Barriers to Gene Flow
• Whether or not a physical barrier
deters gene flow depends upon:
Organism’s mode of dispersal or
locomotion
Duration of time organism can move
Genetic Drift in
Snail Populations
Robert Selander studied Helix
aspersa
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
interbreed?
If so, then they will have become two
distinct species
Speciation & Natural Selection
Natural selection can lead to
speciation
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
Selection
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
concept
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
daughter
species
time A time B time C time D
Reproductive Isolating
Mechanisms
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
Immunis)
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
aggressive…
“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
immunis
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.”
http://www.britannica.com/eb/article-49883
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
http://cas.bellarmine.edu/tietjen/images/speciation.htm
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. “
http://ohoh.essortment.com/reproductiveiso_rcgu.htm
Mechanical Isolation Between Two
Species of Sage
http://cas.bellarmine.edu/tietjen/images/mechanical_
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 ->
mule!
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
species
Allopatric Speciation
in Wrasses
Isthmus of Panama arose and
separated wrasses in Atlantic and
Pacific
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
Texas
Aphelocoma coerulescens Aphelocoma californica
Florida Species are separated by 1600 km
Common Ancestor
Extensive Divergence Prevents
Inbreeding
Species separated by geographic
barriers will diverge genetically
If divergence is great enough it will
prevent inbreeding even if the
barrier later disappears
Archipelagos
Island chains some distance from
continents
Galapagos Islands
Hawaiian Islands
Colonization of islands followed by
genetic divergence sets the stage for
speciation
1
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
1
Speciation on an individuals of the new
species colonize nearby
Archipelago island 2. In this new
2
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
niches
Fruit flies (Drosophila) - 40% of fruit fly
species are found in Hawaii
Hawaiian Honeycreepers
FOUNDER SPECIES
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
Cichlids
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
sympatry
African Cichlids
Speciation by Polyploidy
Change in chromosome number
(3n, 4n, etc.)
Offspring with altered chromosome
number cannot breed with parent
population
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
Page 299
14AA X 14BB 14AB 28AABB X 14DD 42AABBDD
cross-fertilization, followed by a
spontaneous chromosome doubling
Parapatric Speciation
Adjacent
populations
evolve into
distinct species BULLOCK’S
ORIOLE
BALTIMORE
ORIOLE
while
maintaining
contact along a
common border
HYBRID ZONE
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
http://www.amnh.org/exhibitions/expeditions/treasure_fossil/Treasures/Unisexual_Whiptail_Lizards/lizards.html?50
speciation
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
diverge
Stopped here 1/25
Anagenesis
No branching
Changes occur within single lineage
Gene flow throughout process
Evolutionary Trees
new species extinction
(branch
ended
before
branch point present)
(a time of
divergence, branch point a new
species dashed line
speciation) (a time of
(only sketchy
divergence,
evidence of
speciation)
presumed
a single evolutionary
a single lineage relationship)
lineage
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
Extinction
Irrevocable loss of a species
Mass extinctions have played a
major role in evolutionary
history
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
random
Asteroids have repeatedly struck
Earth, destroying many lineages
Changes in global temperature favor
lineages that are widely distributed