the origin of species chapter 24 the origin of species · concept 24.1: the biological species...

The Origin of Species

Chapter 24

Chapter 24

The Origin of Species

Overview: That “Mystery of Mysteries”

• In the Galápagos Islands Darwin discovered

plants and animals found nowhere else on Earth

© 2011 Pearson Education, Inc.

• Speciation, the origin of new species, is at the focal point of evolutionary theory

• Evolutionary theory must explain how new species originate and how populations evolve

• Microevolution consists of changes in allele frequency in a population over time

• Macroevolution refers to broad patterns of evolutionary change above the species level


Concept 24.1: The biological species

concept emphasizes reproductive isolation

• Species is a Latin word meaning “kind” or

“appearance”

• Biologists compare morphology, physiology,

biochemistry, and DNA sequences when

grouping organisms


The Biological Species Concept

• The biological species concept states that a

species is a group of populations whose members

have the potential to interbreed in nature and

produce viable, fertile offspring; they do not breed

successfully with other populations

• Gene flow between populations holds the

phenotype of a population together


Figure 24.2

(a) Similarity between different species

(b) Diversity within a species

Reproductive Isolation

• Reproductive isolation is the existence of

biological factors (barriers) that impede two

species from producing viable, fertile offspring

• Hybrids are the offspring of crosses between

different species

• Reproductive isolation can be classified by

whether factors act before or after fertilization


Figure 24.3_a

Prezygotic barriers

Habitat

IsolationTemporal

Isolation

Behavioral

Isolation

Mechanical

Isolation

Gametic

Isolation

Reduced Hybrid

Viability

Reduced Hybrid

Fertility

Hybrid

Breakdown

Individuals of

differentspecies

MATINGATTEMPT FERTILIZATION

VIABLE,FERTILE

OFFSPRING

Postzygotic barriers

(a) (c) (e)

(d)

(b)

(g)

(k)

(h) (i)

(j)

(l)(f)

Figure 24.3_b

Prezygotic barriers

Habitat

Isolation

Temporal

Isolation

Behavioral

Isolation

Mechanical

Isolation

Gametic

Isolation

Individuals of

differentspecies

MATINGATTEMPT

FERTILIZATION

(a) (c) (e) (f)

(b)

(g)

(d)

Reduced Hybrid

Viability

Reduced Hybrid

Fertility

Hybrid

Breakdown

FERTILIZATION

VIABLE,FERTILE

OFFSPRING

Postzygotic barriers

(k)

(h) (i)

(j)

(l)

Figure 24.3_c

• Prezygotic barriers block fertilization from

occurring by:

– Impeding different species from attempting to

mate

– Preventing the successful completion of mating

– Hindering fertilization if mating is successful


• Habitat isolation: Two species encounter each

other rarely, or not at all, because they occupy

different habitats, even though not isolated by

physical barriers


• Temporal isolation: Species that breed at

different times of the day, different seasons,

or different years cannot mix their gametes


• Behavioral isolation: Courtship rituals and other

behaviors unique to a species are effective

barriers


Albatross Courtship Ritual

Giraffe Courtship Ritual

Blue Footed Boobies Courtship Ritual

• Mechanical isolation: Morphological differences

can prevent successful mating


• Gametic Isolation: Sperm of one species may not

be able to fertilize eggs of another species


• Postzygotic barriers prevent the hybrid zygote

from developing into a viable, fertile adult:

– Reduced hybrid viability

– Reduced hybrid fertility

– Hybrid breakdown


• Reduced hybrid viability: Genes of the different

parent species may interact and impair the

hybrid’s development


• Reduced hybrid fertility: Even if hybrids are

vigorous, they may be sterile


• Hybrid breakdown: Some first-generation hybrids

are fertile, but when they mate with another

species or with either parent species, offspring of

the next generation are feeble or sterile


Limitations of the Biological Species Concept

• The biological species concept cannot be

applied to fossils or asexual organisms

(including all prokaryotes)

• The biological species concept emphasizes

absence of gene flow

• However, gene flow can occur between

distinct species

– For example, grizzly bears and polar bears

can mate to produce “grolar bears”


Figure 24.4Grizzly bear (U. arctos)

Polar bear (U. maritimus)

Hybrid “grolar bear”

Other Definitions of Species

• Other species concepts emphasize the unity within

a species rather than the separateness of different

species

• The morphological species concept defines a

species by structural features

– It applies to sexual and asexual species but relies

on subjective criteria


• The ecological species concept views a species

in terms of its ecological niche

– It applies to sexual and asexual species and

emphasizes the role of disruptive selection

• The phylogenetic species concept defines a

species as the smallest group of individuals on a

phylogenetic tree

– It applies to sexual and asexual species, but it can

be difficult to determine the degree of difference

required for separate species


Concept 24.2: Speciation can take place

with or without geographic separation

• Speciation can occur in two ways:

– Allopatric speciation

– Sympatric speciation


Figure 24.5

(a) (b)Allopatric speciation.

A population forms a

new species while

geographically isolated

from its parent population.

Sympatric speciation.

A subset of a population

forms a new species

without geographic

separation.

Allopatric (“Other Country”) Speciation

• In allopatric speciation, gene flow is

interrupted or reduced when a population is

divided into geographically isolated

subpopulations

– For example, the flightless cormorant of the

Galápagos likely originated from a flying

species on the mainland


The Process of Allopatric Speciation

• The definition of barrier depends on the ability of a

population to disperse

– For example, a canyon may create a barrier for

small rodents, but not birds, coyotes, or pollen


Figure 24.6

A. harrisii A. leucurus

• Separate populations may evolve independently through mutation, natural selection, and genetic drift

• Reproductive isolation may arise as a result of genetic divergence

– For example, mosquitofish in the Bahamas comprise several isolated populations in different ponds


Figure 24.7

(a) Under high predation (b) Under low predation

Evidence of Allopatric Speciation

• 15 pairs of sibling species of snapping shrimp

(Alpheus) are separated by the Isthmus of

Panama

• These species originated 9 to 13 million years

ago, when the Isthmus of Panama formed and

separated the Atlantic and Pacific waters


Figure 24.8

A. formosus

Atlantic Ocean

A. nuttingi

Isthmus of Panama

Pacific Ocean

A. panamensis A. millsae

• Regions with many geographic barriers typically have more species than do regions with fewer barriers

• Reproductive isolation between populations generally increases as the distance between them increases

– For example, reproductive isolation increases between dusky salamanders that live further apart


Figure 24.9

Deg

ree

of

rep

rod

uc

tiv

e i

so

lati

on

Geographic distance (km)

0 50 100 150 200 250 300

2.0

1.5

1.0

0.5

0

• Barriers to reproduction are intrinsic;

separation itself is not a biological barrier


Figure 24.10 EXPERIMENT

RESULTS

Initial population

of fruit flies

(Drosophila

pseudoobscura)

Some flies raised

on starch medium

Mating experiments

after 40 generations

Some flies raised on

maltose medium

Female

Starch Maltose

Ma

leM

alt

ose

Sta

rch

Number of matings

in experimental group

22 9

8 20

Female

Starch

population 1

Ma

leS

tarc

h

po

pu

lati

on

2

Number of matings

in control group

18 15

12 15

Starch

population 2

Sta

rch

po

pu

lati

on

1

Figure 24.10a

EXPERIMENT

Initial population

of fruit flies

(Drosophila

pseudoobscura)

Some flies raised

on starch medium

Mating experiments

after 40 generations

Some flies raised on

maltose medium

Figure 24.10b

RESULTS

Female

Starch Maltose

Male

Malt

ose

Sta

rch

Number of matings

in experimental group

22 9

8 20

Female

Starch

population 1

Male

Sta

rch

po

pu

lati

on

2Number of matings

in control group

18 15

12 15

Starch

population 2

Sta

rch

po

pu

lati

on

1

Sympatric (“Same Country”) Speciation

• In sympatric speciation, speciation takes place

in geographically overlapping populations


Polyploidy

• Polyploidy is the presence of extra sets of

chromosomes due to accidents during cell

division

• Polyploidy is much more common in plants

than in animals

• An autopolyploid is an individual with more

than two chromosome sets, derived from one

species


• An allopolyploid is a species with multiple

sets of chromosomes derived from different

species


Figure 24.11-1

Species A

2n = 6

Species B

2n = 4

Normal

gamete

n = 3

Meiotic error;

chromosome number not

reduced from 2n to n

Unreduced gamete

with 4 chromosomes

Figure 24.11-2

Species A

2n = 6

Species B

2n = 4

Normal

gamete

n = 3

Meiotic error;



Unreduced gamete

with 4 chromosomes

Hybrid with

7 chromosomes

Figure 24.11-3

Species A

2n = 6

Species B

2n = 4

Normal

gamete

n = 3

Meiotic error;



Unreduced gamete

with 4 chromosomes

Hybrid with

7 chromosomes

Unreduced gamete

with 7 chromosomes

Normal

gamete

n = 3

Figure 24.11-4

Species A

2n = 6

Species B

2n = 4

Normal

gamete

n = 3

Meiotic error;



Unreduced gamete

with 4 chromosomes

Hybrid with

7 chromosomes

Unreduced gamete

with 7 chromosomes

Normal

gamete

n = 3

New species:

viable fertile hybrid

(allopolyploid) 2n = 10

• Many important crops (oats, cotton, potatoes,

tobacco, and wheat) are polyploids


Habitat Differentiation

• Sympatric speciation can also result from the

appearance of new ecological niches

• For example, the North American maggot fly can

live on native hawthorn trees as well as more

recently introduced apple trees


Sexual Selection

• Sexual selection can drive sympatric speciation

• Sexual selection for mates of different colors has

likely contributed to speciation in cichlid fish in

Lake Victoria


Figure 24.12

Normal light

Monochromatic

orange light

P. pundamilia

P. nyererei

EXPERIMENT

Allopatric and Sympatric Speciation:

A Review

• In allopatric speciation, geographic isolation

restricts gene flow between populations

• Reproductive isolation may then arise by natural

selection, genetic drift, or sexual selection in the

isolated populations

• Even if contact is restored between populations,

interbreeding is prevented


• In sympatric speciation, a reproductive barrier

isolates a subset of a population without

geographic separation from the parent species

• Sympatric speciation can result from polyploidy,

natural selection, or sexual selection


Concept 24.3: Hybrid zones reveal factors

that cause reproductive isolation

• A hybrid zone is a region in which members of

different species mate and produce hybrids

• Hybrids are the result of mating between species

with incomplete reproductive barriers


Patterns Within Hybrid Zones

• A hybrid zone can occur in a single band where

adjacent species meet

– For example, two species of toad in the genus

Bombina interbreed in a long and narrow hybrid

zone


Figure 24.13

EUROPE

Yellow-bellied

toad, Bombina

variegata

Fire-bellied

toad range

Hybrid zone

Yellow-bellied

toad range

Fire-bellied toad, Bombina bombina

Fre

qu

en

cy o

f

B. v

ari

eg

ata

-sp

ecif

ic a

lle

le

Yellow-bellied

toad range

Hybrid

zone

Fire-bellied

toad range

Distance from hybrid zone center (km)40

0.99

0.9

0.5

0.1

0.01

30 20 10 0 10 20

Figure 24.13a

EUROPE

Fire-bellied

toad rangeHybrid zone

Yellow-bellied

toad range

Figure 24.13b

Fre

qu

en

cy o

f

B.

va

rie

ga

ta-s

pecif

ic a

llele

Yellow-bellied

toad range

Hybrid

zone

Fire-bellied

toad range

Distance from hybrid zone center (km)

40

0.99

0.9

0.5

0.1

0.01

30 20 10 0 10 20

Figure 24.13c

Fire-bellied toad, Bombina bombina

Yellow-bellied toad, Bombina variegata

• Hybrids often have reduced fitness compared with

parent species

• The distribution of hybrid zones can be more

complex if parent species are found in patches

within the same region


Hybrid Zones over Time

• When closely related species meet in a hybrid

zone, there are three possible outcomes:

– Reinforcement

– Fusion

– Stability


Figure 24.14-1

Gene flow

PopulationBarrier to

gene flow

Figure 24.14-2

Gene flow


gene flow

Isolated

population

diverges

Figure 24.14-3

Gene flow


gene flow

Isolated

population

diverges

Hybrid

zone

Hybrid

individual

Figure 24.14-4

Gene flow


gene flow

Isolated

population

diverges

Hybrid

zone

Hybrid

individual

Possible

outcomes:

Reinforcement

OR

OR

Fusion

Stability

Reinforcement: Strengthening Reproductive

Barriers

• The reinforcement of barriers occurs when hybrids are less fit than the parent species

• Over time, the rate of hybridization decreases

• Where reinforcement occurs, reproductive barriers should be stronger for sympatric than allopatric species

– For example, in populations of flycatchers, males are more similar in allopatric populations than sympatric populations


Figure 24.15

Females choosing between

these males:Females choosing between

these males:

Sympatric pied male

Sympatric collared male

Allopatric pied male

Allopatric collared male

(none)

Female mate choice Female mate choice

Own

speciesOwn

species

Other

species

Other

species

Nu

mb

er

of

fem

ale

s

28

24

20

16

12

8

4

0

Fusion: Weakening Reproductive Barriers

• If hybrids are as fit as parents, there can be substantial gene flow between species

• If gene flow is great enough, the parent species can fuse into a single species

• For example, researchers think that pollution in Lake Victoria has reduced the ability of female cichlids to distinguish males of different species

• This might be causing the fusion of many species


Figure 24.16

Pundamilia nyererei Pundamilia pundamilia

Pundamilia “turbid water,”

hybrid offspring from a location

with turbid water

Figure 24.16a

Pundamilia nyererei

Figure 24.16b

Pundamilia pundamilia

Figure 24.16c

Pundamilia “turbid water,” hybrid offspring from a

location with turbid water

Stability: Continued Formation of Hybrid

Individuals

• Extensive gene flow from outside the hybrid zone

can overwhelm selection for increased

reproductive isolation inside the hybrid zone


Concept 24.4: Speciation can occur rapidly

or slowly and can result from changes in

few or many genes

• Many questions remain concerning how long it

takes for new species to form, or how many genes

need to differ between species


The Time Course of Speciation

• Broad patterns in speciation can be studied

using the fossil record, morphological data, or

molecular data


Patterns in the Fossil Record

• The fossil record includes examples of species

that appear suddenly, persist essentially

unchanged for some time, and then apparently

disappear

• Niles Eldredge and Stephen Jay Gould coined the

term punctuated equilibria to describe periods of

apparent stasis punctuated by sudden change

• The punctuated equilibrium model contrasts with a

model of gradual change in a species’ existence


Figure 24.17

(a) Punctuated

pattern

Time

(b) Gradual

pattern

Speciation Rates

• The punctuated pattern in the fossil record and

evidence from lab studies suggest that speciation

can be rapid

– For example, the sunflower Helianthus anomalus

originated from the hybridization of two other

sunflower species


Figure 24.18

Figure 24.19

H. annuus

gameteH. petiolarus

gamete

F1 experimental hybrid

(4 of the 2n = 34

chromosomes are shown)

EXPERIMENT

RESULTS

Chromosome 1

H. anomalus

Chromosome 2

H. anomalus

Experimental hybrid

Experimental hybrid

Figure 24.19a

H. annuus

gameteH. petiolarus

gamete

F1 experimental hybrid

(4 of the 2n = 34

chromosomes are shown)

EXPERIMENT

Figure 24.19b

RESULTS

Chromosome 1

H. anomalus

Chromosome 2

H. anomalus

Experimental hybrid

Experimental hybrid

• The interval between speciation events can range

from 4,000 years (some cichlids) to 40 million

years (some beetles), with an average of 6.5

million years


Studying the Genetics of Speciation

• A fundamental question of evolutionary biology

persists: How many genes change when a new

species forms?

• Depending on the species in question, speciation

might require the change of only a single allele or

many alleles

– For example, in Japanese Euhadra snails, the

direction of shell spiral affects mating and is

controlled by a single gene


• In monkey flowers (Mimulus), two loci affect flower

color, which influences pollinator preference

• Pollination that is dominated by either

hummingbirds or bees can lead to reproductive

isolation of the flowers

• In other species, speciation can be influenced by

larger numbers of genes and gene interactions


Typical

Mimulus

lewisii

(a)

Typical

Mimulus

cardinalis

(c)

M. lewisii with an

M. cardinalis flower-color

allele

(b)

M. cardinalis with an

M. lewisii flower-color

allele

(d)

Figure 24.20

From Speciation to Macroevolution

• Macroevolution is the cumulative effect of many

speciation and extinction events


the origin of species chapter 24 the origin of species · concept 24.1: the biological species...

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