A Global Dynamic Model For The Neolithic Transition (Kai W.wirtz, C.lemmen, ICBM 2002)

A Global Dynamic Model for the Neolithic Transition
Kai W. Wirtz (wirtz@icbm.de)

and Carsten Lemmen (c.lemmen@fz-juelich.de)
Institute for Chemistry und Biology of the Marine Environment (ICBM),
University of Oldenburg, Germany
Phone: +49-441-798-3778, Fax:+49-441-798-3404
Institut f
ur Chemie und Dynamik der Geosph
are, Forschungszentrum J
ulich,
Germany
Abstract. During the Holocene strong gradients in the distribution of technology
including subsistence ways emerged on a global scale. These patterns were further
amplified in historic times and are still visible through worldwide differences in
national wealth. In order to evaluate major factors responsible for the shift from
foraging to food production we here employ quantitative methods by developing a
deterministic but simple model. After compiling existing maps of potential vegetation at 5 000 BP the inhabited world is split into 197 regions with homogeneous
environmental conditions. Suitable variables for the macro-economic and cultural
development in the Neolithic period are found to be farming to hunting-gathering
ratio, number of agricultural economies and a technological development index. The
model explicitly describes economic adaptation, growth and migration of human
populations together with the spread of their cultural characteristics; it accounts for
over-exploitation of natural resources, crowding mortality and the climate variability
on a millennium scale. In a thorough model validation region specific trajectories
are compared to archaeological evidence revealing a high correspondence. Major
parts of the known sequence of Neolithic centers including the timing differences
are robustly reproduced. A series of known problems in prehistory is discussed
comprising the lag between domestication and full scale farming, the off-leveling of
the technological boost following the transition, the emergence of distinct migration
waves and sensitivity to climate fluctuations. Not mere population pressure but
continuous innovation and competition between subsistence strategies is identified
as a prime mover of agricultural development. The results suggest that few aspects
of biogeography may have determined the observed continental gradients in the
number of domesticable species ultimately leading to an increasing differentiation
in technology and demography.
c 2002 Kluwer Academic Publishers. Printed in the Netherlands.

wirtz_3rd.tex; 20/12/2002; 14:13; p.1
K.W. Wirtz and C. Lemmen
1. Introduction
The development of gradients in subsistence intensity between different
world regions after 12 000 years before present (yr BP) can be regarded
as causative for the divergence of subsequent historical pathways. Inherent in this evolvement is the onset of agriculture, which Childe (1928)
termed the Neolithic Revolution. Environmental changes might have
triggered the independent emergence of agriculture accompanied by a
wave of technological advance in at least five places. A chronology of
these events can be found in, e.g., Smith, Harris, Blumler (1998, 1996,
1992), but the sparsity of data does not allow for a complete picture.
An overview of conceptual models explaining the rise of agricultural food production is given by Wright (1992). According to the
most prominent hypothesis there exists a strong linkage with population pressure emerging at the beginning of the Holocene. However,
the questions of how, where and when agriculture rose is not simply of historical interest: The Neolithic shift defines one of the most
effective large-scale human transformations of the biosphere in the
past (Goudie, 1998; Louwe Kooijmans, 2001). Its study is therefore of
high relevance to understanding the revolution in human-environment
interaction modes currently in progress.
Considerable scientific effort during the last decade was directed to
the assessment of human impacts on global climate (IPCC, 2001). It
is, in particular, a primer task of climate change modeling to embed
the anthroposphere into the worlds physical, chemical and ecological
systems (Kane and Yohe, 2000; Ramankutty and Foley, 1998). First
steps toward transdisciplinary methods were undertaken by image, an
integrated assessment model of climate change (Alcamo et al., 1996,
1998). But the integration of human-environment interaction modes
is still lopsided in image or comparable assessment schemes like the
one of Prinn et al. (1999). These models prescribe the distribution of
regional industrial economies rather than simulate their adaptation to
changing climate conditions. Facing such a limitation, simulation tools
able to deal with cultural or economic change in relation to endogenous
dynamics and environmental variability are in strong demand in Earth
System Science.
wirtz_3rd.tex; 20/12/2002; 14:13; p.2
On smaller spatial scales models for the adaptation of human populations are under development. Therein, the method of agent-based
modeling provides the basic starting point; Dean et al. (2000) presented
one of the most forthcoming studies in this regard when simulating
Anasazi household movements over several centuries in a river valley.
However, we doubt that agent-based modeling can be as successful on
larger spatial and temporal scales. One reason is its dependency on fixed
rules which undermine the dynamic nature of long-term innovation
in technology and economy. Secondly, when resolving environmental
gradients on the globe, any agent-based simulation requires a numeric
effort far from being applicable at present.
It should be clear that human-environment interactions work in
both directions; environmental impacts on human development are
recently brought into sharper focus (Binford, 2001). Two groups of
factors, stationary as well as dynamic ones, are generally considered
separately. Static biogeographic determinants of cultural development
were made popular in the influential book by Diamond (1999). Actual
differences in the wealth of regional civilizations are traced back to
continental topology. In the second group of studies, implications of
fluctuating climate for human development during the Holocene are
sought (deMenocal, 2001; Olsson and Hibbs, 2000; Hs
u, 2000). Their
main methodology is to detect synchronizations between environmental hazards and fall of single civilizations, eventually paralleled by
population migrations.
Historical details in these studies are treated under raw working
assumptions which inevitably provokes criticism among historians and
makes thorough verification nearly impossible. By reducing mosaics of
historic contingencies to few relationships, the conceptual approaches
nevertheless may serve as a basis for developing a future generation of
quantitative methods. These implicate new perspectives in coping with
and understanding better the long-term rise of new human interaction
modes such as the Holocene subsistence intensification.
In this work we present a first attempt for such a highly aggregated
and quantitative description. Past environmental boundary conditions
(Section 3.1) are lumped to few variables on regional and continental scales (Sections 2.52.6). Their changes reflect imposed long-term
wirtz_3rd.tex; 20/12/2002; 14:13; p.3
fluctuations in climate (Section 3.3) but also feedbacks due to human

population growth and resource use (Sections 2.12.4). In Section 2.7
we describe how migration and trade patterns relate to changes in
climate, local environment and population structure. The prime goal
is to test to what extent the temporal macro-economic, cultural and
migration patterns can be reproduced as adaptations to the local environment. From this, various regional phenomena reconstructed from
archaeological evidence can be interpreted afresh in reference to few
mathematical and globally acting relationships.
2. Model description
2.1. Effective variables for the Neolithic transition
The post Ice Age transition is characterized by changes in subsistence
intensity (SI). Subsistence intensity describes a communitys effectiveness in generating consumable food and secondary products, for example bones, cotton and leather; this can be achieved based on an
agricultural-pastoral (AP) or a hunting-gathering (HG) life style or a
combination thereof.
In order to quantify an altered allocation pattern exhibited by a
group of individuals or whole populations in spending their energy,
time or manpower to one of the two activities we introduce the relative
proportion Q of effort put into AP food production; a similar variable
was used in the early attempt of Cooke and Renfrew (1979) to simulate
cultural development in ancient Greece.
The return of both competing strategies farming and foraging can
be increased by a set of technological features allowing for a net intensification of resource use. Examples in this context are tools like
arrows or plows and communication or storage abilities like writing
and architecture. In part, societal organization belongs to this feature
set, since it can promote SI by division of labor, for example. We introduce an aggregated variable T representing the mean technological
efficiency relative to its level in the late Paleolithic. This dimensionless
variable equals unity at simulation start by definition, while higher
wirtz_3rd.tex; 20/12/2002; 14:13; p.4
values implicate increased subsistence effectiveness made possible by

the invention and application of new technological and organizational
facilities.
Since T includes only non-biological aspects of energy procurement,
the domestication process as a key element in the Neolithic transition
has still to be encompassed. For this we define the number of potential
agro-pastoral economies (PAE), which are represented, for example, by
grassland pastoralism, cereal cultivation or fruit plantation. A value of
PAE around unity denotes a subsistence way able to sustain a population of agro-pastoralists. Thus, also the caloric efficiency of specific
cultivation or herding styles is implicitly incorporated. Because PAE
corresponds to the richness in domesticable and energy-rich animal or
plant species within a given environment it is not affected by human
actions beside imports as explained later.
The population intrinsic process of domestication itself is followed
by a third effective variable F . The new measure soundly relates to the
number of important species actually brought under domestication by
a local population. More precisely, F is defined as a fractional number
(0 F 1) with regard to PAE such that F times PAE gives the
number of actual agro-pastoral economies (AAE).
AAE = F PAE
(1)
Summarizing, the model considers three effective variables Q, T and

F (see also Table III). These are time variable characteristics of local
populations and aggregate a multidimensional developing stage with
respect to SI. Further important elements of the Neolithic transition
are exploitation of natural resources and diffusion of the developed
achievements. The assessment of these processes requires the introduction of population density P . Gradients in P will, for example, promote
migration in traits (Section 2.7). Success or failure of different food
supplying techniques can be indicated by rising or shrinking population
densities, which in turn may generate feedbacks on local conditions as
outlined in Section 2.4.
wirtz_3rd.tex; 20/12/2002; 14:13; p.5
2.2. Subsistence intensity

In our study, subsistence intensity is dimensionless and appropriately
scaled such that a value of SI = 1 expresses the mean subsistence
intensity of a hunter-gatherer society equipped with tools typical for
that period living in an affluent natural environment. The agro-pastoral
contribution to SI increases linearly with AAE; the more economies
there are, the better are sub-regional scale niches like mountains (e.g.,
for goat herding) or alluvial soils (e.g., for wheat cultivation) utilized.
The coupling of diverse cultivation practices also ensures minimization
of risk under variable annual climates. Finally, the simple linear approach roughly accounts for the synergistic effect of merging different
cultivation practices. An example for agricultural synergy is the surplus
achieved via feeding corralled animals with crop by-products.
The effect of technology as conceptually defined in Section 2.1 on
the return of calories and secondary products is more subtle. Consider,
for example, prevailing humidity-limitation so that the farming contribution in SI can be significantly enlarged by irrigation. If, however,
cold temperatures are constraining vegetative growth, technology can
only moderately promote agricultural productivity. So let TLI be an
index for the absence of temperature limitation or, equivalently, the
susceptibility of farming to technological advances. TLI equals unity in
warm climates and approaches zero at permafrost conditions. Following
the original definition of T and Q, total SI sums over a HG and AP
contribution:
SI = (1 Q) T + Q (TLI T ) (PAE F )
(2)
While a variety of techniques is able to remarkably increase harvests
of domesticated species, analogous benefits for foraging productivity are
less pronounced. Farmed land produces 10 to 100 times more calories
per area than does foraging which to a large extent can be attributed to
technological advances (Geisler, 1988). Apart from storing or hunting
facilities, already the limited availability of natural resources makes it
reasonable to assume a less than linear dependence of the HG calorie
procurement on T . We use a square root formulation with the desired
behavior T < T since T is generally larger than unity. Alternative
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functions could be sought; however, tests have indicated that these left
main model results unaffected.
2.3. Evolution equations
A fundamental property of living systems consists in the ability to adaptively react to environmental changes. It was recently demonstrated
that such adaptive trait shifts often follow a general rule which can
be written as a mathematical equation. Consider a growth affecting
trait X of a biosystem and its relative growth rate RGR; i.e. RGRis
a function of X. The derivative d RGR/d X can be interpreted as the
marginal benefit of adjustments in X. Then it turns out that the rate
of adaptation in X linearly increases with a higher marginal benefit for
the biosystem (Wirtz and Eckhardt, 1996; Wirtz, 2000; Wirtz, 2002b).
Transforming this relation into an evolution equation for X, we have
dX
d RGR
= X
dt
dX
(3)
where X is the specific adaptation coefficient for trait X. In plain

terms, adaptation d X/d t of an effective variable X is directed to maximizing RGR by following the gradient d RGR/d X; with Eq. (3) , classical growth parameters like affinity constants or specific reproduction
rates turn to effective variables of the (bio)system.
The generality of the approach was proven through a broad spectrum
of case studies. These covered changes in phytoplankton community
structure (Wirtz and Eckhardt, 1996), in resource allocation patterns
and in physiological or morphological traits as observed in many plants
and trees (Wirtz, 2000; Wirtz, 2002a). Even the complex regulation
of metabolic pathways, visible through alterations in microbial kinetic
properties can be successfully captured (Wirtz, 2002b). Here we test
for the first time an application to the much different question of
farming development and civilization rise. Our model equations for
macro-economic and cultural development follow from inserting X =
F, Q, T into Eq. (3) .
Applying the general adaptation rule one has to take care of the
case sensitivity of the specific adaptation coefficient X . For X representing an averaged community characteristic, X equals the variance
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of X (Wirtz and Eckhardt, 1996). If, more specifically, X describes

a dimensionless fraction ranging from zero to unity (like F and Q),
one has in place of the variance the product X = X (1 X). In
all other circumstances, X should be treated as a phenomenological
constant (Wirtz, 1998). This is the case for the third specific adaptation
coefficient T . Its relatedness to the variance in T with T 1 confines
the range of reasonable values to [0.1,0.5]. Our choice for the standard
simulation can, like for all other parameters, be found in Table IV.
In order to preserve the capability for adaptation in reasonable times
finite values of F and Q are required; these two variables should not
diminish to arbitrary small levels: In particular, we assume that HG
societies persistently conducted pre-agricultural food production via
the deliberate manipulation of natural resources (Smith, 2001). In the
model, a steady minimal contribution of natural resource management
is represented by using the non-zero initial value of the farming ratio as
lower bound for the later development. Note that a uniform distribution
of F and Q as currently used by the model at simulation start, i.e. the
beginning of the Holocene, can be regarded as strong simplification. Its
accuracy limit is indicated by the recent dating of first crop domestication in the Fertile Crescent to the late Paleolithic (Hillman et al.,
2001). In addition, previously ice covered areas as well as some parts of
South America may not have been populated yet (Binford, 2001). On
the other hand, assigning to each region a common set of initial values
as given in Table III helps to trace back the major causes of regional
differentiation within the simulated period and to keep the focus on
the endogenous development within the Holocene.
2.4. Food productivity and human growth rate
The relative growth rate has to be thoroughly assessed for two purposes.
On the one hand, it generates the derivative terms for the evolution
equations Eq. (3) . Secondly, RGR enables the integration of the population density dynamics since we have, according to its definition,
d P/d t = RGR P .
Links between reproductive success and nutritional status are since
Malthus works in the early nineteenth century often employed by
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demography models for pre-industrial periods (Wood, 1998). In our

model, the growth rate of a regional population relative to its size
(RGR) is mainly controlled by SI introduced in Section 2.2. Total calorie yield resulting from food procuring activity furthermore depends
on regional resource availability, here expressed by the food extraction
potential (FEP). The model roughly resolves variations in FEP due to
over-exploitation of land and resources (van Andel et al., 1990; Louwe
Kooijmans, 2001). When generalizing humanitys impact on the environment, Hardin (1993) introduced a term equivalent to T P n with
n > 1. However, supposing a decrease more than linear in P acts as a
strong growth damping factor which produces a constant distribution
of densities. This is inconsistent with observations of highly variable
population numbers. In addition, technology contributes to local land
degradation, resource depletion or species extinction in some but not
all of its aspects: If the power expression is transformed by taking the
square root, we arrive at a (more realistic) linear relationship with
regard to population density. A proportionality constant (all symbols are described in the following paragraphs and are furthermore
listed in Table V) quantifies the specific fertility loss per capita due
to non-sustainable use. The resulting over-exploitation term has to be
subtracted from the natural FEP.
RGR = (FEP
T P ) (1 T ) SI
P
T
(4)
As technology increases, more and more people neither farm nor hunt.
Construction, maintenance, administration and crafting are time- and
energy consuming tasks. The trade-off which during the Neolithic period presumably regulated the development of high technology societies
is simulated by adding a factor 1 T into Eq. (4) . The constant
denotes a specific labor demand. Assuming that a highly developed
Neolithic community (T 8) allocates between 10 and 40% of its
energy into the technology and organizational sector, one gets an upper
and lower boundary for the specific labor trade-off: 0.01 < < 0.05.
Similarly, a reasonable value for the specific FEP loss should approximately yield a 25% reduction at P 10 Ind km2 , leading to an
estimate of 0.01 Ind km2 .
wirtz_3rd.tex; 20/12/2002; 14:13; p.9
10
Following the above made conventions, the specific growth rate ()

quantifies the survival rate of a well-nourished foraging population in
the late Paleolithic. In addition to the birth rate, also accounts for
non density-dependent mortality. The product of the density-dependent
mortality rate () and P collects negative consequences of crowding;
if one supposes a HG population with P 0.1 Ind km2 to be at
the carrying capacity, should be on the order of 10 Ind1 km2 .
Density-dependent mortality which resulted from infectious diseases
and conflicts (Groube, 1996; Diamond, 1999) can be mitigated by societal differentiation, so that mortality P is divided by T . For example,
death toll to intertribal or larger scale conflicts might be diminished by
recruiting only a certain quota of the male population; other examples
for mortality reducing effects of the technology and organization sector
are housing, prohibition of violence within the community, or medical
knowledge.
2.5. Regional biogeography
In this study spatial gradients in climate impacts on human development are mediated through the regional distribution of vegetation
classes. For reaons of data availability we use the distribution at 5 kyr BP
(see Section 3). This singular choice, albeit blinding out transient regimes
after the onset of the Holocene as well as following abrupt climatic
changes, may still reflect long term climate characteristics during the
Holocene.
So let environmental conditions of each region i be characterized by
its ecosystem type classification which on the other hand corresponds
to a specific mean value of net primary productivity (NPPi ). In our
study, NPP serves as a surrogate for a set of determinants. Its regional
values connect existing biome classification schemes with the major
model inputs FEP, TLI and PAE.
Like NPPi , also local food extraction potential FEPi breaks down a
multidimensional environment including, but not limited to, soil type,
water availability and solar energy input. First evidence suggested a
steady increase of FEPi with growing NPPi . However, with increasing
NPP the relative amount of non-usable biomass rises (Bloom and Sachs,
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11
1998): Tropical rain forests as highly productive biomes offer a lower

fraction of their biomass gain to AP or HG food procurement compared
to less fertile but open vegetation types. Recent studies even suggest
that absolute yields are lower in tropical than in temperate ecozones
(Gallup and Sachs, 1999).
To conform to these findings the model employs a uni-modal dependency of FEP on NPP with a maximum at NPPF . NPPF characterizes
the ecozone which is most profitable for diverse practices as hunting
on big game, collecting plants, farming and herding. According to the
conventions made above, one of the most simple functional realizations
is given by (cmp. Figure 1)
FEPi =
2x
x2 + 1
with x =
NPPi
NPPF
(5)
Linking NPP to the number of potential economies PAE requires a

more elaborate evaluation. Following the concepts of Gaston (2000)
and Caley and Schluter (1997) who analyzed species richness on two
spatial scales separately, PAE is split into two parts,
PAEi = LAEi CAEk
(6)
While CAEk defines an absolute scale (see next subsection), the number
LAEi with 0 LAEi 1 represents the local diversity in domesticables. Huston (1979, 1999) suggested a generic uni-modal relationship
between local NPP and species richness which here is adopted analogously to Eq. (5) with a higher exponent of x in the denominator because reduction of LAEi in tropical habitats is much more pronounced
compared to FEP.
LAEi = TLIi
4x
+3
x3
with x =
NPPi
.
NPP
(7)
NPP denotes the NPP value at which LAE reaches its maximum.
Diversity in domesticable species decreases in ecozones which are either too marginal or too densely forested (Smith, 1998); it attains a
maximum in open woodland (Wright, 1992). Temperature limitation
index TLIi is included into the calculation of LAEi since cold adapted
plants do generally not carry large seeds, fruits or tubers.
Characterizing an open vegetation type, NPP has to be in between
zero and NPPF which represents a temperate steppe or woodland, thus
wirtz_3rd.tex; 20/12/2002; 14:13; p.11
12
Food extraction potential (FEP)
Number of local
agropastoral economies
relative to continental maximum
(LAE)
.5
0
0
500
1000
1500
2000
NPP [g m2 yr1]
Figure 1. The two main environmental determinants of the model which are local
number of potential AP economies (LAE) and food extraction potential (FEP) as
uni-modal functions of regional NPP. The functions are shown for parameter values
of NPP = 0.5 NPPF = 450 g m2 yr1 and TLI = 1.
300 g m2 yr1 < NPP < 600g m2 yr1 ; we simply set NPP = 0.5
NPPF . The resulting behavior in LAE and FEP as functions of NPP
is plotted in Figure 1.
2.6. Continental scale biogeography
The number of continental potential AP economies (CAEk ) is, by definition, equal for all regions i belonging to the same large land mass k,
here denoted as i Ik . This way a second spatial scale is introduced.
Proper boundaries for large scale units of distinct species distributions are difficult to assess (Caley and Schluter, 1997). Especially the
accessibility of both Americas as well as whole Africa from Eurasia
in the Quaternary makes an unequivocal identification of continental
model areas difficult. On the other hand, glacier barriers in northern
America, the tropical Mesoamerican bottleneck and extreme desert
conditions in the Sahara during most periods throughout the Quaternary acted as hardly permeable barriers. As a result, flora and
fauna differ significantly between (1) Eurasia including North Africa,
wirtz_3rd.tex; 20/12/2002; 14:13; p.12
13
(2) South America, (3) North America, (4) Subsaharan Africa, and
(5) Australia. Resolving also (6) Greenland and (7) large islands as
independent land units we arrive at a total of seven continents (Ik , k =
1, . . . , 7).
The continent specific value CAEk should, of course, reflect the
richness in domesticables (LAEi ) of all regions which are member of Ik
what is most easily achieved through summation over LAEi :
CAEk = CAE
max
0
k
iIk0
Ai LAEi
!1
Ai LAEi
(8)
iIk
The region areas Ai serve as weighting factors and, furthermore,

account for a linearized species-area relationship (Begon et al., 1993).
Due to the normalization in Eq. (8) the coefficient CAE denotes the
global maximum of CAEk independent of the distribution of regions.
It is reasonable to assume a value of CAE between 4 and 10. Our particular choice CAE = 8.5 leads to an effective number of 8 economies
(PAEi = 8) in the most suitable Eurasian regions. One may interpret
these, for example, as four productions forms based on cereal crops,
three on herding and one on a mixture of lower-return plants like
squash, sunflower and beans.
Eqs. (7) (8) are constructed in such a way that merely two coefficients (CAE and NPPF ) together with the global arrangement of
regions determine the biogeographic model input. Furthermore, the
recurrence of LAEi in the derivation of CAEk ensures consistency and
compatibility of two differently scaled diversity measures.
2.7. Local spread
Like for all model formulations derived so far, also the part describing
diffusion between regions strongly aggregates over a variety of diverse
mechanisms. These comprise trade, colonization, warfare, migration of
individuals or entire populations. Their net effect on the dynamics of
effective and density variables are calculated using a combination of
rules and methods. To construct a simple but realistic formulation for
exchange we modify the concept of influence theoretically described
and tested by Renfrew and Level (1979). In our approach, influence
wirtz_3rd.tex; 20/12/2002; 14:13; p.13
14
is defined as density P times the local technology T . By including

technology, classical human diffusion models such as proposed by Ammerman and Cavalli-Sforza (1973) are conceptually broadened: The
model resolves mobility enhancement due to technological achievements
such as wheels or route infrastructure.
Differences in influence between adjacent regions not only reflect
unbalanced population pressures but also disparate mercantile ranges,
military powers and efficiencies of transport. The differences act as a
driving force which causes a long-range matter exchange. This flux
is formulated as a first order relaxation process like in a number of
reaction-transport models (Wang and van Cappellen, 1996), and depends on (1) the influence force, (2) common boundary length of the
two regions, (3) the distance of their centers, and (4) a specific exchange
coefficient (Appendix Eq. (9) ). A reasonable range for this coefficient
was derived by de Vries et al. (2002) who also studied effects of changes
within this range.
Crises may lead to additional population movements: We suppose
that people leave their habitats when environmental conditions do not
anymore allow for survival (Dean et al., 2000; Wood, 1998). In the
model, the influence of neighbors is simply set to zero as long as a
populations local relative growth rate is negative and lower than its
neighbors. We prefer to modify the influence based algorithm instead
of introducing a new term or parameter for crisis escape assuming a
similar disposition to mobility in good as well as bad times.
Fluxes induced by differential influences or by crisis control exchange
rates of all model variables. Effective trait variables uniformly follow
a simple transport formulation; population density exchange is formulated accordingly (Appendix Eqs. (10) (11) ). In contrast to the spread
of density, trait variable exchange has to be unidirectional; otherwise
exporting regions would loose developmental status what is rather unrealistic. Imports of AP economies may lead to F > 1 pushing AAE
above its upper local limit PAE. In such a case PAE is increased to the
value of AAE.
Note that remote overseas transport is omitted in the current version
of the model.
wirtz_3rd.tex; 20/12/2002; 14:13; p.14
15
3. Environmental boundary conditions

3.1. Global ecosystem complexes
Global maps of NPP are currently available through different types of
sources. Model based calculations for present conditions are performed
by an increasing number of research groups (Field et al., 1998). However, inherent uncertainty is still considerable, for what large differences
in NPP patterns as simulated by a series of vegetation models are
powerful indicators (Cramer et al., 1999). On the other hand, fieldbased global NPP maps like the one compiled by Bazilevich (1994) can
not easily be extrapolated from the present state to past periods within
the Holocene as well as Pleistocene. As such hindcasts are requested by
envisaged subsequent applications of our dynamical approach, NPP was
estimated using an existing atlas of reconstructed ecosystem complexes.
Adams and Faure (1997a,b) have made available several maps of global
ecosystem complexes from the last glacial maximum to present potential. Their vegetation classification closely follows the carbon inventory
by Olson et al. (1983) modified for the description of paleovegetation.
In the present study, only a single time slice (5 kyr BP) is explicitly
used so far. These maps interrelate all available paleoproxy data and
provide an interpolated expert estimate on paleovegetation in contrast to the pollen-based point vegetation reconstruction in the biome
6000 project (Prentice and Webb III, 1998). A point comparison with
6 kyr BP data for Africa (Jolly et al., 1998) and Europe (Prentice
et al., 1996) showed general agreement between biome 6000 and the
paleovegetation reconstructed by Adams and Faure.
Additionally, the relative change in each vegetation type since the
last glacial maximum was assessed using digital maps. These relative
changes led to small modifications in evaluating CAEk in that Eq. (8)
is calculated at post and mid Ice Age conditions. The mean of both
values yields a more reliable measure for continental biodiversity than
a single time slice (see also Table II).
The second operation comprises the data conversion from the ecosystem type classification scheme used by the authors into values for NPP
and temperature limitation. For each of the biome types in Table I
wirtz_3rd.tex; 20/12/2002; 14:13; p.15
16
estimates for NPP were obtained from Bazilevich (1994); values for
TLI are derived referring to the seasonal frost duration.
Table I. Ecosystem types used in the model and their representation by
Adams and Faure (1997a), conversion to net primary productivity (NPP)
and temperature limitation index (TLI).
Ecosystem type
Adams enumeration
NPP [g m2 yr1 ]
TLI
Tropical rain forest
2000
Tropical woodland
1600
Warm forest
10
1400
Cool forest
14
1200
0.4
Savannah
89
1100
Forest steppe
25
1000
0.5
Grassland
900
Taiga
1517
800
0.8
Med. woodland/forest
13
800
Semi-arid woodland
18
600
Dry steppe
24
500
(Mediterranean) scrub
4,23
250
Forest tundra
26
150
0.95
Tundra
19,20
120
0.95
Semi-desert
50
Hot desert
10
Polar desert
21
Temperate desert
22
0.3
3.2. Elementary region definition

The choice of regions is based on the vegetation type boundaries inherent to the maps of Adams and Faure (1997a,b). To avoid an extremely
unequal distribution of area sizes, an algorithm was developed which
connects smaller patches of unequal biome classification into heterogeneous clusters or subdivides huge biomes like the Siberian Taiga,
wirtz_3rd.tex; 20/12/2002; 14:13; p.16
17
into homogeneous patches. This procedure resulted in a typical area

of 300 000 km2 , corresponding to a division of the globe into 197
land units. Antarctica and some smaller oceanic islands were excluded.
Beside a regions specific area Ai also boundary lengths with each
topological neighbor (Lij ) were calculated by summing along the edges
of the underlying grid cells. In the case of heterogeneous patches, NPP
and temperature limitation TLI were evaluated at each cell according
to Table I and then spatially averaged. Spatial resolution of the grid is
half a degree in latitude as well as longitude.
3.3. Climate fluctuations
Shifts of biomes on a millennium scale are superposed by remarkable
fluctuations in climate throughout the Holocene. Although climate appeared to be more stable during the last 11 kyr than at any other time
during the Pleistocene (Keigwin and Boyle, 2000), many decades or
even centuries lasting cooling events, e.g., the Little Ice Age, repeated
at a period between 11.3 kyr (Fairbridge and Hillaire-Marcel, 1977),
1.5 kyr (deMenocal, 2001) and 2.6 kyr (OBrien et al., 1995), depending
on the world region, the fluctuation magnitude and the climate-related
parameter under consideration. Intense shifts are to some extent spatially correlated on a global scale. However, their appearance in terms
of changed precipitation patterns is ambiguous, even if reduced temperature often produced a drier climate. Nevertheless, beside the negative
effect of lowered temperatures, probably in combination with increasing
storm intensities, conditions which were too dry or too wet may have
equally harmed locally established vegetation (Hs
u, 2000). The overall
impact of a quasi-periodic climate cooling is simulated by an oscillating reduction index c with amplitude c and period . The function
c = 1c cos4 (t/ ) confines reduction to a small interval. When time t
equals 0.5 (2n + 1) (n = 0, 1, . . .), c reaches unity and reduction
vanishes. The omission of an additional phase parameter constrains
the simulation start to distinct time frames. After choosing = 1500
yr, a satisfying overlap of c with existing data is found by dating the
simulated time onset t = 0 with 12 kyr BP, i.e. at the out-phase of the
Younger Dryas.
wirtz_3rd.tex; 20/12/2002; 14:13; p.17
18
Under the assumption that Holocene climate fluctuations left species

composition and, thus, PAE substantially unchanged, c only affects
two measures. First, the temperature limitation index TLI is lowered
(TLI cTLI) in order to simulate simple cooling effects, often accompanied by less precipitation. FEP in all regions is affected, but mediated
through changes in the underlying NPPi . If oscillations are acting on
local NPPi instead of FEP directly, accomplished by FEP(NPP)
FEP(c NPP) in Eq. (5) , a possible positive effect for human food
productivity in tropical or semi-tropical zones is preserved since here
a decrease in NPPi yields higher FEP (see Figure 1). As a consequence, the globally averaged FEP always reveals small variations,
whereas large scale reductions in agricultural subsistence are mainly
synchronized by shifts in TLI.
3.4. Model implementation
The integration of the evolution equations was facilitated by an operator splitting method treating development Eqs. (3) (4) and exchange
Eqs. (10) (11) functions in sequence. A forward Euler scheme with
adaptive time step of maximally one year was used for numerical integration. Initial values of state variables for all regions are listed in
Table III.
4. Model results vs. archaeological evidence

4.1. Distribution of potential agricultures
One of the major results illustrated in this work was reached before
starting the dynamical model. The compilation of regional boundary
conditions, in particular for PAE (PAEi = LAEi CAEk ), provides
important gradients on the two spatial scales under consideration. Continent specific maxima as shown in Table II reveal a subdivision of
the globe into three classes. The first class is solely given by the huge
Eurasian land mass which according to the simple integration algorithm
Eq. (8) offers about twice the potential agricultures as each member of
the second class made up of both Americas and Subsaharan Africa.
wirtz_3rd.tex; 20/12/2002; 14:13; p.18
19
Large islands and small continents form the third class which is most
discriminated in terms of CAE. The ranking of continents is for the
most part consistent with the original distribution of important crops
and domesticated animals (Rotter and van de Geijn, 1999; Diamond,
1999). Only the relative high African value CAE4 overrates the actual
number of relevant species independently domesticated south of the
Sahara.
Table II. Continental number of agricultural production styles CAEk calculated using vegetation maps for 5 and
18 kyr BP.
Index k
Continent
CAEk
Eurasia
8.5
South America
3.7
North America
3.3
Subsaharan Africa
4.8
Australia
2.5
Greenland
0.0
Large islands
0.2
Recent surveys on global agricultural origins pinpoint not more than

five independent farming centers (Smith, 1998). In order of first emergence these are the Fertile Crescent, the middle reaches of the Huang-he
in North China, lower Yang-tse-kiang regions in South China, central
Mexico and the central Andes, which are pictured as red boxes in
Figure 3 for comparison. Excluded from this list are areas where local
species were domesticated first but the concept of domestication and/or
other founder crops had presumably been introduced from elsewhere,
e.g., the African Sahel, the Indus Valley and eastern North America.
Except for the Yang-tse-kiang region, all independent as well as semiindependent centers are already highlighted with respect to other
regions of the same land mass (see map of PAE, Figure 2); it further
wirtz_3rd.tex; 20/12/2002; 14:13; p.19
20
2.0
NPP
[kg yr1m2]
8.3
Potential
AgroPastoral
Economies (PAE)
0
Figure 2. Global distribution of NPP (top, cmp. Table I, compilation of the
original vegetation maps of Adams and Faure is documented in Section 3)

and potential AP economies PAE (bottom, derived from Eqs. (7) (8) ) which
both are used as biogeographic model input. Border lines of all 197 regions
are included.
indicates a high suitability of the western Mediterranean which indeed

adopted agricultural life style very early.
Other regions which according to the simple model algorithm were
privileged in terms of potential agricultures but lack early evidence
of farming are northeast Brazil, southern Argentina and Tanzania. Although the latter was among the first subequatorial areas with agrarian
communities, it is here obviously overestimated with respect to the
more prominent African core lands for millet and sorghum domestication south of the Sahel zone.
The overall coincidence between Figure 2 and the currently known
pattern is striking. Especially the moderate number of autonomous kernels is robustly predicted even though most of the 197 regions resolved
in the model potentially sustain agricultural production. The location
of kernels selected by our algorithm is controlled by the value of NPP
wirtz_3rd.tex; 20/12/2002; 14:13; p.20
21
which describes optimal NPP for LAE. Setting its value to the upper
estimate (cmp. Table IV), the AP developing centers in Eurasia are
shifted westwards to the zone around the Black Sea, whereas in South
America they are moved eastwards, i.e. the Pampas. A lower estimate
for NPP advantages the North China kernel over the Fertile Crescent.
Finally, intercontinental differences remain qualitatively unaffected by
both variations.
The latter result underlines the importance of the region definition.
Much smaller or larger typical area sizes lead to a different global
NPP distribution. The major mode inputs FEP and LAE depend on
the map resolution, since regional NPPi is subjected to the nonlinear
response functions drawn in Figure 1. For example, the value of NPP
for the Fertile Crescent region dominated by arid woodland is made
more favorable due to the contribution of small areas characterized as
grassland or semi-desert (see map preparation in Section 3.2).
One can infer from this scale dependency together with the high
coincidence of projected and actual centers that the typical area size of
3105 km2 may define a relevant spatial scale for the interaction between
human societies and their environment during Neolithic times.
4.2. Agricultural transition

The transition from HG to AP societies develops at different speeds
across the continents. This is visualized by the three time slices in
Figure 3. To understand such inequalities in development rates it is
useful to first analyze time trajectories for a single region. In terms of
the two model variables farming quota Q and relative number of agricultures F , the transition evolves along a typical dynamical pathway
where the temporal evolutions in Q and F are clearly separated. Shifts
in F , i.e. domestication successes, are in general positively directed.
Their velocity linearly increases with higher growth rate derivative
d RGR/d F and, this way, is proportional to the diversity measures
PAE or CAE, respectively. These two factors govern regional and intercontinental differences in the rate of the domestication process from
its early start.
wirtz_3rd.tex; 20/12/2002; 14:13; p.21
22
1.0
8000 yr BP
0
1.0
Farming
quota
6000 yr BP
0
1.0
4000 yr BP
0
Figure 3. Synoptic maps of simulated farming quota Q for 8, 6, and 4 kyr BP. For
comparison, red boxes show the actual five independent agricultural centers. They
are included into a time slice in accordance to latest evidence of their temporal
emergence.
The differential benefit of agricultural food production d RGR/d Q

is negative as long as F remains below a critical value around unity. As
soon as one economy is established, the relative fraction of farming expense explodes. The short duration of the transition in Q (Figure 4) was
graphically anticipated by Zvelebil (1996) using a sigmoidal curve. The
wirtz_3rd.tex; 20/12/2002; 14:13; p.22
23
author based his projection of farming quota dynamics on ethnographic

sources revealing a lack of societies which show equal proportion of HG
and AP subsistence.
For the Fertile Crescent, the abrupt shift is hindcasted at 9 kyr BP as
can be seen in Figure 4. First domestication success was accomplished
for Triticum species such as wild emmer by the Natufian culture dating
back up to 12.5 kyr BP and einkorn wheat in northern Syria around
10.5 kyr BP. New evidence suggest that cereal cultivation may have
started even in the late Pleistocene (Hillman et al., 2001). With some
time lag animal species like goats and sheep followed (Harris, 1996;
Zeder and Hesse, 2000). A marking point of the transition is visible at
the oldest Neolithic town known, C
atal Hoy
uk, where around 10 kyr BP
farming and hunter-gathering techniques temporarily coexisted (Balter,
1999).
In contrast, domestication in the New World centers, e.g., Ecuador
and central Mexico, needed some additional millennia to reach the
critical value in AAE. Since NPP equally approaches NPP in the
Old as well as New World kernels, the lower rate exclusively derives
from a lower continental potential CAE, what, finally, results in a
significant delay of full adaptation to agricultural life style. The division
into Q and F dynamics together with its mathematical analysis puts
the differential onset between Old and New World into a new light.
It is ultimately related to domestication speed which in turn is determined by the domestication potential: The latter controls the necessary
time for reaching a new dynamical state where agricultural practices
pay off and, thus, Q sharply increases. The model study hence offers
some new arguments for the debate about the Old to New World lag
and its site specific extension recently started by anthropologists and
archaeobotanists (Pringle, 1998; Smith, 2001).
Again, the absolute predicted timing is in nearly exact accordance
with archaeological evidence from different parts of the globe. In part,
this reflects the choice of parameter values used, such as for the specific
growth rate . But as already discussed above, other parameterizations
in general leave the relative timing unchanged.
In Figure 4 also simulation results for two regions adjacent to the
Fertile Crescent are displayed. Starting from 9 kyr BP, both regions
wirtz_3rd.tex; 20/12/2002; 14:13; p.23
24

1.2
Farming quota Q
Fertile Crescent
Asia Minor
Northern Arabia
Ecuador
0.8
0.4
Agricultural economies AAE
8
6
4
2
0
12
11
10
Time [kyr BP]

Figure 4. Dynamics of actual agricultures AAE and relative farming expense Q
simulated according to Eq. (3) . The first AP centers of the New (Ecuador) and the
Old World (Fertile Crescent) are shown. In order to focus on local interaction effects,
two neighbors of the latter are added, both with low (Northern Arabia) and high
FEP (Asia Minor).
import initial founder crops as well as skills for their productive use. In
the forested part of Asia Minor the lag between domestication and full
scale farming is hardly recognizable since the intruding farming wave
transports domesticates simultaneously with production style. Due to
the regions high fertility the adaptation proceeds at high speed without
leaving room for remaining HG societies. The impact is by far more
dampened in northern Arabia, where development is retarded for about
two millennia. The low potential of this area is further declined by
the periodically deteriorating climate. Several long lasting periods of
wirtz_3rd.tex; 20/12/2002; 14:13; p.24
25
incomplete importation of AP economies can be seen between 7.5 and

4 kyr BP.
Northern China, the second independent Eurasian center, develops in parallel with the Fertile Crescent (Figure 3). Whereas earliest evidence of genetically changed livestock in East Asia lags behind
the evidence for the Fertile Crescent about one millennium, some researchers have put forward a roughly simultaneous timing of both
centers (Shelach, 2000). However, regional peculiarities for East Asia
are less well reproduced. A prominent example is the late development
of the semitropical Yang-tse-kiang basin as projected by the model.
A main reason for the obvious disagreement with observation (and
current opinion) was already touched above: A NPP to LAE relation
with a maximum at more open biomes favors large seeded annuals
whereas less importance is attributed to the progenitors of small seeded
semitropicals or tropicals such as Oryza rufipogon. As a consequence,
the nowadays global relevance of rice cultivation is underestimated.
Western Europe trajectories clearly anticipate the actual transition.
As far as the Iberian Peninsula is concerned, no evidence for the abandonment of HG life style on a larger spatial scale before 6 kyr BP as
suggested by the mid panel of Figure 3 can be found (Zilh
ao, 1993).
The northwest African dependence on Fertile Crescent founder crops
and animal husbandry is yet under discussion (Smith, 1998). Note that
according to our simulation study the agricultural life style in the Western Mediterranean area is substantially endogenous. The Kapsai began
to herd sheep and goats around 7 kyr BP, other residents of the western
Sahara herded cattle and cultivated barley as early as 8 kyr BP. Recent
genetic studies suggest an early independent domestication of cattle in
the eastern Sahara (Hanotte et al., 2002). These early stages are only
visible in coastal areas. Differences in the hinterland stem from treating
the Sahara in its semi-desert appearance at 5 kyr BP and not as fecund
grassland which it was till 6 kyr BP.
Simulated transition of East Africa is almost complete at 4 kyr BP.
In fact, pastoralism emerged there (Hanotte et al., 2002), probably
at 34 kyr BP (Diamond, 1999), followed by Bantu cereal farming at
32 kyr BP (Smith, 1998). The coincidence with the model prediction
wirtz_3rd.tex; 20/12/2002; 14:13; p.25
26
surprises given that richness in native African domesticables is overestimated by the model.
Two distant centers in the Andes, the Ecuador highlands and the
Lake Titicaca Altiplano provided the basis for the first pre-Colombian
societies such as the Valdivan culture or the Tiahuanacan Empire.
Records for monumental architecture and irrigation agriculture at Caral,
a coastal site 500 km south of the Ecuadorian region, can be dated back
to 45 kyr BP (Solis et al., 2001). In contrast, secondary regions with
relatively early predicted AP food production, the western Pampas and
California, lack prominent cultural relics. Model projections for highand lowland Mexico are again compatible with classical theories and
recent findings (MacNeish, 1992; Pope et al., 2001).
Though domestication or field cultivation may have occurred within
isolated spots in the Amazon tropical forest (Mann, 2000), no evidence
for agrarian societies occupying huge areas have been found. The exclusion of the tropics similar to the African and Southeast Asian case
is therefore fully consistent with archaeological data.
By 43 kyr BP, only parts of the western margin of South America
as well as of the Southwest of North America have adopted farming.
Currently available records suggest more agricultural activities in the
eastern part of the United States where squash was brought under
domestication very early, followed by other crops such as goosefoot
and sunflower (Smith, 1998). Note that following our definition of production style, labor intensive cultivations of the listed species are not
regarded as efficient energy sources and, thus, do not sum up to a
value of AAE equal to unity as required for the transition to full scale
farming. According to this, large North American societies rose not
until they were based on maize. The Cahokians temporally flourished
along river banks in the Southeast (Milner, 1998), approximately contemporaneous with the southwestern Anasazi civilization (Wills, 1988).
In the standard model run full scale farming in the zone between the
Rocky Mountains and the western Great Plains started at 4 kyr BP
independently from the Mexican kernel (Figure 3), raising doubts about
the current concept of an import of subsistence intensification into that
region.
wirtz_3rd.tex; 20/12/2002; 14:13; p.26
27
Australia retains regions with predominantly HG life styles similar

to large but remote islands like Madagascar or New Guinea.
4.3. Technology evolution
The overall development of technology differs from the temporal pattern observed for the agricultural transition in that it develops more
steadily with respect to either Q or F (cmp. Figure 5 and Figure 4)
and is not restricted to AP societies. On a global scale (cmp. Figure 6),
the regional differences in technological development are hence not
as pronounced as those for other model variables. Before 9 kyr BP,
many later innovation centers are slightly less developed than their
more fertile vicinal zones. Yet T benefits greatly from the establishment of first agricultural economies and exhibits its fastest increase
within 1 000 to 2 000 years after the establishment of full-scale farming.
Thereafter, technology advance is more and more dampened as obvious for the Near East trajectories in Figure 5. These findings are in
good accordance with the observations made by Childe (1936, 1942):
He noticed that the Neolithic revolution was everywhere accompanied
with a two millennia lasting avalanche of technological innovations as
exemplified by the invention of irrigation systems in north Iraq by
8 kyr BP. For the subsequent period only minor improvements could
be claimed by the local societies. Nolan and Lenski (1998) interpret
this class of phenomena as a divorce between expertise and incentive
in increasingly stratified systems. Our model calculations are able to
simplify such hypothetical causalities by following Woods hypothesis
of a declining marginal productivity of innovation (Wood, 1998). A
balance between differential costs and benefits of further innovations is
mathematically inherent to the evolution equation of T derived from
our basic adaptation assumption.
As a result of imposed climate fluctuations, the rate of change in T
varies only slightly with c. Apparently, T increases more slowly, when
climatic conditions deteriorate and local population densities decline
(Figure 5). This is in contrast to the suggestion by Moran (1979), that
the demand for resource intensification is expected to rise when the
carrying capacity of a habitat is exceeded. Also, absolute values of T
wirtz_3rd.tex; 20/12/2002; 14:13; p.27
28
Technology index T
10
8
Fertile Crescent
Asia Minor
Northern Arabia
Ecuador
6
4
2
Density P [Ind km ]
0
18
16
14
12
10
8
6
4
2
0
12
11
10
Time [kyr BP]

Figure 5. Temporal evolution of technology T and population density P for four
regions as simulated by applying Eq. (3) and calculating the population growth rate
RGR for each region.
do generally not show significant reversal phenomena which have been

documented for a small number of civilizations or regions (Richerson
et al., 2001; Zvelebil, 1996) and even mathematically formulated in
the appendix of Richerson et al. (2001). In this study, the minute
reversals observed for the established agricultural economies (Figure 4)
are mainly due to populations with lower AAE which invade from
neighboring areas threatened by worsening climate.
4.4. Population growth and migration
Oscillations in local (Figure 5) and global (Figure 7) population density
are driven by imposed climate fluctuations. In regions with intensive
wirtz_3rd.tex; 20/12/2002; 14:13; p.28
29
9.4
8000 yr BP
1
9.4
Technology
6000 yr BP
1
9.4
4000 yr BP
1
Figure 6. Snapshots of global technological advancement described via the model

variable T .
agriculture and high population densities near carrying capacity, climatic deterioration can cause a decline of P by up to 30 per cent,
as shown in Figure 5 for the Fertile Crescent. Large scale population
decline may be tracked back to the collapse of a series of civilizations in
the New and Old World in response to climatic change or long lasting
drought, respectively (deMenocal, 2001; Weiss and Bradley, 2001). A
wirtz_3rd.tex; 20/12/2002; 14:13; p.29
30
critical examination of correlating cultural collapse and natural hazards

was performed by Erickson (1999).
In developed world regions, highest densities occur nearby AP centers but not therein. With an increasing number of regions practicing
intense food production global population size advances exponentially
from about 2.5 million at 12 kyr BP to 300 million at 3.5 kyr BP (Figure 7). Both numbers fit within the scatter of published estimates
(Groube, 1996; Ambrose, 1998) very well. For the early Holocene, data
are most uncertain; the cited sources presume an average density of
foragers on the order of 0.1 Ind km2 , at least one order of magnitude
higher than the data calculated for the western Europe Magdalenian
by Bocquet-Appel and Demars (2000) or compiled by Binford (2001).
Modern agrarian communities typically have a density more than twice
as high as the numbers given in Figure 4 which are more characteristic
for horticultural societies (Nolan and Lenski, 1998). Rather than
relating this discrepancy to model shortcomings one has to consider
that model regions span over a variety of archaeological sites, making
it difficult to compare model and data derived numbers.
The course in total global migration between regions is shown in
Figure 7. Remote migration follows the initial farming wave and the
same corridor between the two Eurasian centers can be observed for
population density (Figure 8). Arid regions south of this corridor do
not sustain high densities. Deserts surrounding the more fecund Indian
subcontinent act as migration barriers so that development projections failed there. In particular, the missing overseas transport retards
the simulated development of the Indus valley for several millennia
with respect to the actual one. Also other coastal areas reveal timing
mismatches to archaeological evidence (e.g., Italy, British islands, Mississippi valley). This can be taken as a strong indication that coastal,
long-range trading routes indeed played a dominant role for development pathways as was, for example, stressed by Hanotte et al. (2002).
It is likely that remote transportation will also affect the time course
of global migration rate given in Figure 7.
Under deteriorating conditions a net migration occurs away from
more marginal zones like the Fertile Crescent into better suited areas
like Asia Minor. Crisis induced population movement accounts for more
wirtz_3rd.tex; 20/12/2002; 14:13; p.30
31
Global population [108 Ind]

Global migration rate[105 Ind yr1]
Global FEP TLI
0
12
11
10
Time [kyr BP]

P
Figure 7. Simulated global population size and migration rate as given by i Pi Ai

P
and i |d Pi /d ttrans Ai |, respectively. Climate fluctuations, expressed by FEPTLI,
are shown for comparison.
than 70% of total capita exchange. Massive and distinct peaks occur
when climate deteriorates, but long before it reaches its millennia minimum. Thus, many densely inhabited regions are near their carrying
capacity, making their residents prone to famine in the face of a nonoptimal climate (Wood, 1998). Relatively small fluctuations have to
be immediately regulated through emigration. Together with massive
population decline, outward migration alleviates over-exploitation and,
this way, the pressure on local FEP; it leads to a recuperation of RGR
to positive values at a time when climate is still most unfavorable. This
phase dependent pattern is heavily in line with recent findings. The
Tiwanaku collapse in the Peruvian Altiplano and the Akkadian collapse
wirtz_3rd.tex; 20/12/2002; 14:13; p.31
32
18.0
8000 yr BP
0
18.0
Density
[Ind km2]
6000 yr BP
0
18.0
4000 yr BP
0
Figure 8. Global development of population density P . The second time slice

(6 kyr BP) coincides with a period of imposed climate deterioration.
in Mesopotamia are both dated at the onset of multi-century droughts

(deMenocal, 2001). More importantly, subsequent Akkadian resettlement took indeed place while dry conditions were still prevailing. In
decoupling the climate index c and migration flux, the model realistically disintegrates a static one-to-one relationship between cultural
behavior and environmental conditions.
wirtz_3rd.tex; 20/12/2002; 14:13; p.32
33
Except for the coastal Ecuador region, New World kernels as well
as secondary Eurasian centers like the Hindus valley suffer from the
negative influence of forested neighbor regions. Before the full adoption
of an AP production style, these surrounding areas are privileged in
terms of T and P (Figures 6 and 8) and therefore exert a high influence
which is, for example, visible through their export of low Q and F
status. Such negative migration effects on the local development are in
more detail described by de Vries et al. (2002).
The migration submodel also separates population and cultural dynamics. On a global scale, correlation between P and T made visible
in Figures 6 and 8 turns out to be inhomogeneous. For example, according to our simulation study the continuously sparsely inhabited
northern part of the Arabian peninsula is among the most developed
areas in the mid Holocene. The status is maintained through exhaustive
import of technology from its southern and northern neighbor regions
indicating alternating colonization and/or the existence of vital trade
routes. However, it remains difficult to relate any single trajectory like
the North Arabia case to historical knowledge. The combination of
strictly periodic fluctuations and a stationary vegetation distribution
yields a picture which may be representative on a larger time and
spatial perspective but is far from being confidential when it comes
to detail. Single events, albeit important for macro-history, keep their
unpredictable nature.
4.5. Model sensitivity and climate variability
The timing difference between the Neolithic shift in the New and Old
World provides a simple but effective measure for the global diversification of coarse historic pathways. In the standard simulation, the
time lag is, quite realistically, about 3.5 kyr. In order to detect the
robustness of this projection, model coefficients were manually varied.
In many cases, no dramatic changes in the duration of the lag occur,
but also exceptions were found. The initial value of the farming ratio
Q(t = 0), for instance, has a great influence on the overall timing since
the number linearly affects the evolution in F (see Section 4.2). It is
therefore crucial to rely estimates in Q(t = 0) on a more profound
wirtz_3rd.tex; 20/12/2002; 14:13; p.33
34
basis, for example by including also pre-Ice Age conditions into the
simulation.
Agriculture was developed independently in several areas within a
relatively short period after millions of years of hominid and human
hunting, scavenging and gathering. This striking coincidence has led
some researchers to the hypothesis that the stability of the Holocene
climate might have been a precondition for the Neolithic transition
(Richerson et al., 2001). In order to test the strength of this argument
we varied the climate fluctuation intensity from zero to one. The variation result on the New to Old World timing as visualized in Figure 9
suggests a different picture: Though acute and recurrent deteriorations
in climate retard the onset in all centers, they are nowhere able to prevent it. In addition, retardation is stronger in the first American kernel
(Ecuador) than in the Fertile Crescent indicating a higher vulnerability
to external hazards of regions developing at a lower pace.
5. Summary and conclusions

Adaptation of populations to changing and heterogeneous environmental conditions is a complex process. One therefore should be cautious
in asserting that the many facets of the human past can be reduced to
a simple formula (Taylor, 2001). However, Toynbee (1989) in his monumental work demands that single historical events, personages or ethnic
processes must be disregarded when looking at long-lasting processes.
Only then functional aspects may become evident. Here, to predict the
different timings of Neolithic kernels, a common rule set is applied to
few biogeographic factors with regionally different values. The ability of
the simple model to robustly produce a pattern of agrarian civilizations
very similar to those of archaeological sites substantiates the view of
functional similarity in the many different pre-historical trajectories. A
similar approval is provided to the old concept of culture being defined
as an acclimatization to a particular environment (Steward, 1955).
Two types of applications may take profit out of this approach. The
first is linked to the mere reproduction of large scale historic processes.
Transient shifts in production style due to innovations are realistically
wirtz_3rd.tex; 20/12/2002; 14:13; p.34
35
Farming onset [kyr BP]
10
Eurasia
America
2
0
0.2
0.4
0.6
0.8
Climate fluctuation intensity c*
Figure 9. The effect of climate fluctuation intensity on simulated farming onset

for Eurasia (Fertile Crescent) and America (Ecuador). The value of c used in the
standard run is marked by a dotted line.
calculated in terms of environmental conditions and of adaptability as

well as current developing state of human communities. It is hitherto
demonstrated that estimates of population growth and migration patterns are achievable with dynamical models. Though these are not yet
adjusted to the (post)industrial age, a foundation is laid for future models able to include the human dimension into climate change scenarios
in a fully dynamical way.
Even though in this study the emphasis lays on model development
and validation, causal analysis has been shown to be a second application field. A selection of known problems in prehistory is discussed: (i)
We propose an explanation for the lag between domestication and full
scale farming. Both activities are mutually dependent, but a positive
shift in farming requires that the number of energy rich domesticates
exceeds a critical value. In the opposite direction, no threshold in the
wirtz_3rd.tex; 20/12/2002; 14:13; p.35
36
farming ratio exists which could prevent the monotone improvement in

the domestication of wild plants and animals; (ii) Neolithic populations
have reached carrying capacity which is in turn feedback-controlled
by over-exploitation. Then even a moderate deterioration of climate
soon leads to the formation of distinct migration waves. Technology
evolution as potential elevator of carrying capacity is dampened out
as long as crisis intensity increases but resumes afterwards. If climate
change is synchronized world-wide, the globally integrated spreading
rate can explode or implode by more than one order of magnitude
within few decades. (iii) Population dynamics is much more sensitive
to external fluctuations than macro-economic or technological trajectories. (iv) Our simulation study rejects the widespread hypothesis that
local population pressure due to immigration or climate change
was a prime mover of agricultural development. Instead, we confirm
the view of a steady innovation and competition between different ways
of life (Boserup, 1981). Also the cultural maintenance of sub-optimal
subsistence strategies on a low level may have played an important
role due to a net increase in adaptability. (v) Regional and continental
differences in the rise of agrarian societies, followed by high technology civilizations can be ultimately linked to continental geography
and the vegetation distribution during the Holocene. We proposed a
simple method to upscale from biogeography to the maximal number
of regional agricultures and proved its key role for the overall dynamics.
Of course, many other applications can be sought on the base of
the first model version presented here. For example, the late Pleistocene, or more precisely the whole period following the appearance of
the anatomically modern genus Homo sapiens should be taken under
consideration.
The high explanatory power reached by the presented model supports our initial goal to build a scheme which is as simple as possible.
We recommend that future improvements save this strategy. These
can be made in many directions like the implementation of overseas
transport or a dynamic coupling to a climate model. But yet through
careful examination of the respective model shortcomings like dependence on long-range trading or on climate variability, for example,
wirtz_3rd.tex; 20/12/2002; 14:13; p.36
37
our understanding of human prehistory in its coarse appearance has

progressed.
Acknowledgements
Our special thanks go to Peter Kohler and Maximilian Schafer for a
critical reading of the manuscript. We are grateful to Andrew Sherrat, Kees Klein Goldewijk, Michael Coe and Navin Ramankutty who
provided valuable comments on an earlier version of this manuscript.
We wish to acknowledge the encouragement and support of Wolfgang
Ebenh
oh and Bert de Vries.
wirtz_3rd.tex; 20/12/2002; 14:13; p.37
38
Tables
Table III. Model variables
Symbol
description
initial values
ratio of established economies
0.2
population density
0.03
farming ratio
0.04
technology index
1.0
unit
Ind km2
Table IV. Model parameters

Symbol
description
value
range
amplitude in climate fluctuation
0.3
00.5
Global max. of PAE
8.5
510
specific technology adaptability
0.15
0.10.5
over-exploitation coefficient
0.01
0.0050.02
Ind1 km2
maximum specific growth rate
0.0025
' 0.1
yr1
NPP
NPP where LAE = 1
450
300600
g m2 yr1
specific density mortality rate
0.025
' 10
Ind1 km2 yr1
exchange coefficient
0.08
0.010.3
Ind1 km3 yr1
period of climate fluctuations
1500
12002600
yr
production loss coefficient
0.04
0.010.05
CAE
unit
wirtz_3rd.tex; 20/12/2002; 14:13; p.38
39
Table V. Auxiliary variables and symbols

Symbol
description
unit or range
region area
km2
AAE
number of actual economies
climate fluctuation
CAE
Continental maximum of PAE
specific adaptability in F
]0; 1[
specific farming ratio adaptability
]0; 1[
fij
migration/exchange flux
yr1
[0; 1]
FEP
food extraction potential
[0; 1]
i, j
region indices
1 . . . 197
continental set of regions
index for continents
1...7
boundary length
km
LAE
region specific part of PAE
[0; 1]
neighborhood set of regions
NPP
net primary production
kg m2 yr1
NPPF
NPP where FEP = 1
kg m2 yr1
PAE
potential number of economies
RGR
relative growth rate
SI
subsistence intensity
TLI
temperature limitation index
yr1
[0; 1]
wirtz_3rd.tex; 20/12/2002; 14:13; p.39
40
Appendix: Transport flux calculation

The transportation driving force fij is proportional to the difference
between local influence Ti Pi and the spatial average hT P iij with
hT P iij = (Ai Ti Pi + Aj Tj Pj )/(Ai + Aj )
fij grows linearly with boundary length Lij between i, j and is inversely
related to the product of distances from gravity center to border line,
approximated by and proportional to the square root of the area,

Lij
hT P iij Ti Pi
Ai Aj
fij = p
(9)
where collects all constants of proportionality. Transport of trait X

between i and its neighborhood j Ni is formulated as

X
d Xi
=
fij (Xj Xi ) (fij )

d t transport jN
(10)
where is a binary filter function defined as

(fij ) =
1, if fij > 0;
0, otherwise.
and ensures unidirectional flux. Aiming, but not perfectly reaching

mass conservation during bidirectional flow of individuals, the force
fij is divided by the technology index of the exporting region so that
population density changes due to migration are expressed as

X fij Pi
d Pi
fij Pj Ai
=
(fij ) +
(fij ) .

d t transport jN
Ti
T j Aj
(11)
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41
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