UNIVERSITY OF SOUTHERN MINDANAO

EVOLUTION IN
SPACE

Bio 221a- Evolutionary Biology

Mangaoang & Manceras, 2021
 Topic Outline
- Patterns in Space
- Gene flow
- Genetic Divergence between populations
- Gene flow and selection
- Gene flow and drift
- The evolution of dispersal
- The evolution of species’ ranges

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 Intended Learning Outcomes

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 Patterns in Space
White clover (Trifolium repens) has
a smooth gradient (cline) in the
frequency of a gene needed to
produce cyanide, which protects
the plant from herbivores. The
white portion of each pie diagram
shows the fraction of the
population in which the gene
CYP79D15 has been deleted.
Frequencies of deletions are much
higher in populations growing in
cold climates. The solid curve
shows the 0°C isotherm for
January.

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 • Cline
- A smooth change in space of a trait mean or an
allele frequency
- A very common pattern
- Clines in body size can be hundreds or even
thousands of kilometers long.

Bio 221a- Evolutionary Biology

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 • Moose (Alces alces) are larger in the north of
Sweden than in the south. This cline is an example
of Bergmann’s rule, which says that the body
sizes of mammals and birds tend to increase with
distance from the equator.
Bio 221a- Evolutionary Biology
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 A grass called common bent
(Agrostis capillaris) grows on
and around an abandoned
surface mine in Wales. Soil
on the mine has high
concentrations of copper,
and plants growing there
have evolved tolerance of
this toxic element.

Bio 221a- Evolutionary Biology

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 Remember this….
• Most species show geographic variation in allele
frequencies and the means of phenotypic traits.
Clines, which are smooth changes in an allele
frequency or trait mean, are a very common
pattern.
• Clines and other patterns can result from local
adaptation, which results when selection varies
in space.

Bio 221a- Evolutionary Biology

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 Gene Flow
• The solitary and
gregarious forms of
the desert locust
(Schistocerca gregaria,
shown at left) differ in
color, morphology,
and behavior. High
population densities
cause individuals to
develop into the
gregarious form,
which gathers in huge
swarms that disperse
long distances

Bio 221a- Evolutionary Biology

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 Roles of Gene flow in
Evolution
• Equalizes allele frequencies
• Introduce new alleles into a population

Bio 221a- Evolutionary Biology

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 How is gene flow measured?
• Migration rate
- how gene is quantified
- Symbolized by m
- The fraction of individuals in a population that
arrives from another population in each
generation.

Bio 221a- Evolutionary Biology

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 Where:

- change in allele frequency

- the allele’s frequency in the migrants
p - its frequency in the focal population before
migration
m – migration rate
Bio 221a- Evolutionary Biology
Mangaoang & Manceras, 2021
 Remember this…..

• Gene flow is the mixing of alleles from different

populations, eroding differences caused by selection
and drift. It results from the dispersal of individuals
and their gametes. Gene flow is measured by the
migration rate (m) when populations are discrete or
patchy, and by the migration variance (σm2) when
populations are continuously distributed in space.

Bio 221a- Evolutionary Biology

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 Genetic Divergence between
Populations
FST is a statistic used to measure
genetic differences between two
or more populations. In this
schematic, two alleles at a locus
are represented by red and blue
circles. Top: FST = 0 when allele
frequencies are equal in the
populations. Middle: FST = 0.36
when allele frequencies are 0.2
and 0.8 in the two populations.
Bottom: FST = 1 when the
populations are fixed for different
alleles
Bio 221a- Evolutionary Biology
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 Isolation-by-distance

• Isolation-by-distance in human populations. The horizontal axis

has been corrected for large bodies of water that could not be
crossed when humans first spread across Earth. The vertical axis
gives FS estimated from 783 loci. Each dot represents a
comparison between a pair of populations from the indicated
region(s).
Bio 221a- Evolutionary Biology
Mangaoang & Manceras, 2021
 Remember this……
• FST is a statistic commonly used to describe
genetic divergence between two or more
populations. In many species with broad
geographic ranges, FST increases with the
distance between two populations, a pattern
called isolation-by distance. FST varies across
the genome, and genomic regions with high
FST can be used to find loci that are locally
adapted.

Bio 221a- Evolutionary Biology

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 Gene Flow and Selection

A grass called common bent

(Agrostis capillaris) grows on
and around an abandoned
surface mine in Wales. Soil
on the mine has high
concentrations of copper,
and plants growing there
have evolved tolerance of
this toxic element.

Bio 221a- Evolutionary Biology

Mangaoang & Manceras, 2021
• The rock pocket mouse
(Chaeto- Light rock Dark lava
Light rock Dipus intermedius)
typically has light-colored fur
where it lives on light-
colored granite, but is dark
colored where it lives on
dark lava flows in Arizona.
The dark coloration, which
results from a melanic allele
at the Mc1r locus,
camouflages mice on the
lava from predators.
Selection favoring the
melanic allele on the lava is
intense, with a selection
coefficient estimated to be
as high as s = 0.4.
Bio 221a- Evolutionary Biology
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 • When both gene flow and local selection are at
work, allele frequencies evolve toward a
compromise between them.
• If gene flow is weak relative to selection, allele
frequencies will evolve to what selection favors
at each location.
• If gene flow is relatively strong, allele frequencies
will be equalized.
• Strong gene flow can cause gene swamping,
which is when a locally favored allele is lost
because migration overwhelms local selection.
Bio 221a- Evolutionary Biology
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 • In continuous habitats, the widths of clines
are determined by the ratio of the migration
variance to the strength of local selection.
• When there is a patch of habitat that selects
for a different allele than that favored
outside the patch, the locally adapted allele
will be lost by gene swamping if the size of
the patch is smaller than a critical size
determined by the relative strengths of
migration and selection.
Bio 221a- Evolutionary Biology
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 Tension zones

• Tension zones are clines

that form when there is
selection against
heterozygotes.

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 • Tension zones are clines in allele
frequencies that result from selection
against heterozygotes (underdominance)
that acts uniformly in space.

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 • Drift can cause allele frequencies at selectively
neutral loci to diverge between populations. Very
small rates of migration prevent divergence at
neutral loci. The amount of divergence can be
used to estimate the amount of gene flow.

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 Evolution of Dispersal
• Factors that favors dispersal
a. Allows individuals to find habitat that is
better now or that will be in the future.
b. Competition with relatives
c. Another kind of interaction between
individuals

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 • Factors that select against dispersal
a. Passive dispersal can land an individual in
hostile habitat
b. Active dispersal is also dangerous when
patches of good habitat are separated by
regions of bad habitat.
c. Energetic trade-offs

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 • The Barton Springs salamander (Eurycea sosorum) has lost
the terrestrial phase of its life cycle. It cannot survive on land
and so is unable to move to another spring. This species is
found only in the aquifer that feeds a group of springs in
Austin, Texas, that provide water to a large swimming pool
(inset). The salamander’s entire range is only a few square
kilometers.
Bio 221a- Evolutionary Biology
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 • Roesel's bush-cricket (Metrioptera roeselii) has two
morphs, one with normal wings (left) and one with
highly reduced wings (right). The morph with normal
wings can disperse long distances by flying, but has
lower fecundity because of the energy it invests in the
growth and maintenance of flight muscles.
Bio 221a- Evolutionary Biology
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 When a species’ range expands, the most
rapidly dispersing genotypes automatically
become more common at the range edge,
causing the rate of spread to increase.

Bio 221a- Evolutionary Biology

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 Remember this……
• Dispersal rates evolve.
• Higher dispersal is favored by habitat
disturbance that causes extinction of local
populations, competition between related
individuals, and inbreeding.
• Lower dispersal is favored because
movement is often risky and energetically
expensive.
• In a species that is expanding its range, there
is an automatic increase at the range’s edge
of alleles that enhance dispersal.
Bio 221a- Evolutionary Biology
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 The Evolution of Species’
Ranges
• Possible explanation on species’ ranges limit
a. Populations may simply lack genetic variation in a
trait necessary for adapting to a new environment.
b. Gene swamping caused by migration from other
parts of a species’ range can prevent local
adaptation to the extreme conditions at the range
edge and prevent the species from expanding
outward
c. biotic interactions can set range boundaries where
a species encounters a new competitor, predator,
or pathogen.

Bio 221a- Evolutionary Biology

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 How species richness
respond to environmental?
• By changing where they live
• By adapting to the new conditions.

Bio 221a- Evolutionary Biology

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 • Species living along an environmental gradient
(for example, correlated with latitude) can
respond to climate change in several ways.
These cartoons depict a species living in the
northern hemisphere. ( Figure next slide)

Bio 221a- Evolutionary Biology

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Bio 221a- Evolutionary Biology
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 Remember this……
• Species ranges evolve.
• Factors that prevent ranges from expanding
outward include dispersal barriers, genetic
constraints and gene flow that prevents
adaptation to more extreme environments, and
competition with other species that have
adjacent ranges.
• Global climate change is causing shifts in the
ranges of many species, but there is little
evidence that species can generally avoid
extinction by adapting to the new conditions.

Bio 221a- Evolutionary Biology

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 References
Futuyma, D.J. and Kirkpatrick, M. Evolution,
4th Edition. Pp 191-211

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