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THE

NATURE OF
SPECIES AND
THE BIOLOGICAL
SPECIES CONCEPT

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

Formulated by ERNST MAYR,


an evolutionary biologist (1942)

Gene flow between populations holds the


phenotype of a population together

(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

Prezygotic

barriers block fertilization from


occurring by:
Impeding

different species from attempting to

mate
Preventing
Hindering

the successful completion of mating

fertilization if mating is successful

Prezygotic barriers
Habitat Isolation

Temporal Isolation

Individuals
of
different
species

(a)

Postzygotic barriers

Behavioral Isolation

Mechanical Isolation

Gametic Isolation

Mating
attempt

(c)

(d)

(e)

(f)

Reduced Hybrid Viability

Reduced Hybrid Fertility

Hybrid Breakdown

Viable,
fertile
offspring

Fertilization

(g)

(h)

(i)

(j)

(b)

(k)

(l)

Habitat

isolation: Two species encounter each


other rarely, or not at all, because they
occupy different habitats, even though not
isolated by physical barriers

(a)

Water-dwelling Thamnophis

(b)

Terrestrial Thamnophis

Temporal

isolation: Species that breed at


different times of the day, different seasons,
or different years cannot mix their gametes

(c)

Eastern spotted skunk


(Spilogale putorius)

(d)

Western spotted skunk


(Spilogale gracilis)

(e)

Behavioral
isolation:
Courtship rituals
and other
behaviors unique
to a species are
effective barriers
Courtship ritual of bluefooted boobies

(f)

Mechanical
isolation:
Morphological
differences can
prevent
successful
mating
Bradybaena with shells
spiraling in opposite
directions

(g)

Gametic isolation:
Sperm of one species
may not be able to
fertilize eggs of
another species
Sea urchins

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 hybrids development

(i)

Donkey

( j)

Horse

(k)

Mule (sterile hybrid)

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

(l)

Hybrid cultivated rice plants with


stunted offspring (center)

Limitations of the Biological


Species Concept

The biological species concept cannot be applied to fossils or asexual organisms (including all
prokaryotes)

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

Speciation can take place with or without


geographic separation

Speciation can occur in two ways:

Allopatric speciation

Sympatric speciation

SPECIATION- The process of Biological Species Formation.

(a) Allopatric speciation

(b) Sympatric speciation

Allopatric
Country)
In
allopatric (Other
speciation,
gene flow

is
interrupted
Speciationor reduced when a population is
divided into geographically isolated
subpopulations

The Process of Allopatric


Speciation
The definition of barrier depends on the ability of a
population to disperse
Separate populations may evolve independently
through mutation, natural selection, and genetic drift

A. harrisi

A. leucurus

Evidence of Allopatric
Speciation
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

Mantellinae
(Madagascar only):
100 species
Rhacophorinae
(India/Southeast
Asia): 310 species
Other Indian/
Southeast Asian
frogs

100

60

80
1

40

20

Millions of years ago (mya)


1

India

Madagascar
88 mya

65 mya

56 mya

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
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

2n = 6

4n = 12
Failure of cell
division after
chromosome
duplication gives
rise to tetraploid
tissue.

2n
Gametes
produced
are diploid..

4n
Offspring with
tetraploid
karyotypes may
be viable and
fertile.

Species B
2n = 4

Unreduced
gamete
with 4
chromosomes
Meiotic
error

Species A
2n = 6

Normal
gamete
n=3

Hybrid
with 7
chromosomes

Unreduced
gamete
with 7
chromosomes

Normal
gamete
n=3

Viable fertile
hybrid
(allopolyploid)
2n = 10

Polyploidy

is much more common in plants


than in animals

Many

important crops (oats, cotton, potatoes,


tobacco, and wheat) are polyploids

Sexual Selection
Sexual

selection can drive sympatric


speciation

Sexual

selection for mates of different colors


has likely contributed to the speciation in
cichlid fish in Lake Victoria

EXPERIMENT
Normal light

P.
pundamilia

P. nyererei

Monochromatic
orange light

Allopatric
Sympatric
In
allopatric and
speciation,
geographic

isolation
restricts
gene flow
between populations
Speciation:
A Review

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

Hybrid zones provide opportunities to study


factors that cause reproductive isolation
A

hybrid zone is a region in which members of


different species mate and produce hybrids

Patterns Within Hybrid Zones


A

hybrid zone can occur in a single band


where adjacent species meet

Hybrids

often have reduced fitness compared


with parent species

The

distribution of hybrid zones can be more


complex if parent species are found in
multiple habitats within the same region

EUROPE

Fire-bellied
toad range
Hybrid zone

0.99
Allele frequency (log scale)

Yellow-bellied toad,
Bombina variegata

Yellow-bellied
toad range

Fire-bellied toad,
Bombina bombina

0.9
0.5
0.1

0.01
40

20
30
10
10
20
0
Distance from hybrid zone center (km)

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

Pundamilia nyererei

Pundamilia pundamilia

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

In

cases where hybrids have increased fitness,


local extinctions of parent species within the
hybrid zone can prevent the breakdown of
reproductive barriers

GAME!
(by

group)
2 points for every
correct
Answer

Cynotilapia afra, introduced at West Thumbi Island in Lake


Malawi in the 1960s, has split into two genetically distinct
populations, located at the north and south ends of the island.
How can scientists determine whether these populations are
now different species, according to the biological species
concept?
a)

See whether the two populations are morphologically


different from each other: coloring, bone structure, and
so on.

b)

Determine whether captured individuals from the two


different populations will mate and produce offspring in
a laboratory fish tank.

c)

Determine whether individuals from one population


will interbreed with individuals from the other
population when introduced into each others native
habitats.

Cynotilapia afra, introduced at West Thumbi Island in Lake


Malawi in the 1960s, has split into two genetically distinct
populations, located at the north and south ends of the island.
How can scientists determine whether these populations are
now different species, according to the biological species
concept?
a)

See whether the two populations are morphologically


different from each other: coloring, bone structure, and
so on.

b)

Determine whether captured individuals from the two


different populations will mate and produce offspring in
a laboratory fish tank.

c)

Determine whether individuals from one population


will interbreed with individuals from the other
population when introduced into each others native
habitats.

Cichlids in Lake Victoria are thought to have


diversified from a relatively few species to
600 species in about 12,000 years (the last
time the lake dried up). What is the best way
Examine the fossil record in lake sediments.
toa)test
this hypothesis?
b)

Look for morphological characteristics shared among Lake Victoria species


that are distinct from the species present in other lakes.

c)

Look for genetic characteristics shared among


Lake Victoria species that are distinct from
species present in other lakes.

d)

This is not a testable hypothesis.

Cichlids in Lake Victoria are thought to have


diversified from a relatively few species to
600 species in about 12,000 years (the last
time the lake dried up). What is the best way
Examine the fossil record in lake sediments.
toa)test
this hypothesis?
b)

Look for morphological characteristics shared among Lake Victoria species


that are distinct from the species present in other lakes.

c)

Look for genetic characteristics shared among Lake Victoria species


that are distinct from species present in other lakes.

d)

This is not a testable hypothesis.

In what type of environment is a scientist


most likely to find examples of ongoing
a)
areas that have a large range of altitudes, such as mountains and deep
speciation?
valleys
b)

areas such as large plains that facilitate migration of animals

c)

extreme physical environments that are marginal for supporting life

d)

areas that are already species-rich

e)

areas that have undergone recent environmental disturbance or change

In what type of environment is a scientist


most likely to find examples of ongoing
a)
areas that have a large range of altitudes, such as mountains and deep
speciation?
valleys
b)

areas such as large plains that facilitate migration of animals

c)

extreme physical environments that are marginal for supporting life

d)

areas that are already species-rich

e)

areas that have undergone recent environmental disturbance or change

Sea urchins are broadcast spawners: They


release their gametes into the water without
courtship. Which of these reproductive
isolation mechanisms is most likely affecting
a)
behavioral isolation or mechanical isolation
sea
urchins?
b)

gametic isolation or temporal isolation

c)

habitat isolation

Sea urchins are broadcast spawners: They


release their gametes into the water without
courtship. Which of these reproductive
isolation mechanisms is most likely affecting
a)
behavioral isolation or mechanical isolation
sea
urchins?
b)

gametic isolation or temporal isolation

c)

habitat isolation

You spot a sea snake while you are in a boat


on the ocean and remark about how similar it
looks to a snake that lives in central Illinois.
However, you know that they are most likely
separate species due to which of the
a)
behavioral
isolation
following
reproductive
barriers?
b)

gametic isolation

c)

ecological/habitat isolation

d)

temporal isolation

You spot a sea snake while you are in a boat


on the ocean and remark about how similar it
looks to a snake that lives in central Illinois.
However, you know that they are most likely
separate species due to which of the
a)
behavioral
isolation
following
reproductive
barriers?
b)

gametic isolation

c)

ecological/habitat isolation

d)

temporal isolation

The century cactus flowers every 100 years.


A similar cactus flowers every 50 years. If the
50-year cactus pollen has the correct
characteristics to fertilize the century
cactus, which reproductive barrier will most
a)
behavioral
isolation against that possibility?
likely
be working
b)

gametic isolation

c)

ecological/habitat isolation

d)

temporal isolation

The century cactus flowers every 100 years.


A similar cactus flowers every 50 years. If the
50-year cactus pollen has the correct
characteristics to fertilize the century
cactus, which reproductive barrier will most
a)
behavioral
isolation against that possibility?
likely
be working
b)

gametic isolation

c)

ecological/habitat isolation

d)

temporal isolation

Several species of amphibians are able to


read each others mating rituals and
understand them, tend to be fertile at the
same time, and are usually similar in size.
However, when they attempt to mate, they
never produce offspring. Which of the
a)
behavioral isolation
following
barriers
is
most
likely
the
cause
of
b)
gametic isolation
this?
c)
reduced hybrid fertility
d)

hybrid breakdown

Several species of amphibians are able to


read each others mating rituals and
understand them, tend to be fertile at the
same time, and are usually similar in size.
However, when they attempt to mate, they
never produce offspring. Which of the
a)
behavioral isolation
following
barriers
is
most
likely
the
cause
of
b)
gametic isolation
this?
c)
reduced hybrid fertility
d)

hybrid breakdown

When differentiating between two species of


fish, you notice that one species stay near
the surface and feed on floating algae, while
the other species are bottom feeders. This is
a)
morphological species concept.
using
the
b)

ecological species concept.

c)

phylogenetic species concept.

When differentiating between two species of


fish, you notice that one species stay near
the surface and feed on floating algae, while
the other species are bottom feeders. This is
a)
morphological species concept.
using
the
b)

ecological species concept.

c)

phylogenetic species concept.

Which of the following is believed to be the


primary reproductive isolation mechanism for
causing sympatric speciation in North
American apple maggot flies (Rhagoletis
pomonella)?
a)
polyploidy
b)

sexual selection

c)

phylogenetic species concept

d)

habitat differentiation

Which of the following is believed to be the


primary reproductive isolation mechanism for
causing sympatric speciation in North
American apple maggot flies (Rhagoletis
pomonella)?
a)
polyploidy
b)

sexual selection

c)

phylogenetic species concept

d)

habitat differentiation

For FIVE POINTS!

1.

What do you call an offspring of two different animals or plants of different


races, breeds, varieties and species.

2. Who formulated the biological species concept?

3. Defines a species by structural features. It applies to sexual and asexual species but
relies on subjective criteria

4. It views a species in terms of its ecological


niche. It applies to sexual and asexual species and emphasizes the role of
disruptive selection.

5. The differentiation of geographically


Isolated populations into distinct
Species.

THE END!