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Ecology, Economy and Society–the INSEE Journal 4 (1): 5–12, January 2021

COMMENTARY

Mathematical Ecology, Evolution, and the Social

Sciences
Simon A. Levin 

Abstract: The last few decades have seen an enhanced partnership between
ecologists and social scientists, especially economists, in addressing the
environmental challenges facing societies. Not only do ecology and economics, in
particular, need each other; but also the challenges they face are similar and can
benefit from cross-fertilization. At the core are scaling from the micro- to the
macro, the development of appropriate statistical mechanics to facilitate scaling,
features underlying the resilience and robustness of systems, the anticipation of
critical transitions and regime shifts, and addressing the conflicts of interest
between individual agents and the common good through exploration of
cooperation, prosociality and collective decision-making. Confronting these issues
will be crucial in the coming years for all nations, especially those in South Asia that
will suffer in major ways from the consequences of overpopulation, climate change
and other environmental threats.
Keywords: Mathematical Ecology and Evolution; Economics; Public Goods;
Scaling; Complex Adaptive Systems.

1. INTRODUCTION
Though my academic training was as an applied mathematician, I was
drawn even as a graduate student to problems in biology, especially in
ecology. I received my Ph.D. in 1964, two years after the appearance of
Rachel Carson‘s monumental Silent Spring and just a few years before the
influential Population Bomb of Paul and Anne Ehrlich (Carson 1962; Ehrlich
1968). In this context of increasing societal attention to environmental


James S. McDonnell Distinguished University Professor, Department of Ecology and
Evolution, Princeton University, Princeton, New Jersey 08544-1003; [email protected].
Copyright © Levin 2021. Released under Creative Commons Attribution-NonCommercial
4.0 International licence (CC BY-NC 4.0) by the author.
Published by Indian Society for Ecological Economics (INSEE), c/o Institute of Economic
Growth, University Enclave, North Campus, Delhi 110007.
ISSN: 2581-6152 (print); 2581-6101 (web).
DOI: https://doi.org/10.37773/ees.v4i1.401
 Ecology, Economy and Society–the INSEE Journal [6]

degradation, I was increasingly drawn to studying the environment, and, by

1970, I was fully committed to the field of ecology, and in particular the
development of theoretical foundations.
Theoretical ecology is a subject that is at least a century old. Its earlier roots,
going back several centuries, were in demography and the biology of
populations, spurred on by the need for more quantitative approaches to
these subjects. The first fundamental mathematical advances were made by
the great mathematician Vito Volterra and the physical chemist and
mathematician Alfred Lotka, who independently derived sets of differential
equations to describe the dynamics of species in competition and
exploitation (Lotka 1925; Volterra 1926). Volterra, in particular, was
interested in the fluctuations of the Adriatic fisheries, a problem brought to
him by Umberto d‘Ancona, a formidable fisheries biologist, and his simple
equations were able to explain those fluctuations. The equations of Lotka
and Volterra still are at the core of much work in theoretical ecology today
and have found application in economics and other social sciences
(Samuelson 1971, 1974).
I enjoyed pure theoretical ecology and still do; but I soon realized that to
address environmental issues substantively, I needed to turn more
effectively towards economics and other social sciences. The key issues that
needed to be addressed included how to value biodiversity and convince
others of that value; matters of discounting the future, equity, and the
interests of others; and the role of cooperation and social norms in the
development of management strategies in local and global commons. Thirty
years ago, I was invited to join the nascent Beijer Institute at the Swedish
Academy of Sciences, and that interdisciplinary paradise changed my
research trajectory, and those of countless others, forever. I learned how
much ecology needed economics and how much economics needed
ecology, and the collaborations that followed have been invaluable.
However, I also learned something else. The problems of ecology and
economics are two sides of the same coin; this is no accident, because both
disciplines address the self-organization and evolution of communities of
individual agents competing, exploiting, and cooperating with emergent
dynamics at higher levels of organization that produce systems that cycle
various kinds of capital, natural and created, and provide a context in which
individual behaviours change. Kenneth Arrow, one of the key members of
the Beijer community, pointed out to us that the fact that both disciplines
begin with ―eco-,‖ meaning household, was not coincidence; they both face
similar issues—both confront complex adaptive systems (CAS) and the
challenges of understanding and managing them (Arrow, Ehrlich, and Levin
2014).
[7] Simon A. Levin

2. SOME COMMON PROBLEMS IN ECOLOGY AND

ECONOMICS
Because similar problems have been addressed in both disciplines, there are
reciprocal benefits from revisiting the different approaches that have been
developed in the two. Such mutualisms are increasing in number, and I will
touch on only a few of them in this section. These include but are not
limited to the matter of scaling across levels and across space and time
(Chave and Levin 2003); foraging theory and search theory (Pyke, Pulliam,
and Charnov 1977); the potential for critical transitions (Steele 1998;
Scheffer 2009); and identifying the structural features that make systems
robust and resilient (Holling 1973; Levin et al. 1998; Levin and Lubchenco
2008).
Evolutionary ecology to a large extent addresses how genomes have been
adapted to deal with uncertain environments. The vertebrate immune
system is a wonderful example of evolution‘s hierarchical response to the
certainty of uncertain events and is a model for how societies might deal
with everything from pandemics to terrorist attacks to financial collapse
(Levin and Lo 2015). More generally, life history theory in evolution deals
in large part with the allocation of resources over an evolutionary lineage
(Stearns 1992) and has strong parallels with models of the theory of interest
(Haberler and Fisher 1931) and of resource allocation and dynasty theory in
economics (Becker 1965; Heckman 2015). Kenneth Arrow and I merged
these by examining through dynamic programming how uncertainty of
various kinds would affect consumption patterns and deferral of resources
to one‘s offspring, a fundamental problem in evolution and economics alike
(Arrow and Levin 2009).
More generally, evolution deals with fitnesses and the processes that select
more fit genotypes over less fit ones, while economics does the same, with
the word ―fitnesses‖ replaced by ―utilities‖. Both ecology and economics
therefore utilize techniques from optimization and game theory to
understand what strategies can be expected to prevail; indeed, cross-
fertilization between the two perspectives on such problems is rapidly
gaining traction.

3. COMPLEX ADAPTIVE SYSTEMS

Ecological systems, the socio-economic systems with which they interact,
and the interconnected global socio-ecological system that emerge, are all
examples of CAS (Holland 1995; Levin 1998, 1999; Arrow, Ehrlich, and
Levin 2014). Such systems are composed of individual agents that interact
 Ecology, Economy and Society–the INSEE Journal [8]

with one another at local scales, leading to the emergence of higher order
patterns and processes that feed back to modify individual behaviours. The
agents—be they genomes, organisms, institutions, or nations—adapt their
behaviours, in some cases over evolutionary time, generally in response to
differential payoffs, utilities, or fitnesses; hence, the term ―adaptive‖ in the
name. The term, however, can be misleading, since the myopic benefits to
individuals may not advance the collective good; the adaptation is at levels
below the whole system.
Whatever the natures of the CAS, similar questions arise: What factors
underlie the robustness and resilience of these systems to change,
exogenous or endogenous? Are there early warning indicators of impending
transitions? How do patterns vary across scales, and how can we use
coarse-grained models to understand the emergent properties as statistical
consequences of behaviours and interactions among large numbers of
agents? And perhaps most challenging, how do conflicts between the
interests of individual agents and the collective good play out, and to what
extent do they undermine equity or system resilience?
These are, of course, all familiar challenges to economists, who are used to
dealing with the interplay between microeconomics and macroeconomics
and with the challenges of public goods and common-pool resources. It
might be surprising, however, for economists to learn that such questions,
especially of cooperation in the face of public goods, are at the centre of
much evolutionary theory. Bacteria produce extracellular polymers that
provide collective benefits as well as antibiotics that poison susceptible
neighbours; plants fix nitrogen, transforming it into a form more widely
available to free-riders; and species from cellular slime moulds to birds, fish,
and humans engage in collective behaviours that improve access to
resources or mutual defence. All involve costs and all are susceptible to
free-riding. Understanding how and under what conditions such behaviours
are maintained, and when they break down (as in tumour growth), are
among the deepest problems in evolutionary theory—puzzles that worried
Charles Darwin enough to delay publication of The Origin of Species for many
years.

4. PUBLIC GOODS, COMMON-POOL RESOURCES, AND THE

TRAGEDY OF THE COMMONS
Our planet is threatened by a litany of interlocking problems—climate
change, biodiversity loss, overpopulation, and increasing inequity—that
must be solved if we are to achieve a sustainable future. William Forster
Lloyd first spoke of the commons nearly two centuries ago (Lloyd 1833),
[9] Simon A. Levin

and Garrett Hardin famously introduced the notion of the ―tragedy of the
commons, arguing that the solution was in ―mutual coercion, mutually
agreed upon‖ (Hardin 1968). Views about managing the commons were
changed, however, by the landmark work of another core Beijer participant,
Elinor Ostrom, who showed, through empirical and theoretical arguments,
that societies could self-regulate through the development of social norms
and other institutions that limited overexploitation (Ostrom 1990). Dealing
with the commons is at the centre of the partnership that must grow
between biology and the social sciences in the coming decades. Economists
have with great success developed frameworks for dealing with public
goods by addressing questions such as: What is the social optimum?
Without top-down regulation, can the social optimum be approximated and
sustained as a Nash equilibrium or does discounting of the future
undermine it? Where it cannot be sustained, are there second-best solutions
that can be? In cooperative games, can co-operators develop social norms,
for example, the punishment of defectors, that can flip the system to more
favourable equilibria? What is the role of prosociality and how can it be
achieved (Gintis 2003; Akcay et al. 2009; Dixit, Levin, and Rubenstein 2013;
Dixit and Levin 2017)? Evolutionary biologists have had to tackle the same
problems from different perspectives, focusing attention on the evolution
of altruism and cooperation and the importance of genetic relatedness
(Hamilton and Axelrod 1981; Nowak, Tarnita, and Wilson 2010; Levin,
2014). We now need to bring these separate approaches together to develop
a theory of governance for the global commons. (Polasky et al. 2019).
Ostrom‘s ideas of polycentric governance (Ostrom 2009) are likely to play a
central role in what must be at the top of the agenda for ecological and
environmental ecologists for many years to come.

5. CONCLUDING THOUGHTS
The problems discussed in this paper are, of course, of general relevance to
all nations, but they pose special challenges for India and other parts of
South Asia. Overpopulation, biodiversity loss, and the growing inequity in
well-being are especially acute in these parts of the world, as documented in
the Dasgupta report for the British government on the economics of
biodiversity (H M Treasury 2020). Infectious diseases, like the current
COVID-19 pandemic, threaten devasting consequences. Climate change
will impact India, China, Pakistan, and Bangladesh especially hard, and
expanding populations and their resource needs will exacerbate the climate
crisis, unless steps towards promoting alternative energy are successful. It is
no surprise then that there has been increasing attention on implementing
decarbonization as rapidly as possible, especially in India and China, and in
 Ecology, Economy and Society–the INSEE Journal [10]

addressing the social norms issues that will be crucial to people‘s willingness
to accept it. Thus, the marriage of ecology, economics, and other social
sciences is especially crucial in South Asia and stands to benefit enormously
from the efforts of the readers of this journal. I hope this call to action will
find enthusiastic acceptance.

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