16.2 How Is Reproductive Isolation Between Species Maintained?

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Chapter 16
The Origin of Species
Lecture Outlines by Gregory Ahearn,
University of North Florida
Copyright © 2011 Pearson Education Inc.
Chapter 16 At a Glance
 16.1 What Is a Species?
 16.2 How Is Reproductive Isolation Between
Species Maintained?
 16.3 How Do New Species Form?
 16.4 What Causes Extinction?
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.1 What Is a Species?
 Biologists need a clear definition of species
– In Pre-Darwinian times, the term “species”
referred to different kinds of organisms
– Species were classified initially based upon
appearance
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.1 What Is a Species?
 Each species evolves independently
– Today, a species is defined as a group of
actually or potentially interbreeding natural
populations that are reproductively isolated
from other such groups (the biological species
concept)
–A group that is reproductively isolated is
unable to breed successfully outside the group
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.1 What Is a Species?
 Each species evolves independently (continued)
– There are two limitations to the biological
species definition
–Because it is based on the ability to
interbreed, this definition cannot be used to
determine species identity among asexually
reproducing organisms or among fossils
–Also, it is often difficult to observe whether
members of two different groups interbreed
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.1 What Is a Species?
 Appearance can be misleading in determining a
species
– Organisms of similar appearance sometimes belong to
different species
– The cordilleran flycatcher and Pacific slope flycatcher are
so similar that birdwatchers can’t tell them apart
– These birds were considered to be a single species,
but research revealed that they do not interbreed and
are in fact two different species
– A butterfly considered for two centuries to be a single
species known as the two-barred flasher butterfly is now
known to be 10 separate species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Members of Different Species May Be Similar in
Appearance
Fig. 16-1
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.1 What Is a Species?
 Appearance can be misleading in determining a
species (continued)
– Organisms that differ in appearance may belong
to the same species
– The myrtle warbler and Audubon’s warbler, which
have different throat colors, were listed in bird
books as two species in the 1970s
–It was then discovered that the warblers
interbreed where their ranges overlap
–Today, they are considered a single species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Members of a Species May Differ in Appearance
Fig. 16-2
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Traits that prevent interbreeding and maintain
reproductive isolation are called isolating
mechanisms
 Mechanisms that prevent mating between
species are called premating isolating
mechanisms
 Mechanisms that prevent formation of vigorous,
fertile hybrids between species are called
postmating isolating mechanisms
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms prevent mating
between species
– Reproductive isolation occurs when members of
one population are unable to interbreed with
members of another population
– For most species, two or more isolating
mechanisms interact to prevent the formation of
fertile offspring
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms include the
following
– Geographical isolation
– Ecological isolation
– Temporal isolation
– Behavioral isolation
– Mechanical incompatibility
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Geographical isolation prevents interbreeding
between populations that do not come into
contact because they live in physically separated
places
– This type of isolation is usually considered to be
a mechanism that allows new species to form
rather than one that maintains reproductive
isolation between species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Geographically separated populations aren’t
necessarily distinct species
–The Kaibab and Abert squirrels living in
geographically separate areas of the Grand
Canyon are physically separated, but still very
similar
–It is currently unknown if the two populations
have diverged into two species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Geographic Isolation
Fig. 16-3
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Ecological isolation occurs when species don’t
mate because they occupy different habitats
–White-crowned sparrows inhabit fields and
meadows, while white-throated sparrows
inhabit dense thickets
–In another example, each species of fig wasp
breeds in the fruit of a different species of fig,
and the wasps thus do not come into contact
with one another
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Ecological Isolation
Fig. 16-4
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Temporal isolation occurs when species can’t
mate because they breed at different times
– In nature, Bishop pines and Monterey pines do
not interbreed
–Bishop pine pollination occurs in summer
–Monterey pine pollination occurs in early
spring
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Temporal Isolation
Fig. 16-5
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Behavioral isolation occurs when species can’t
mate because they have different courtship and
mating rituals
– The songs and plumage of male songbirds are
species-specific
–They attract females of the same species
–Females of other species are unresponsive
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Behavioral Isolation
Fig. 16-6
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Behavioral isolation occurs when species can’t
mate because they have different courtship and
mating rituals (continued)
–Male frogs embrace any female regardless of
species
–Female frogs encountering males of a
different species utter the “release call,” which
causes the males to let go
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Premating isolating mechanisms (continued)
– Mechanical incompatibility occurs when species cannot
mate because their reproductive structures are
incompatible
– In animals with internal fertilization, male and female
sexual organs may not fit together
– For example, snails of species whose shells have lefthanded spirals may be unable to copulate
successfully with snails whose shells have righthanded spirals
– In plants, differences in flower size or structure may
attract different pollinators
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Mechanical Isolation
Fig. 16-7
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Postmating isolating mechanisms limit hybrid
offspring
– Postmating isolating mechanisms include the
following
–Gametic incompatibility
–Hybrid inviability
–Hybrid infertility
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Postmating isolating mechanisms limit hybrid
offspring (continued)
– Gametic incompatibility occurs when sperm from
one species cannot fertilize eggs of another
–In animals, fluids of the female reproductive
tract my weaken or kill sperm of another
species
–In plants, pollen from one species may fail to
germinate when it lands on the stigma of
another species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Postmating isolating mechanisms limit hybrid offspring
(continued)
– Gametic incompatibility may be an especially important
isolating mechanism in species such as marine
invertebrates and wind-pollinated plants that reproduce
by scattering gametes in the water or in the air
– Sea urchin sperm cells contain a protein that allows
them to bind only to eggs of their own species
– Among plants, chemical incompatibility may prevent
the germination of pollen from one species that lands
on the stigma (pollen-catching structure) of the flower
of another species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Postmating isolating mechanisms limit hybrid offspring
(continued)
– Hybrid inviability occurs when hybrid offspring fail to
survive to maturity
– The hybrid may abort early in development or die shortly
after birth
– The hybrid may be unable to reproduce because it
displays behaviors that are mixtures of the two parental
types
– Lovebird hybrids have great difficulty learning to carry
nest materials during flight and probably could not
reproduce in the wild
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.2 How Is Reproductive Isolation Between Species
Maintained?
 Postmating isolating mechanisms limit hybrid
offspring (continued)
– Hybrid infertility occurs when hybrid offspring are
sterile or have reduced fertility
–Mule hybrids (a cross between a horse and a
donkey) are sterile
–Liger hybrids (a zoo-based cross between a
male lion and a female tiger) are sterile
– Infertility is often caused by the failure of
chromosomes to pair properly during meiosis, so
eggs and sperm never develop
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Hybrid Infertility
Fig. 16-8
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Table 16-1
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Speciation is the process by which new
species form
– Speciation depends on two factors
–The isolation of populations must prevent
gene flow from spreading between them and
keeping them similar
–Genetic divergence of the populations must
occur through the evolution of isolating
mechanisms, driven by genetic drift or natural
selection
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Author Animation: Divergent Traits
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Speciation is the process by which new species
form (continued)
– Evolutionary biologists group the different
pathways to speciation into two broad categories
–In allopatric speciation, the isolating
mechanism is a physical barrier
–In sympatric speciation, isolation occurs
without geographical separation
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Geographical separation of a population can lead to
allopatric speciation
– Allopatric speciation occurs when two populations of a
species become separated by a geographical barrier
– The colonization of remote habitats, such as islands or
reefs far out to sea, can lead to separate populations
– Geological changes caused by such factors as
volcanoes, earthquakes, continental drift, and change
of course by rivers can lead to the separation of
populations
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Geographical separation of a population can
lead to allopatric speciation (continued)
– Allopatric speciation occurs when isolated
populations diverge genetically because of
natural selection and genetic drift
– Allopatric speciation is believed to be the most
common type of speciation, especially among
animals
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Author Animation: Allopatric Speciation 1
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Author Animation: Allopatric Speciation 2
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Allopatric Isolation and Divergence
Part of a mainland
population reaches
to an isolated island
The isolated populations begin
to diverge due to genetic drift
and natural selection
Divergence may eventually
become sufficient to cause
reproductive isolation
Biology: Life on Earth, 9e
Fig. 16-9
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Ecological isolation without geographical separation
can lead to sympatric speciation
– Sympatric speciation may occur when a geographical
area contains two distinctly different habitats (ecological
isolation)
– Two populations of a species living in the
geographical area may occupy the two habitats
– Through the action of natural selection, different
members of the species begin to specialize in one
habitat or the other
– The two isolated populations may diverge sufficiently
genetically so that they can’t interbreed and are thus
considered separate species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Ecological isolation without geographical
separation can lead to sympatric speciation
(continued)
– Two sympatric populations of fruit flies
(Rhagoletis pomonella) may be evolving into two
separate species
–One population lays its eggs in hawthorn fruit,
while the other prefers apples
–The two populations experience very little
interbreeding
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Two sympatric populations of fruit flies (Rhagoletis
pomonella) may be evolving into two separate species
(continued)
– Males and females prefer the same type of fruit in which
they developed
– Apples mature two or three weeks later than hawthorn
fruit (flies mature and mate at different times)
– These differences in fruit preference and maturity mean
the two varieties of flies have very little chance of
meeting and are well on their way to becoming separate
species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Author Animation: Sympatric Speciation
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Sympatric Isolation and Divergence
Part of a fly population that lives only on
hawthorne trees moves to an apple tree
The flies living on the apple tree
do not encounter the flies living
on the hawthorne tree, so the
populations diverge
Biology: Life on Earth, 9e
Fig. 16-10
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Mutations can lead to sympatric speciation
– New species may arise almost instantaneously as a
result of mutations that change the number of
chromosomes in their cells
– The acquisition of multiple copies of each chromosome is
known as polyploidy and is a frequent cause of
sympatric speciation
– In general, polyploid individuals cannot mate
successfully with normal diploid individuals and so are
genetically isolated from the parent species
– Polyploid plants are more likely than polyploid animals
to be able to reproduce
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Author Animation: Speciation by Polyploidy
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Under some conditions, many new species may
arise in a short period of time
– The mechanisms of speciation and reproductive
isolation lead to forking branches in the
evolutionary tree of life, as one species splits into
two
– Adaptive radiation is the rise of many new
species over a relatively short period of time
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Interpreting Evolutionary Trees
present
time
Each line
represents
a species
Forks
represent
speciation
events
past
(a) Evolutionary tree
In an adaptive
radiation, multiple
speciation events
may occur rapidly
enough that
biologists cannot
be certain of their
order
(b) Evolutionary tree representing
adaptive radiation
Fig. 16-11
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.3 How Do New Species Form?
 Under some conditions, many new species may
arise in a short period of time (continued)
– This process occurs when populations of one
species invade a variety of new habitats with few
competitors
–Finch colonization of the Galápagos Islands
resulted in 13 species
–Cichlid fish colonization of Lake Malawi
resulted in more than 300 species
–Tarweed plant colonization of the Hawaiian
Islands resulted in 30 species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Adaptive Radiation
(a) Ahinahina
(b) Waialeale dubautia
(c) Kupaoa
(d) Na’ena’e ’ula
Biology: Life on Earth, 9e
Fig. 16-12
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Extinction is the death of all members of a
species
 At least 99.9% of all species that ever existed
are now extinct
 The immediate cause of extinction is probably
always environmental change
–Environmental changes that can lead to
extinction include habitat destruction and
increased competition among species
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Localized distribution make species vulnerable
in changing environments
– Species inhabiting extremely limited ranges may
become extinct if the area is disturbed
–The Devil’s Hole pupfish is found in only one
spring-fed waterhole in the Nevada desert
– Wide-ranging species normally do not succumb
to local environmental catastrophes
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Very Localized Distribution Can Endanger a
Species
Fig. 16-13
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Overspecialization increases the risk of
extinction in changing environments
– Species that develop adaptations that favor
survival in a specific environment are at risk of
becoming extinct
–The Karner blue butterfly feeds only on the
blue lupine plant
–The habitat of the lupine has been significantly
reduced by development
–Loss of the lupine will lead to extinction of the
Karner blue butterfly
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
Extreme Specialization Places Species at Risk
Fig. 16-14
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Interactions with other species may drive a
species to extinction
– Species that are unable to exploit resources
more efficiently and effectively than their
competitors may become extinct
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Interactions with other species may drive a
species to extinction (continued)
– 2.5 million years ago, a land bridge (the isthmus
of Panama) formed between North and South
America
– North American species displaced the vast
majority of South American species, many of
which became extinct
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.
16.4 What Causes Extinction?
 Habitat change and destruction are the leading
causes of extinction
– Extinctions due to prehistoric habitat change
have had a significant effect on the evolution of
organisms
– Human activities are the primary cause of
present-day habitat destruction
–Clearing of tropical rainforests could lead to
loss of up to half of all current species over the
next 50 years
Biology: Life on Earth, 9e
Copyright © 2011 Pearson Education Inc.

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