Evolution of life in urban environments
Welcome to the city
Human populations are shifting en masse to cities, which is leading to rapid increases in the number and extent of urban areas. Such changes are well known to cause declines in many species, but they can also act as alternative selection pressures to which some species are able to adapt. Johnson and Munshi-South review the suite of pressures that urban environments exert, the ways in which species may (or may not) adapt, and the larger impact of these evolutionary events on natural processes and human populations. Understanding such urban evolution patterns will improve our ability to foster species persistence in the face of urbanization and to mitigate some of the challenges, such as disease, that adaptation can bring.
Science, this issue p. eaam8327
Structured Abstract
BACKGROUND
The extent of urban areas is increasing around the world, and most humans now live in cities. Urbanization results in dramatic environmental change, including increased temperatures, more impervious surface cover, altered hydrology, and elevated pollution. Urban areas also host more non-native species and reduced abundance and diversity of many native species. These environmental changes brought by global urbanization are creating novel ecosystems with unknown consequences for the evolution of life. Here, we consider how early human settlements led to the evolution of human commensals, including some of the most notorious pests and disease vectors. We also comprehensively review how contemporary urbanization affects the evolution of species that coinhabit cities.
ADVANCES
A recent surge of research shows that urbanization affects both nonadaptive and adaptive evolution. Some of the clearest results of urban evolution show that cities elevate the strength of random genetic drift (stochastic changes in allele frequencies) and restrict gene flow (the movement of alleles between populations due to dispersal and mating). Populations of native species in cities often represent either relicts that predate urbanization or populations that established after a city formed. Both scenarios frequently result in a loss of genetic diversity within populations and increased differentiation between populations. Fragmentation and urban infrastructure also create barriers to dispersal, and consequently, gene flow is often reduced among city populations, which further contributes to genetic differentiation between populations.
The influence of urbanization on mutation and adaptive evolution are less clear. A small number of studies suggest that industrial pollution can elevate mutation rates, but the pervasiveness of this effect is unknown. A better studied phenomenon are the effects of urbanization on evolution by natural selection. A growing number of studies show that plant and animal populations experience divergent selection between urban and nonurban environments. This divergent selection has led to adaptive evolution in life history, morphology, physiology, behavior, and reproductive traits. These adaptations typically evolve in response to pesticide use, pollution, local climate, or the physical structure of cities. Despite these important results, the genetic basis of adaptive evolution is known from only a few cases. Most studies also examine only a few populations in one city, and experimental validation is rare.
OUTLOOK
The study of evolution in urban areas provides insights into both fundamental and applied problems in biology. The thousands of cities throughout the world share some features while differing in other aspects related to their age, historical context, governmental policies, and local climate. Thus, the phenomenon of global urbanization represents an unintended but highly replicated global study of experimental evolution. We can harness this global urban experiment to understand the repeatability and pace of evolution in response to human activity. Among the most important unresolved questions is, how often do native and exotic species adapt to the particular environmental challenges found in cities? Such adaptations could be the difference as to whether a species persists or vanishes from urban areas. In this way, the study of urban evolution can help us understand how evolution in populations may contribute to conservation of rare species, and how populations can be managed to facilitate the establishment of resilient and sustainable urban ecosystems. In a similar way, understanding evolution in urban areas can lead to improved human health. For example, human pests frequently adapt to pesticides and evade control efforts because of our limited understanding of the size of populations and movement of individuals. Applied evolutionary studies could lead to more effective mitigation of pests and disease agents. The study of urban evolution has rapidly become an important frontier in biology, with implications for healthy and sustainable human populations in urban ecosystems.

A gradient in urbanization showing the skyline of Canada’s sixth largest city (Mississauga, Canada) on the horizon, and the Credit Valley and the University of Toronto Mississauga campus in the foreground.
PHOTO CREDIT: ARJUN YADAV
Abstract
Our planet is an increasingly urbanized landscape, with over half of the human population residing in cities. Despite advances in urban ecology, we do not adequately understand how urbanization affects the evolution of organisms, nor how this evolution may affect ecosystems and human health. Here, we review evidence for the effects of urbanization on the evolution of microbes, plants, and animals that inhabit cities. Urbanization affects adaptive and nonadaptive evolutionary processes that shape the genetic diversity within and between populations. Rapid adaptation has facilitated the success of some native species in urban areas, but it has also allowed human pests and disease to spread more rapidly. The nascent field of urban evolution brings together efforts to understand evolution in response to environmental change while developing new hypotheses concerning adaptation to urban infrastructure and human socioeconomic activity. The next generation of research on urban evolution will provide critical insight into the importance of evolution for sustainable interactions between humans and our city environments.
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Science
Volume 358 | Issue 6363
3 November 2017
3 November 2017
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Copyright © 2017, American Association for the Advancement of Science.
This is an article distributed under the terms of the Science Journals Default License.
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Published in print: 3 November 2017
Acknowledgments
We thank A. Yadav for permission to use the photo in the summary, B. Cohan for the artwork in Fig. 2, and J. Beninde for donating the wall lizard photo. All other art was original or obtained under creative commons licenses. The paper was improved by three anonymous reviewers, D. Carmona, M. Combs, C. Fitzpatrick, P. Grant, M. Hetherington-Rauth, J. Losos, E. Puckett, R. Rivkin, J. Santangelo, and K. Thompson. We thank L. Miles and B. Verrelli for identifying additional studies included in table S1. This paper was supported by grants from the National Science Foundation (NSF DEB 1457523) to J. Munshi-South and Natural Sciences and Engineering Research Council of Canada to M. Johnson.
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Funding Information
National Science Foundation: award351416, 1457523
Natural Sciences and Engineering Research Council of Canada: award351421, 491174
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