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Andres
Species in urban settings are evolving along subtly different lines than their rural counterparts, researchers are finding. An example is the white
clover plant, shown here being sampled in Mexico City. The plant has evolved to produce far less hydrogen cyanide in urban environments than
in rural ones. CREDIT: DIEGO CARMONA
Urban evolution: How species adapt
to survive in cities
Plants and animals are evolving in cities around the world — offering ways to study
longstanding scientific questions and clues to where climate change is taking us
By Eric Bender 03.21.2022
From Knowable Magazine (from Annual Reviews)
B rown rats in New York City may be evolving smaller rows of teeth. Tiny fish across the
Eastern US have adapted to thrive in polluted urban waters. Around the globe, living things
are evolving differently in cities than in the surrounding countryside.
3inYea
It’s happening in plants: White clover in downtown Toronto is less likely than clover in
surrounding rural areas to produce a cyanide that deters herbivores — a trend mirrored in cities
in many countries, a new study finds. And it’s going on in birds: Songbirds in Europe and owls
in Argentina show evidence of natural selection in genes associated with cognition.
penin
All are examples of urban evolution: genetic changes that may help living things adapt to life
explci in big city environments. “A city changes an environment dramatically. It creates a completely
positive connotation
novel ecosystem,” says Marc Johnson, an evolutionary ecologist at the University of Toronto
Mississauga.
The city is also the fastest-growing ecosystem on the planet, home to more than half of the
world’s people. So perhaps it’s no surprise that studying the evolution of species in urban settings,
a field that barely existed at the start of the millennium, now is a focus for many biology labs.
Cities can act as test beds to address longstanding questions in evolution. Do different
populations of the same species evolve in similar ways when faced with the same environmental
pressures? And do different species in the same locations evolve similar characteristics?
Many environmental factors are similar across thousands of cities, says Johnson: things like
higher temperatures, pollution and habitats fragmented by buildings and roads. But cities also
differ in age, amount of green space, climate and more.
“You can look at these similarities and these differences and start to ask, how can this drive
evolution?” Johnson says.
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A city rat enjoys a tasty snack. In New York City, brown rats in different neighborhoods have accrued genetic differences. The populations
appear to be kept apart by intervening areas with more intense rat control and less human food to feast on.
CREDIT: ERNIE JANES / ALAMY STOCK PHOTO
idea
Main
Observing how creatures respond to urban life also may help to improve conservation -
management or pest control, and to plan cities with functioning ecosystems that are
-
environmentally more robust and better places for people to live.
And urban evolution may hold hints about our future world. “Cities are kind of the key for
understanding responses to global climate change,” says Sarah Diamond, an evolutionary
ecologist at Case Western Reserve University in Cleveland, Ohio, and coauthor of an article on
urban evolution research in the Annual Review of Ecology, Evolution, and Systematics. “You
can step through time. You can say, ‘This city is giving you the global climate warming that we
would expect by 2050 or 2070 or 2100.’”
People often feel that city life is removed from nature, says Colin Garroway, an evolutionary
ecologist at the University of Manitoba in Winnipeg. “But cities are nature.” - >
she is arguing for
Looking out our back doors citics
E
excess adapted Probably the best-known example of urban evolution is the English peppered moth whose
coloration darkened in the 19th century in response to coal pollution. In a famous 1955 paper,
to its environment
British geneticist Bernard Kettlewell presented evidence that this was a case of natural selection
in which darkness helped the moths evade bird predation as they rested on sooty tree trunks.
Rapid in
3
crese
But the field of urban evolutionary ecology remained tiny until recently: “Most evolutionary
is Resecrat s
biologists would not be caught dead in a city,” says Johnson. That began to change with the rapid
new urban like
growth of urban ecology studies in the 1990s and accelerated with discoveries of surprisingly
CKKIKH Aninch.
quick cases of evolution, such as Caribbean lizard populations that displayed larger toepad area,
crucial for clinging to surfaces, after two major hurricanes in 2017.
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It didn’t hurt that professors of evolutionary biology generally are employed in urban universities
and curious about what is happening in their backyards. “These dynamics are happening all
around you,” says Ryan Martin, an evolutionary ecologist at Case Western Reserve and coauthor
with Diamond of the Annual Reviews article. “Go out and look in your garden, and you’ll
see a bunch of native pollinators that are all presumably evolving in response to these changes shows that
in the city…. You don’t have to do anything special to see these cool dynamics; you walk out evolution is everywhere
- -
-
your door.” in cities.
In industrial parts of England, black-bodied forms of the peppered moth Biston betularia became more abundant as air pollution increased
after the Industrial Revolution, blackening trees and buildings. The frequency of black moths decreased again when the air became cleaner.
CREDIT: CHISWICK CHAP / WIKIMEDIA COMMONS
S
The water flea Daphnia magna — a freshwater crustacean up to a few millimeters in size — is
positiveevolute
ne
one species busy evolving in cities in response to heat, pollution and even local predators. These
are good.
zooplankton can prevent algal blooms that overload ponds with toxic cyanobacteria, so this
adaptation may have a big effect on freshwater ecosystems, says Kristien Brans, an evolutionary
ecologist at KU Leuven in Belgium, who studies the water fleas.
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One basic challenge in such urban investigations is to distinguish between two modes of
response to altered environments: evolution (genetic alterations that appear across generations)
and phenotypic plasticity (the flexibility to alter physical and/or behavioral characteristics in an
organism’s lifetime).
E
For water fleas, it turns out that both are at play. Fleas raised in lab experiments at temperatures
Shows how Achange
matching urban ponds are smaller, and mature and reproduce more quickly, than fleas reared
environment
in
at rural pond temperatures that tend to be several degrees cooler. (That’s phenotypic plasticity
only
not
Alters but
physicale — no genetic changes have occurred.) But over time, urban water fleas living generation after
generation in warmer, urban pond waters have genetically changed to have those same kinds of
evolutions
alterations. (That’s evolution.)
In a 2017 paper, for example, Brans and her coworkers took populations of water fleas from overall better
3
a range of habitats — some more rural and some more urban — and reared them for many eudlution for the
generations before testing their ability to survive in urban-temperature water and rural- Heg
·
is they
temperature water. Fleas collected from urban ponds displayed higher heat tolerance in the warm are able to handle
ponds than those collected from rural ponds, along with smaller body size and other changes. environment
tougher
better.
A follow-up study published in 2018 showed that urban Daphnia have significantly higher
concentrations than rural water fleas of total body fat, proteins and sugars, trait changes that are
associated with handling stresses such as heat as well as with faster life cycles.
Scientists studying the water flea Daphnia magna in rural and urban ponds have identified gene-based differences in traits such as tolerance
to water temperature. CREDIT: HAJIME WATANABE FLICKR
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Brans and coworkers have also recently found that urban water fleas are more likely than their
rural cousins to survive exposure to a common pesticide, and that populations of Daphnia display
different genetic adaptions to pesticides depending on whether they grow in ponds surrounded
by conventional farms, organic farms or nature reserves. In lab tests, water fleas taken from
ponds surrounded by conventional farmland displayed higher resistance to a pesticide called
chlorpyrifos that’s routinely employed in such farming. Fleas near organic farms were more
resistant to two pesticides allowed in organic agriculture.
E
Looking up the food chain, Brans and colleagues have evidence that urban water fleas
Althoughtheyare and predatory insects that eat them — damselflies — are evolving in step with each other.
Urban damselfly larvae are far better than rural damselfly larvae at encountering and gobbling
dres
equal which e
up rural water fleas, for example. But they have a tougher time preying on the urban fleas.
nch disrupt the -
-
In other words, when rural or city damselfly and flea populations are matched, there seems
Ford chain
to be more balance — as you’d expect if two populations are evolving in step with each other.
-
Brans also is studying how the microbes that live in Daphnia guts differ between city and
countryside. These microbial communities — or microbiome — shape what the water fleas
can eat, and some flea genotypes encourage microbiomes that enable fleas to digest toxic
cyanobacteria that can overrun ponds.
- -
-
Better overall
Adapting successfully, or maybe not
Acorn ants offer another case of adaptive urban evolution. With colonies so tiny they can live
inside a single acorn, they are easy to study. (“Put them in a little plastic cup with some sugar
water and a little dead mealworm and they’re totally happy,” Martin says.) Colonies in Cleveland,
Ohio — whose downtown temperatures average about 4 degrees Celsius warmer year-round
than the rural surroundings — have higher heat tolerances but lower cold tolerances than rural
ants, Martin and Diamond found. “We’re pretty confident that it’s due to underlying genetic
differences,” Martin says.
Brown rats in Manhattan offer yet another case of urban evolution, though it may not impart
advantages to the unloved creatures. Jason Munshi-South, an evolutionary ecologist at Fordham
University in New York, and colleagues analyzed the genomes of 262 rats and found that the
animals have evolved distinct genomic profiles in different neighborhoods. The scientists believe
it’s because the rat populations don’t move freely between these spots, and slowly, over time,
accrue differences.
What’s keeping them apart? Midtown Manhattan may act as a kind of soft barrier between
Lower and Upper Manhattan, the scientists say, because it is less residential (providing less food)
and the site of intense rat control efforts. Roads and waterways also can genetically split up rat
populations, according to studies in New Orleans, Salvador in Brazil and Vancouver in Canada,
where rats also show genetic variations by neighborhoods.
Such insights may prove useful in designing measures to suppress rat populations. “If you
understand how rats move around and what facilitates or prevents their movement, you can
break the city down into more manageable units for rodent control,” Munshi-South says.
Other changes in rats may be adaptive. Munshi-South’s lab has evidence that natural selection
-
is changing the skulls of the rats such that they have longer noses and shorter sets of teeth.
-
These might be adaptations to colder environments and a diet of human leftovers respectively,
the scientists speculate. Similar changes in teeth have been spotted in urban white-footed mice,
so this might be a general phenomenon in rodents in cities, Munshi-South says.
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In the clover
Urban plants are on the genetic move too — such as white clover, a perennial plant that thrives
in human landscapes. The plant, due to the activity of two known genes, can produce hydrogen
cyanide, if it invests the resources to do so. This protects it from browsing herbivores.
-
Sampling the plants from the center of Toronto out to surrounding rural areas, Johnson’s lab
discovered a striking inherited correlation: The closer to the center, the less cyanide gets
-
produced. Johnson and his colleagues suggest this happens because the center is colder in winter,
u
due to less snow cover, and plants that make hydrogen cyanide are more susceptible to freezing.
(His lab found generally similar results in several dozen other North American cities.)
To delve more deeply into urban evolution, a few years back Johnson and colleagues launched
the Global Urban Evolution Project (GLUE), bringing together 287 scientists in 26 countries.
(Many responded to tweets Johnson sent out while pursuing another project in the Galapagos.)
“GLUE is the largest collaborative study in evolutionary biology ever attempted, if you don’t
include the human genome project,” Johnson says.
GLUE took white clover’s cyanide production as a model to study three questions. Do instances
of urbanization in different cities lead to similar local environments? Do those similar
environments lead the clover to evolve along the same lines — display parallel evolution — in
a trait of interest (in this case, cyanide production)? And if so, what environmental factors are
driving the pattern?
Je
In a new Science paper, the collaborators showed that urban environments do indeed end up
quite similar to each other, with less vegetation, more impervious surfaces and higher summer
temperatures than their outlying rural areas. (In fact, downtowns of cities such as Beijing and
Boston are more similar to each other in such factors than they are to their rural areas, Johnson
comments.) Analyzing more than 110,000 clover plants from 160 cities in 26 countries, the
GLUE investigators also demonstrated a strong link between urbanization and clover cyanide
production. And after sequencing more than 2,000 clover genomes and analyzing the urban-rural
differences, the researchers showed that natural selection truly is at work.
But what are the environmental factors driving this change in cyanide? “The answer is pretty
complicated,” Johnson says, and may not be the same for all cities. The most important ones
the team uncovered were changes in overall vegetation (probably related to the abundance
and diversity of herbivores that eat clover) and the aridity of the environment. “We don’t see
temperature clearly popping out, which is what we had identified when we looked at Boston,
Toronto, Montreal and New York,” he says.
The first GLUE results show that white clover is a powerful global model for understanding
evolution and ecology in response to urbanization, he adds.
Disparities within cities
E
research that As researchers continue to study evolution in the big city, some are focusing on the effects of
Nuanced
in to consideration social and economic inequality. The question, says Simone Des Roches, an evolutionary ecologist
takes
at the University of Seattle in Washington, is whether plants and animals evolve differently in
neighbouring
Economy
low- versus high-income neighborhoods. Lead author on a 2020 paper on the interaction of social,
ecological and evolutionary dynamics in cities, Des Roches notes that racial discrimination in the
United States has produced strikingly different urban environments.
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The evolution of a species in a city involves an array of interacting factors, as shown here for mosquitoes. Factors can be social (blue),
evolutionary (red) or ecological (green). In the case of mosquitoes, these interactions are important to understand due to the ability
of the insects to carry dangerous pathogens such as the malaria parasite and the viruses causing Zika and West Nile disease.
CREDIT: S. DES ROCHES ET AL / EVOLUTIONARY APPLICATIONS 2020
Impoverished neighborhoods tend to have higher temperatures, greater exposure to pollutants
and other environmental disadvantages. These can act as playgrounds for disease-carrying pests
such as mosquitoes and rats that enjoy human company: Invasive tiger mosquitoes grow larger in
neighborhoods with abandoned buildings in Baltimore, for example. Researchers want to know if,
and how, organisms may evolve differently in these disadvantaged environments. -
Urban evolution studies also can shed light on what lies ahead in this time of the Anthropocene
and suggest steps that might achieve a friendlier world for humans and other forms of life. For
example, in many cities, Diamond says, scientists can date the onset of high levels of warming
from industrialization. Researchers then can measure how much a species’ heat and cold
tolerances have changed over time, infer the rate of evolution of those traits and apply those
inferences to predict how life forms will respond to future climate change.
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Diamond’s work in acorn ants suggests that rural populations may be able to evolve to take the
greater heat. But, she says, urban acorn ants appear less well-adapted to cities than the rural ants
are adapted to their ancestral homes.
Brans, meanwhile, looks to apply her research to preserve urban biodiversity and public health —
since urban conservation managers will want to see ponds growing healthy populations of water
fleas that bolster those ecosystems against toxic algae blooms.
Unfortunately, the genetic biodiversity that can fuel adaptation often dwindles in urban areas.
A genetic survey by Chloé Schmidt working in Garroway’s lab, for example, found this to be
the case, along with lower population sizes, for North American mammals living in more
disturbed environments. That’s a concern during a period when so many populations of animals
-
and plants are seeing their natural habitats degraded or simply destroyed.
-
Scientists don’t take urban environments as precise models for the impacts of climate change.
But they say such studies will provide important clues to how creatures may respond to
dwindling access to water and food, and exposure to pollution, heat, drought and other dangers.
“We’re in the Anthropocene, and we don’t understand how we’re changing the environment on
every level, from greenhouse gas emissions to changing the evolution of life around us,” Johnson
says. “People realize this research is part of the solution.”
10.1146/knowable-031822-1
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