
range extension and a notable increase
in the relative abundance of three east-
ern North American seed-bearing
plants in Late Archaic and Early Wood-
land archeobotanical assemblages (ca.
3000?2000 B.P.) has been suggested as
indicating that they were cultivated and
of considerable economic importance,
even though they did not exhibit any
unequivocal morphological indications
of domestication.33 These three plants
were maygrass, Phalaris caroliniana;
erect knotweed, Polygonum erectum;
and little barley,Hordeum pusillum.
The Northwest Coast of North
America has also witnessed consider-
able recent interest in documentation
of various forms of human niche con-
struction, both present-day and pre-
contact. Studies have included, for
example, documentation of human-
mediated range extension and trans-
Figure 1. Agave management in the Southwest. Grid field features near Stafford Arizona
have yielded evidence for the management of transplanted Agave.
192 Smith ISSUES
planting of wild root crops, evidence
of ??ownership?? and weeding or cultiva-
tion of wild plant resources, and con-
trolled burning.43,47
One of the most frequently en-
countered archeological signatures of
environmental management is the
deliberate use of fire in controlled sea-
sonal burning of vegetation. Such con-
trolled burns, while a uniquely human
form of niche construction, can also be
considered a specific type of conserva-
tive, counteractive, stabilizing pertur-
bation as carried out by a wide range of
other species, in that it involves revers-
ing or neutralizing a prior change in
the environment in order to maintain a
preferred ecosystem state.1 Scheduled
burning has been documented histori-
cally as an important niche-construc-
tion effort employed by human soci-
eties in many world areas and has been
recognized or suspected in the archeo-
logical record of areas as diverse as
Australia,40 eastern North America,54
the Neotropics,55 and the west coast of
North America.43,44 Controlled burning
of vegetation by human societies
appears to extend as far back as the be-
ginning of the Holocene in the Ameri-
cas, perhaps to 30,000 B.P. in Island
Southeast Asia,9 and to 55,000 B.P. in
South Africa.56
DOMESTICATES PROVIDE
EVIDENCE OF NICHE
CONSTRUCTION
Occasionally, human niche-con-
struction efforts in the distant past
developed into sustained relation-
ships of domestication with particu-
lar species. Various aspects of the ar-
cheological record of domestication
worldwide in fact provide the best in-
dication of the existence of compre-
hensive human strategies for the
overall management of ancient biotic
communities. Significantly, and not
surprisingly, when considered within
the broader context of proposed
ubiquitous human niche-construc-
tion activities, the development of
such relationships of domestication
was not limited to a particular place
or a narrow time frame. Each of the
eight to ten environmentally and cul-
turally diverse world regions cur-
rently identified as likely independent
centers of domestication and agricul-
tural origin exhibits a unique multi-
ple-millennia sequence of domestica-
tion of different species.3?10
Further evidence in support of the
proposition that initial domestication
occurred within broad strategies of
resource management is provided by
the diversity of human behavior pat-
terns included under the general
heading of ??domestication.?? For each
of the hundreds of different species
of plants and animals brought under
domestication at widely scattered
times and places around the world,
human societies had to develop a
unique, species-specific set of new
behaviors. As a result, while taxa
with similar attributes (for example,
annual grasses versus tree crops ver-
sus caprines) could be expected to
have responded in similar ways to
generally similar forms of human
intervention in their life cycle,
??domestication?? as a general cate-
gory of human activity encompasses
a remarkable spectrum of different
causal human behavior patterns. A
brief consideration of ten of the ear-
liest domesticates in different world
regions can provide a clear demon-
stration of this diversity.
The domestic dog (Canis famili-
aris) and bottle gourd (Lagenaria
siceraria) comprise an unlikely but
informative pair in this regard. They
are thought to be the first two spe-
cies brought under domestication,
between 12,000 and 15,000 years
ago, somewhere in Asia, and to have
likely colonized the Americas in the
company of Paleoindian popula-
tions.57 Both were primarily of utili-
tarian value, dogs for hunting and
gourds as containers, rather than
food sources. The initial formation
of a relationship of domestication
with the two has often been charac-
terized as not so much involving uni-
lateral intervention on the part of
humans as it was dogs and bottle
gourds being welcomed to colonize
and share the human niche.58
Often described as ??camp fol-
lowers?? and ??dump heap?? plants,
bottle gourds are active pioneers of
disturbed soil situations, and do not
require much care or attention once
introduced into a disturbed human
habitat. The initial phase of domesti-
cation of the bottle gourd may have
been marked by human harvesting of
wild gourds and the subsequent dis-
card of seeds along with other refuse
near habitation sites, establishing
them as refuse area colonizers and
fellow travelers, with subsequent de-
liberate human selection resulting in
thicker rind and larger fruits.
Human refuse may also have
played a role in initially attracting
dogs to come into closer contact with
humans. Movement by canines into
the ??human niche?? can also be
associated with their automatic self-
selection for reduced aggression,
thereby facilitating the development
of a relationship of domestication.
Reduced aggression in dogs as they
occupy the human niche is reflected
in linked morphological markers
such as snout shortening, as well as
crowding and size reduction in
teeth.59,60
As was the case with the dog and
bottle gourd, the initial domestica-
tion of the fig (Ficus carica) in the
Near East by 11,400 years ago also
involved a relatively limited level of
intervention by human societies in
the life cycle of the species in ques-
tion: the simple cutting and planting
of branches from fig trees.61 This
simple human action, however, reflected
a sophisticated biological knowledge
of their environment, since the fig
trees that were selected for such veg-
etative cloning were parthenocarpic,
For each of the
hundreds of different
species of plants and
animals brought under
domestication at widely
scattered times and
places around the
world, human societies
had to develop a
unique, species-specific
set of new behaviors.
ISSUES Behavioral Context of Plant and Animal Domestication 193
and while they produced soft sweet
fruit, they did not set germinative
seeds. As a result, they would have
been reproductive dead-ends unless
humans intervened by deliberately
propagating their shoots.61?63 As was
the case with agave, such replanting
of fig branches could also have taken
place close to human habitation
sites, expanding the niche of the en-
couraged species while making each
year?s fig crop easier to monitor and
harvest. In addition, since such
planted shoots could not be expected
to yield fruit for some years, humans
had clearly accepted the delayed re-
ward associated with a slow-matur-
ing crop.
Like the fig, the banana Musa sp.,
another relatively long-lived and
delayed-yield tree crop, was the sub-
ject of early human management as
part an overall strategy of landscape
modification, but with a substan-
tially greater level of human invest-
ment of labor. The archeological site
of Kuk Swamp, situated at an eleva-
tion of 1,560 meters above sea level
in highland New Guinea, has yielded
Musa phytoliths thought to reflect
experimental cultivation of indige-
nous plants by 10,000 B.P., followed
by the construction of increasingly
substantial water management
canals associated with the cultiva-
tion of bananas and other crops af-
ter 7,000 B.P.9 Kuk Swamp, in
island Southeast Asia, thus provides
very early evidence of the apparent
relocation and transplanting of a
tree crop outside of its natural
range, along with substantial human
investment in sustained manage-
ment paired with a clear awareness
of delayed return.
Although the goat (Capra hircus) is
far different from figs and bananas
in most respects, and required quite
different efforts on the part of
humans in the initial creation of a
relationship of domestication, it
again reflects both a sophisticated
human understanding of its manage-
ment requirements and the acknowl-
edgment of delayed return linked to
a substantial commitment to sus-
tained management. By 10,000 years
ago, human societies in the Zagros
Mountains of present-day Iran had
intervened to a significant extent in
the life cycle of goat herds, and had
taken control of their reproduction.
As is the case with other domesti-
cated livestock species worldwide,
humans selectively culled immature
males, restructuring breeding popu-
lations into the distinctive general
age and sex profile that is the hall-
mark of present-day livestock man-
agement for meat production: a sub-
stantial majority of adult breeding
females are kept along with a few
adult males.21
In the Americas, two species of
squash provide the earliest evidence
of domestication of food crops. In
Mexico, an increase in seed size
marks the initial domestication of
Cucurbita pepo at ca. 10,000 B.P.,64,65
while in Ecuador, at about the same
time, an increase in the size of rind
phytoliths, reflecting an increase in
fruit size, indicates the deliberate
human cultivation of Cucurbita ecua-
dorensis.66 As is the case for seed-
bearing plants worldwide, among
them barley, maize, millets, rice, sor-
ghum sunflower, and wheat, the spe-
cific set of human behaviors involved
in the initial domestication of these
two squashes centered on the sus-
tained planting of stored seed stock
in prepared planting areas.
Plants with starch-rich under-
ground organs, such as manioc
(Manihot esculenta), arrowroot (Mar-
anta arundinacea), and leren (Cala-
thea allouia), were also domesticated
very early in South America, based
on the recovery of diagnostic root
crop starch grains and phytoliths
from 9,000 to 8,000 B.P. con-
texts.32,67 As with other species in
the general category of root crops,
initial human cultivation of manioc,
leren, and arrowroot would have
involved the replanting of clonal
fragments of parent plants in concert
with sustained cultivation and delib-
erate selection for desired attributes
such as larger tubers and preferred
starch types.
Interestingly, at the same time that
recent research in different world
regions has documented an increas-
ingly diverse range of human behav-
ior patterns associated with success-
ful early domestication of plants and
animals, there has also been substan-
tial recent evidence of early efforts
that were not sustained over the long
term. Initial efforts to domesticate
barley (Hordeum spontaneum), oats
(Avena sterilis), and rye (Secale
cerale), for example, dating from
before 11,000 B.P. (and the end-
Pleistocene climatic shift?) have been
recognized in the Near East.68 Each
of these early efforts at plant man-
agement and manipulation was
abandoned, however. Sustained culti-
vation of the species in question did
not occur until several thousand
years later in different locales. In the
Americas a species of runner bean
(Phaseolus sp.) has similarly been
identified as a possible early, ca.
9,000 B.P., subject of human efforts
at cultivation and management that
were not sustained.32 It has also
recently been suggested that the sun-
flower (Helianthus annuus) may have
been domesticated twice, with the
domestication of wild populations in
eastern North America at ca 4,400
B.P. being sustained and leading to
all documented present-day cultivar
varieties; a second, roughly contem-
poraneous and independent domesti-
cation of the sunflower in Mexico
was subsequently abandoned.33 In
eastern North America, three indige-
nous seed plants, maygrass (Phalaris
caroliniana), erect knotweed (Poly-
gonum erectum), and little barley
(Hordeum pusillum), have also been
identified as likely having been culti-
vated and of considerable economic
importance between 3,000 and 2,000
B.P., even though they do not exhibit
any unequivocal morphological indi-
cations of domestication and subse-
quently disappeared from cultiva-
tion.33
This brief consideration of the
diverse array of human activities
involved in the initial domestication
of plants and animals, both sus-
tained and short-lived, when com-
bined with other lines of evidence
discussed, provides considerable sup-
port for the proposition that plants
and animals were initially domesti-
cated within a broad behavioral con-
text of human niche construction.
What then, does placing human
domestication within such a broad
behavioral context add to our overall
understanding of this major transi-
tion in human history?
194 Smith ISSUES
DISCUSSION
Often described under a range of
different terms, including environ-
mental manipulation, indigenous
management, domestication of the
environment, and traditional re-
source management, niche construc-
tion is the broad strategy employed
by humans to shape, enhance, and
sustain their ??natural?? world while
also expanding their developmental
horizons. More than 40 years ago,
James Downs27 noted the substantial
variation that existed among differ-
ent small kin groups of Great Basin
hunter-gatherers in the details of
how they managed and manipulated
their environment. He identified two
factors, variation and manipulation,
as central to the general evolutionary
potential of human societies. More
recently, Odling-Smee, Laland, and
Feldman1 have argued that environ-
mental engineering or niche con-
struction is a major, if widely under-
appreciated general factor in the
overall shaping of biotic commun-
ities and evolutionary processes.
Here I argue that niche construction
provides an important evolutionary
and behavioral context for under-
standing one of the major transfor-
mations in the history of our species,
the initial domestication of plants
and animals.
As outlined earlier, observations
and arguments in support of the
proposition that the human societies
that initially domesticated plants and
animals worldwide did so within a
preexisting broad context of environ-
mental engineering can be organized
under at least eight different catego-
ries of general observations and
arguments. First, niche construction
by a large number of other animal
species has been documented across
all of the earth?s ecosystems and bi-
otic communities. Given that so
many different animal species
manipulate their environments, it is
reasonable to assume that human
populations have been actively man-
aging environments to varying
degrees for tens of thousands of
years. Second, It is also reasonable
to assume that such past patterns of
human resource management were
both sophisticated and constantly
being refined. Humans, after all, are
acknowledged to be the ultimate
niche constructors, both in terms of
the diversity of different ways in
which we manipulate the world
around us and the magnitude of our
resultant impacts. Third, ethohistori-
cal and present-day studies have
documented a growing inventory of
the different ways in which human
societies actively intervene in their
local environments in an effort to
shape them more to their liking.
Fourth, recent research is beginning
to provide much more fine-grained
and comprehensive understanding of
the overall coordinated management
strategies employed by human soci-
eties, strategies that embrace many
different forms of environmental
manipulation and involve a variety of
different organic and inorganic eco-
system components. Fifth, archeolo-
gists and archeobiologists continue
to look for and find increasing evi-
dence of a range of forms of human
niche construction in the archeologi-
cal record, extending across many of
the world?s terrestrial ecosystems.
Sixth, the widely dispersed and iso-
lated record of domestication of dif-
ferent species of plants and animals,
globally scattered in time and space,
suggests the existence of an underly-
ing shared general pattern of human
behavior. Seventh, domestication of
hundreds of species, with different
species-specific management or ma-
nipulation challenges, called for a
wide variety of human behavior sets.
Eighth, increasing documentation of
multiple efforts to establish relation-
ships of domestication, which some-
times were not sustained, under-
scores the extent to which human
societies, with a diverse array of
skills, were consistently experiment-
ing across a broad range of species
in an effort to enhance their environ-
ment. Based on these supporting
arguments and observations, a plau-
sible case can be presented that do-
mestication quite likely occurred
within integrated strategies of eco-
system engineering based on a com-
prehensive storehouse of knowledge
about local biotic communities that
had been acquired over hundreds, if
not thousands, of years of direct ex-
perience.43,44
How then, does this recognition
that relationships of domestication
were forged within a broad behavio-
rial context of coherent environmen-
tal management strategies add to our
understanding of this major develop-
mental episode? First, and perhaps
most importantly, by situating do-
mestication within a broad and well-
documented category of general
human behavior we remove, to a
considerable extent, the proximate
mystery surrounding exactly how
domestication was accomplished.
Whatever the exact nature and role
of the particular macroevolutionary
causal factors that may have been in
play in independent centers of domes-
tication, human societies responded
not in terms of spontaneous and
anomalous acts of isolated inventive
genius, but rather within a coherent
and broadly based approach to man-
aging and enhancing their environ-
ment. Domestication was not the
product of unusual ??outside the en-
velope?? behavior patterns, but em-
Whatever the exact
nature and role of the
particular
macroevolutionary
causal factors that may
have been in play in
independent centers of
domestication, human
societies responded not
in terms of spontaneous
and anomalous acts of
isolated inventive
genius, but rather within
a coherent and broadly
based approach to
managing and
enhancing their
environment.
ISSUES Behavioral Context of Plant and Animal Domestication 195
erged out of coherent preexisting
resource management systems. Mac-
roevolutionary forces did not directly
cause domestication, but rather
resulted in human societies intensify-
ing their niche-construction efforts.
The human societies that initially
domesticated different species in dif-
ferent world regions, I argue, were
involved in the active, ongoing
manipulation of many species in a
variety of ways. As macroevolution-
ary variables caused overall expan-
sion and intensification of efforts at
resource manipulation, a few of the
varied efforts at manipulation of dif-
ferent plants and animals produced
an advantageous enhancement of
abundance and productivity far be-
yond what was achieved with the
vast majority of other species being
managed. Niche construction thus
provides a general universal context
for domestication that is, in fact, rel-
evant across a wide range of differ-
ent environmental zones and within
a broadly diverse set of cultural de-
velopmental trajectories. Even though
human societies in very different
ecosystem settings, from south China
to the highlands of South America,
independently domesticated a wide
range of different species of plants
and animals at different times and in
different sequences, they all share a
common behavioral strategy of niche
construction. Recognition of this broad
context of human ecosystem engi-
neering also underscores the general
manner in which human societies
identified and targeted certain spe-
cies and species groups for further
intervention and manipulation and,
in the process, highlights several
interesting related issues that may
represent promising future areas of
research.
When initially established, for
example, relationships of domestica-
tion cannot be assumed to have dif-
fered from other forms of niche con-
struction either qualitatively, for
example in terms of inherent intel-
lectual complexity, or in terms of
required human labor investment.
The specific human behavior pat-
terns that comprised relationships of
domestication with managed species
were not necessarily either more
challenging or more labor intensive
initially than were other niche-con-
struction activities carried out by
human populations. Within such
long-term sustained strategies of eco-
system engineering, human societies
carried out ??trial and error,??1 ??tin-
kering,??18 or ??experimentation??31 that
involved repeated auditioning of a
wide range of species with a constant
stream of different forms of manage-
ment in an effort to identify new and
better ways of shaping and enhanc-
ing their niche. Most such ecosystem
engineering by human societies, and
most species they manipulated, had
inherent limitations, and did not
hold the promise of open-ended
expansion or lead to impressive
returns, even though they were inte-
gral components of successful, sus-
tainable, long-term human adapta-
tions to local environments. Placing
those particular behavior patterns
that produced domesticates within
the broader comparative context pro-
vided by other parallel human efforts
at landscape management may offer
clues to other relevant aspects of the
process by which humans shifted to a
greater economic reliance on domes-
ticates. Ongoing consideration of why
some species went on to have long
and illustrious careers as domesti-
cates while others did not3,4,6,33 holds
the promise of continuing to provide
substantial new insights regarding
the particular attributes and profiles
of preadaptation that humans were
able to recognize during broad-scale
niche-construction auditioning for
potential star performers.
Situated at this interface of macro-
evolutionary causation on the one
side and the formation of a multi-
tude of species-specific relationships
of domestication on the other,
human niche construction also pro-
vides a vantage point for speculation
on two other major and widely rec-
ognized aspects of the initial domes-
tication of plants and animals: the
extent to which the ??broad-scale rev-
olution?? may reflect intensified
resource management by human
societies and the potential applica-
tion of niche-construction theory in
gaining a better understanding of the
apparent ??rich resource zone?? con-
text of domestication of plants and
animals worldwide.
Establishing the structure, com-
plexity, and impact of human niche
construction strategies back into the
Pleistocene remains a challenging
and potentially rewarding area of in-
quiry. The very early domestication
of utilitarian species such as the dog
and bottle gourd, along with tanta-
lizing evidence of possible deliberate
burning of vegetation, provide most
of the admittedly limited direct evi-
dence that human societies were
active ecosystem engineers well back
into the Pleistocene and likely had
the potential for establishing rela-
tionships of domestication.
Interestingly, both the dog and
bottle gourd fall toward the lower
end of the scale of required human
intervention in establishing and sus-
taining such new relationships. As
the search for evidence of niche con-
struction is extended back into the
Pleistocene in a more concerted
manner, it will be interesting to see
to what extent the ??broad spectrum
revolution??69 may be associated with
intensification of human efforts at
resource management. Reflecting a
dramatic increase in the range of
species being exploited by human
societies and documented in many
world regions as overlapping with
and immediately preceding domesti-
cation, the broad-scale revolution is
often identified as evidence of a
growing human population, with
resource imbalance that necessitated
increased human reliance on higher
cost, lower yield resources.37 This
dramatic increase in the number and
diversity of species, whatever its
cause, may also reflect an associated
expansion and intensification of
human niche construction activities.
At the same time that human soci-
eties were working their way down
the preferred species list and increas-
ing their diversity index of exploited
species, they could well have also
been expanding and intensifying
their ecosystem engineering efforts.
It will be interesting to see to what
extent new markers of human
resource management may be hiding
in plain sight within the increasingly
well documented developmental pat-
terns of the broad-scale revolution.
Niche-construction theory also
provides an interesting new perspec-
196 Smith ISSUES
tive on why a variety of the world?s
first food crops appear to have been
domesticated in relatively similar
habitats. Species that were initially
domesticated for their food value,
rather than utilitarian uses, and
which involved more capital invest-
ment in terms of human effort in
intervention, first appear as domesti-
cates in very similar environmental
settings in many of the independent
centers of domestication so far iden-
tified worldwide. In the Yangtze
River Valley corridor in China, for
example, as well as in the Near East,
Sub-Saharan Africa, eastern North
America, and Mexico, very early evi-
dence domestication of a range of
different species comes from settle-
ments situated in very rich resource
zones associated with river valleys,
lake margins, and springs.6,33,70 In
none of these areas did relationships
of domestication appear to have de-
veloped within a ??necessity is the
mother of invention?? context as pop-
ulation growth forced humans into
marginal environmental zones.69,70
Rather, these relationships developed
within rich resource situations that
enabled expanded and sustained
human experimental intervention in
the life cycle of a broad spectrum of
different species and the adoption
of a sedentary, logistically based
economy.
This rich environmental context
for initial human domestication of
plants and animals fits comfortably
within the expectations of niche-con-
struction theory.1 Local habitat set-
tings that were rich in biotic resour-
ces (species abundance and diversity,
as well as species with high biotic
potential) would have provided the
greatest opportunity for human
societies to expand and enrich their
overall integrated resource-manage-
ment strategies. The greater the
range of species included in human
efforts at intervention auditions, and
the wider the range of different
potential forms of intervention that
could be attempted, the greater the
likelihood that relationships of do-
mestication would have been suc-
cessful and sustained.
According to niche-construction
theory, within the context of post-Pleis-
tocene climatic improvement and the
associated increase in resource gra-
dients within spatially heterogeneous
environments (increasingly nonuni-
form distribution of resources), the
richer resource zones, those that
exhibited a greater capacity for sup-
porting more people in more perma-
nent settlements, could be expected to
have witnessed stronger sustained
niche-construction efforts. Trajecto-
ries of change would have been ??chan-
neled toward adaptation to those
regions of niche space in which abun-
dance is greatest, rather than to other
regions.??1 Similarly, with increased
sedentism and sustained niche con-
struction over many generations, the
impact and potency of human ecosys-
tem engineering, with the sheer per-
sistent repetition of the same niche-
construction activities, ??might act like
a persistent unidirectional ?pump,???1
directing trajectories of human devel-
opment toward greater investment in
and dependence on intensified niche-
construction activities. While it is, of
course, dangerous to place too much
reliance on ??unidirectional pumps??
and ??channeled?? adaptation, any
more than on climatic ??triggers,??
niche-construction theory can be con-
sidered to provide additional concep-
tual insight regarding why domestica-
tion appears to have occurred within
rich resource zones in many different
world regions.
The apparent shared environmen-
tal setting of initial domestication of
food crops across a wide range of
ecosystems worldwide provides a
third common aspect of this major
developmental shift in human his-
tory. Human domestication of crop
plants and livestock species appears
to have occurred in many different
world regions, in resource-rich envi-
ronmental zones, within a broad
general behavioral context of niche-
construction strategies and nested in
an associated broad-scale revolution
in subsistence economies.
In conclusion, while not meant to
represent any sort of overarching ex-
planatory framework for domestica-
tion, these proposed common as-
pects of human domestication of
plants and animals worldwide, when
taken together, do provide new ways
of looking at one part of this com-
plex process: the environmental and
behavioral interface between macroe-
volutional causal factors and the
establishment of new relationships of
domestication. This discussion also
helps focus attention on many cen-
tral and still-open questions regard-
ing the relative importance and ori-
gin of environmental or cultural cir-
cumscription in the initial domestication
of plants and animals. The resource-
rich river valleys and lake or spring
margins that witnessed much of the
early domestication of plants and
animals existed within regional mo-
saics of habitats of varying biotic
abundance, with environmental gra-
dients forming natural clinal boun-
daries. Within the rich resource-
zone pieces of the mosaic that could
support greater residential stability
and ??logistical?? subsistence strat-
egies, it is still difficult to establish
exactly why stronger and less per-
meable cultural territorial bounda-
ries may have been established
around the resource catchment
areas of different communities. Was
the apparent formation of strength-
ened and sustained cultural bounda-
ries in such resource-rich zones the
result of population growth, peer
polity packing, and intergroup com-
petition for resources? Or is it possi-
ble that such ??affluent?? and ??logisti-
cally?? organized communities more
firmly defined their own territorial
boundaries as an aspect of an
ongoing process of investment in,
enhancement of, and conceptual
attachment to an ever-improving
anthropogenic landscape?
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VC 2007 Wiley-Liss, Inc.
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ISSUES Behavioral Context of Plant and Animal Domestication 199