Urban evolution refers to the heritable genetic changes of populations in response to urban development and anthropogenic activities in urban areas. Urban evolution can be caused by mutation, genetic drift, gene flow, or evolution by natural selection. [1] Biologists have observed evolutionary change in numerous species compared to their rural counterparts on a relatively short timescale. [1] [2]
Strong selection pressures due to urbanization play a big role in this process. The changed environmental conditions lead to selection and adaptive changes in city-dwelling plants and animals. [3] [2] Also, there is a significant change in species composition between rural and urban ecosystems. [4]
Shared aspects of cities worldwide also give ample opportunity for scientists to study the specific evolutionary responses in these rapidly changed landscapes independently. How certain organisms (are able to) adapt to urban environments while others cannot, gives a live perspective on rapid evolution. [3] [2]
With urban growth, the urban-rural gradient has seen a large shift in distribution of humans, moving from low density to very high in the last millennia. This has brought a large change to environments as well as societies. [5]
Urbanization transforms natural habitats to completely altered living spaces that sustain large human populations. Increasing congregation of humans accompanies the expansion of infrastructure, industry and housing. Natural vegetation and soil are mostly replaced or covered by dense grey materials. Urbanized areas continue to expand both in size and number globally; in 2018, the United Nations estimated that 68% of people globally will live in ever-larger urban areas by 2050. [6]
Urbanization intensifies diverse stressors spatiotemporally such that they can act in concert to cause rapid evolutionary consequences such as extinction, maladaptation, or adaptation. [7] Three factors have come to the forefront as the main evolutionary influencers in urban areas: the urban microclimate, pollution, and urban habitat fragmentation. [8] These influence the processes that drive evolution, such as natural and sexual selection, mutation, gene flow and genetic drift.
A microclimate is defined as any area where the climate differs from the surrounding area. Modifications of the landscape and other abiotic factors contribute to a changed climate in urban areas. The use of impervious dark surfaces which retain and reflect heat, and human generated heat energy lead to an urban heat island in the center of cities, where the temperature is increased significantly. A large urban microclimate does not only affect temperature, but also rainfall, snowfall, air pressure and wind, the concentration of polluted air, and how long that air remains in the city. [9] [10] [11]
These climatological transformations increase selection pressure. [12] Certain species have shown to be adapting to the urban microclimate. [3] [2]
Many species have evolved over macroevolutionary timescales by adapting in response to the presence of toxins in the environment of the planet. Human activities, including urbanization, have greatly increased selection pressures due to pollution of the environment, climate change, ocean acidification, and other stressors. Species in urban settings must deal with higher concentrations of contaminants than naturally would occur. [13] [14]
There are two main forms of pollution which lead to selective pressures: energy or chemical substances. Energy pollution can come in the form of artificial lighting, sounds, thermal changes, radioactive contamination and electromagnetic waves. Chemical pollution leads to the contamination of the atmosphere, the soil, water and food. All these polluting factors can alter species’ behavior and/or physiology, which in turn can lead to evolutionary changes. [15]
The fragmentation of previously intact natural habitats into smaller pockets which can still sustain organisms leads to selection and adaptation of species. These new urban patches, often called urban green spaces, come in all shapes and sizes ranging from parks, gardens, plants on balconies, to the breaks in pavement and ledges on buildings. The diversity in habitats leads to adaptation of local organisms to their own niche. [16] And contrary to popular belief, there is higher biodiversity in urban areas than previously believed. This is due to the numerous microhabitats. These remnants of wild vegetation or artificially created habitats with often exotic plants and animals all support different kinds of species, which leads to pockets of diversity inside cities. [17]
With habitat fragmentation also comes genetic fragmentation; genetic drift and inbreeding within small isolated populations results in low genetic variation in the gene pool. Low genetic variation is generally seen as bad for chances of survival. This is why probably some species aren’t able to sustain themselves in the fragmented environments of urban areas. [18]
The differing urban environment imposes different selection pressures than the natural setting. [7] These stressors elicit phenotypic changes in populations of organisms which may be due to phenotypic plasticity—the ability of individual organisms to express different phenotypes from the same genotype as a result of exposure to different environmental conditions--or actual genetic changes.
In considering the examples of urban evolution, observed phenotypic divergences or differences in responses to urbanization have to be genetically based and adaptive (increase fitness in that particular environment) to be tagged as evolution and adaptation, respectively. Hence, it will be appropriate to consider neutral/non-adaptive and adaptive urban evolution, with the later needing to be sufficiently proven. [7]
Although there is widespread agreement that adaptation is occurring in urban populations, as of 2021 [update] there are almost no proven examples – almost all are cases of selection, reasoned speculation connecting to adaptive benefit, but no evidence of actual adaptive phenotype. [7] At this time only six examples are demonstrated:
Some interesting cases of possible adaptation which remain insufficiently proven are:
Claimed examples of urban adaptation include:
In one case selection is widely expected to occur and yet is not found:
Evolution is the change in the heritable characteristics of biological populations over successive generations. Evolution occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certain characteristics becoming more or less common within a population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation.
Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within lineages. Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book On the Origin of Species. He also identified sexual selection as a likely mechanism, but found it problematic.
Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and shifts in allele frequencies. A small population is then more susceptible to demographic and genetic stochastic events, which can impact the long-term survival of the population. Therefore, small populations are often considered at risk of endangerment or extinction, and are often of conservation concern.
Population genetics is a subfield of genetics that deals with genetic differences within and among populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and population structure.
Urban ecology is the scientific study of the relation of living organisms with each other and their surroundings in an urban environment. An urban environment refers to environments dominated by high-density residential and commercial buildings, paved surfaces, and other urban-related factors that create a unique landscape. The goal of urban ecology is to achieve a balance between human culture and the natural environment.
In population genetics, gene flow is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent allele frequencies and therefore can be considered a single effective population. It has been shown that it takes only "one migrant per generation" to prevent populations from diverging due to drift. Populations can diverge due to selection even when they are exchanging alleles, if the selection pressure is strong enough. Gene flow is an important mechanism for transferring genetic diversity among populations. Migrants change the distribution of genetic diversity among populations, by modifying allele frequencies. High rates of gene flow can reduce the genetic differentiation between the two groups, increasing homogeneity. For this reason, gene flow has been thought to constrain speciation and prevent range expansion by combining the gene pools of the groups, thus preventing the development of differences in genetic variation that would have led to differentiation and adaptation. In some cases dispersal resulting in gene flow may also result in the addition of novel genetic variants under positive selection to the gene pool of a species or population
Evolvability is defined as the capacity of a system for adaptive evolution. Evolvability is the ability of a population of organisms to not merely generate genetic diversity, but to generate adaptive genetic diversity, and thereby evolve through natural selection.
Biological dispersal refers to both the movement of individuals from their birth site to their breeding site, as well as the movement from one breeding site to another . Dispersal is also used to describe the movement of propagules such as seeds and spores. Technically, dispersal is defined as any movement that has the potential to lead to gene flow. The act of dispersal involves three phases: departure, transfer, settlement and there are different fitness costs and benefits associated with each of these phases. Through simply moving from one habitat patch to another, the dispersal of an individual has consequences not only for individual fitness, but also for population dynamics, population genetics, and species distribution. Understanding dispersal and the consequences both for evolutionary strategies at a species level, and for processes at an ecosystem level, requires understanding on the type of dispersal, the dispersal range of a given species, and the dispersal mechanisms involved. Biological dispersal can be correlated to population density. The range of variations of a species' location determines expansion range.
In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the population during that process. Thirdly, it is a phenotypic trait or adaptive trait, with a functional role in each individual organism, that is maintained and has evolved through natural selection.
In evolutionary biology, the Baldwin effect describes an effect of learned behaviour on evolution. James Mark Baldwin and others suggested that an organism's ability to learn new behaviours will affect its reproductive success and will therefore have an effect on the genetic makeup of its species through natural selection. It posits that subsequent selection might reinforce the originally learned behaviors, if adaptive, into more in-born, instinctive ones. Though this process appears similar to Lamarckism, that view proposes that living things inherited their parents' acquired characteristics. The Baldwin effect only posits that learning ability, which is genetically based, is another variable in / contributor to environmental adaptation. First proposed during the Eclipse of Darwinism in the late 19th century, this effect has been independently proposed several times, and today it is generally recognized as part of the modern synthesis.
Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological processes that slowly alter the layout of the physical environment, and human activity such as land conversion, which can alter the environment much faster and causes the extinction of many species. More specifically, habitat fragmentation is a process by which large and contiguous habitats get divided into smaller, isolated patches of habitats.
The gene-centered view of evolution, gene's eye view, gene selection theory, or selfish gene theory holds that adaptive evolution occurs through the differential survival of competing genes, increasing the allele frequency of those alleles whose phenotypic trait effects successfully promote their own propagation. The proponents of this viewpoint argue that, since heritable information is passed from generation to generation almost exclusively by DNA, natural selection and evolution are best considered from the perspective of genes.
Evidence of common descent of living organisms has been discovered by scientists researching in a variety of disciplines over many decades, demonstrating that all life on Earth comes from a single ancestor. This forms an important part of the evidence on which evolutionary theory rests, demonstrates that evolution does occur, and illustrates the processes that created Earth's biodiversity. It supports the modern evolutionary synthesis—the current scientific theory that explains how and why life changes over time. Evolutionary biologists document evidence of common descent, all the way back to the last universal common ancestor, by developing testable predictions, testing hypotheses, and constructing theories that illustrate and describe its causes.
The white-footed mouse is a rodent native to North America from Ontario, Quebec, Labrador, and the Maritime Provinces to the southwestern United States and Mexico. In the Maritimes, its only location is a disjunct population in southern Nova Scotia. It is also known as the woodmouse, particularly in Texas.
Ecology and evolutionary biology is an interdisciplinary field of study concerning interactions between organisms and their ever-changing environment, including perspectives from both evolutionary biology and ecology. This field of study includes topics such as the way organisms respond and evolve, as well as the relationships among animals, plants, and micro-organisms, when their habitats change. Ecology and evolutionary biology is a broad field of study that covers various ranges of ages and scales, which can also help us to comprehend human impacts on the global ecosystem and find measures to achieve more sustainable development.
Genetic assimilation is a process described by Conrad H. Waddington by which a phenotype originally produced in response to an environmental condition, such as exposure to a teratogen, later becomes genetically encoded via artificial selection or natural selection. Despite superficial appearances, this does not require the (Lamarckian) inheritance of acquired characters, although epigenetic inheritance could potentially influence the result. Waddington stated that genetic assimilation overcomes the barrier to selection imposed by what he called canalization of developmental pathways; he supposed that the organism's genetics evolved to ensure that development proceeded in a certain way regardless of normal environmental variations.
The medium ground finch is a species of bird in the family Thraupidae. It is endemic to the Galapagos Islands. Its primary natural habitat is tropical shrubland. One of Darwin's finches, the species was the first which scientists have observed evolving in real-time.
A genetic isolate is a population of organisms with little genetic mixing with other organisms within the same species due to geographic isolation or other factors that prevent reproduction. Genetic isolates form new species through an evolutionary process known as speciation. All modern species diversity is a product of genetic isolates and evolution.
Forest migration is the movement of large seed plant dominated communities in geographical space over time.
In evolutionary biology, developmental bias refers to the production against or towards certain ontogenetic trajectories which ultimately influence the direction and outcome of evolutionary change by affecting the rates, magnitudes, directions and limits of trait evolution. Historically, the term was synonymous with developmental constraint, however, the latter has been more recently interpreted as referring solely to the negative role of development in evolution.