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Barriers to Gene Flow

• Whether or not a physical barrier
deters gene flow depends upon:
 Organism’s mode of dispersal or
 Duration of time organism can move
Genetic Drift in
Snail Populations

 Robert Selander studied Helix
 Collected snails from a two-
block area
 Analyzed the allele frequencies
for five genes
Genetic Divergence in
Snail Populations
Snail Speciation?
 Will the time come when the snails
from opposite sides of the street are
so different that they can no longer

 If so, then they will have become two
distinct species
Speciation & Natural Selection

 Natural selection can lead to
 Speciation can also occur as a result
of other microevolutionary processes
 Genetic drift
 Mutation
 Sexual Selection
Morphology & Species

 Morphological traits may not be useful in
distinguishing species
 Members of same species may appear
different because of environmental conditions
 Morphology can vary with age and sex
 Different species can appear identical
Darwin's Explanatory Model of
Evolution Through Natural
 Refer to Excel File
Variable Morphology

Grown in water Grown
on land
Biological Species Concept

“Species are groups of interbreeding
natural populations that are
reproductively isolated from other
such groups.”
Ernst Mayr
Reproductive Isolation
 Cornerstone of the biological species
 Speciation is the attainment of
reproductive isolation
 Reproductive isolation arises as a
by-product of genetic change
Genetic Divergence
 Gradual accumulation of differences
in the gene pools of populations
 Natural selection, genetic drift, and
mutation can contribute to divergence
 Gene flow counters divergence
Genetic Divergence

parent species

time A time B time C time D
Reproductive Isolating

 Prezygotic isolation
 Mating or zygote formation is prevented

 Postzygotic isolation
 Takes effect after hybrid zygotes form

 Zygotes may die early, be weak, or be sterile
Prezygotic Isolation

Ecological Isolation
Temporal Isolation
Behavioral Isolation
Mechanical Isolation
Gametic Mortality
Ecological Isolation and Competitive Exclusion in
Two Crayfish (Orconectes Virilis and Orconectes

 Ecology: Vol. 51, No. 2, pp. 225–236.

 Richard V. Bovbjerg
 Abstract. The two crayfish, Orconectes virlis and O.
immunis, have similar ranges but are ecologically
isolated within these ranges; the former species
inhabits streams and lake margins, the latter inhabits
ponds and sloughs. Field and experimental data
suggest that the stream species, O. virilis, is
excluded from ponds by summer drying and periodic
low oxygen periods of those ponds.
O. Virilis is not tolerant of low
oxygen levels and is more
 “It (O. virilis) is not as competent a burrower as O.
immunis nor is its as tolerant of low oxygen levels.
Neither current nor substratum excludes the pond
species, O. immunis, from the stream. But both field
and laboratory observations strongly indicate a
competitive exclusion of O. immunis by O. virilis. The
latter was demonstrated to be intrinsically more
aggressive in interspecific contacts, and in other
experiments, evicted O. immunis from crevices in the
substratum. “
Sister Species

Streams and lake margins

Sloughs and ponds

Orconectes virilis and Orconectes
Flowers and Temporal Isolation
 “Populations may mate or flower at
different seasons or different times of
day. Three tropical orchid species of
the genus Dendrobium each flower
for a single day; the flowers open at
dawn and wither by nightfall.”
Flowers Respond to the Same
Meteorological Stimuli…
 Flowering occurs in response to
certain meteorological stimuli, such
as a sudden storm on a hot day. The
same stimulus acts on all three
species, but the lapse between the
stimulus and flowering is 8 days in
one species, 9 in another, and 10 or
11 in the third.
Dendrobiums are Epiphytes
Behavioral Isolation
 “Behavioral isolation is another isolating
mechanism. This mechanism operates
through courtship behavioral patterns. If
one species displays a certain courtship
pattern, it won’t be recognized by those of
the other species. This is the mechanism
which separates wolfs from dogs, their
courtship patterns are different in the wild.”
Sonographs of Two Closely Related
Tree Frogs
Mechanical Isolation
 “Mechanical isolation deals with the
actual mechanics of the reproductive
organs. Mechanical isolation acts a
prezygotic barrier by preventing
sexual intercourse between two
different species. “
Mechanical Isolation Between Two
Species of Sage
Gametic Mortality
 Sperm are immobilized or destroyed
before fertilization can take place.
Postzygotic Mechanisms

 Zygotic mortality

 Hybrid inviability

 Hybrid sterility
Zygote Mortality
Hybrid Inviability
Hybrid Sterility
 Hybrid sterility: Horse + Donkey ->

Mechanisms of Speciation

 Allopatric speciation

 Sympatric speciation

 Parapatric speciation
Allopatric Speciation

 Speciation in geographically
isolated populations
 Some sort of barrier arises and
prevents gene flow
 Effectiveness of barrier varies with
Allopatric Speciation
in Wrasses

 Isthmus of Panama arose and
separated wrasses in Atlantic and
 Since separation, genes for certain
enzymes have diverged in structure
 Divergence may be evidence of
speciation in progress
Blue-Headed Wrasse
Sister Species – Florida and
Western Scrub Jays
 These

Aphelocoma coerulescens Aphelocoma californica

Florida Species are separated by 1600 km

Common Ancestor
Extensive Divergence Prevents

 Species separated by geographic
barriers will diverge genetically

 If divergence is great enough it will
prevent inbreeding even if the
barrier later disappears
 Island chains some distance from
 Galapagos Islands
 Hawaiian Islands
 Colonization of islands followed by
genetic divergence sets the stage for
A few individuals of a
species on the mainland 3
reach isolated island 1. 2
Speciation follows genetic
divergence in a new habitat. 4

Later in time, a few
Speciation on an individuals of the new
species colonize nearby
Archipelago island 2. In this new
habitat, speciation follows
genetic divergence.

Speciation may also
follow colonization of 1
islands 3 and 4. And it
may follow invasion of 3
island 1 by genetically 2
different descendents
of the ancestral species. 4
Hawaiian Islands
 Volcanic origins, variety of habitats
 Adaptive radiations:
 Honeycreepers - In absence of other bird
species, they radiated to fill numerous
 Fruit flies (Drosophila) - 40% of fruit fly
species are found in Hawaii
Hawaiian Honeycreepers

Speciation without a Barrier

 Sympatric speciation
 Species forms within the home range
of the parent species
 Parapatric speciation
 Neighboring populations become
distinct species while maintaining
contact along a common border
Sympatric Speciation in African

 Studied fish species in two lakes
 Species in each lake are most likely
descended from single ancestor
 No barriers within either lake
 Some ecological separation but
species in each lake breed in
African Cichlids
Speciation by Polyploidy

 Change in chromosome number
(3n, 4n, etc.)
 Offspring with altered chromosome
number cannot breed with parent
 Common mechanism of speciation in
flowering plants
Possible Evolution of Wheat
T. aestivum (one of the
common bread wheats)
Triticum monococcum T. tauschii
(einkorn) (a wild relative)
Unknown species T. turgidum
of wild wheat (wild emmer)

Figure 18.9
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cross-fertilization, followed by a
spontaneous chromosome doubling
Parapatric Speciation

evolve into
distinct species BULLOCK’S
contact along a
common border
Recall: Most products of
crossbreeding, such as the mule,
are sterile…
 “New Mexico Whiptail, is an all-
female species that is actually a
mixture (hybrid) of … the Western
Whiptail, which lives in the desert,
and the Little Striped Whiptail, a
denizen of grasslands.”

 This is another example of parapatric

We’re All Related

 All species are related by descent

 Share genetic connections that
extend back in time to the
prototypical cell
Patterns of Change
in a Lineage

 Cladogenesis
 Branching pattern
 Lineage splits, isolated populations
 Stopped here 1/25
 Anagenesis
 No branching
 Changes occur within single lineage
 Gene flow throughout process
Evolutionary Trees

new species extinction
branch point present)
(a time of
divergence, branch point a new
species dashed line
speciation) (a time of
(only sketchy
evidence of
a single evolutionary
a single lineage relationship)
Gradual Model
 Speciation model in which species
emerge through many small
morphological changes that
accumulate over a long time period

 Fits well with evidence from certain
lineages in fossil record
Punctuation Model
 Speciation model in which most
changes in morphology are
compressed into brief period near
onset of divergence

 Supported by fossil evidence in
some lineages
Adaptive Radiation
 Burst of divergence
 Single lineage gives rise to
many new species
 New species fill vacant
adaptive zone
 Adaptive zone is “way of life”
Adaptive Radiation
 Irrevocable loss of a species
 Mass extinctions have played a
major role in evolutionary
 Fossil record shows 20 or more
large-scale extinctions
 Reduced diversity is followed by
adaptive radiation
Asteroid Imparter

Mass Extinction
Who Survives?
 Species survival is to some extent
 Asteroids have repeatedly struck
Earth, destroying many lineages
 Changes in global temperature favor
lineages that are widely distributed