Species

Diversity is a
consequence of
cladogenesis, the
branching or
multiplication of
lineages, each of
which the evolved by
anagenesis along it
own path.

Each branching in the

phylogenetic tree

Cladogenesis vs.
Anagenesis

What are Species?

Species are particular kinds or
types of organisms.

The word species in Latin means

kind.

There are many definitions of

species.

Table 17.1

And: Morphological Species Concept

Phylogenetic Species
Concept - PSC
"An irreducible (basal) cluster of organisms
diagnosably different from other such clusters,
and within which there is a parental pattern
of ancestry and descent.
The smallest possible groups whose members
are descended from a common ancestor and
who all possess defining or derived
characters that distinguish them from other
such groups. Zimmer and Emlen

Phylogenetic species concept emphasizes the

phylogenetic history of organisms: common

Phylogenetic Species
Concept - PSC
A species is the smallest lineage that
can be united by synapomorphic
characters.

Synapomorphies are characters shared

by two or more taxa that are derived
from a common ancestor.

The members of the species should

share characteristics that other groups
lack; these characteristics are
diagnostic.

This definition makes no reference to

Biological Species
Concept - BSC

As defined by Ernest Mayr

(1942): Species are
groups of actually or
potentially interbreeding
populations, which are
reproductively isolated
from other such groups.

Populations of similar
organisms that interbreed
in the wild and produce
viable and fertile

Biological Species
Concept - BSC
Based on Buffon's ideas.
Reproductive isolation (no genetic
exchange) is a key element of this
definition.
Based on sexual reproduction.
Reproduction without human
interference.

Morphological similarities and

Biological Species Concept BSC

Biological Species
Concept - BSC

Variation within populations:

characteristics vary among the
members of a single population of
interbreeding individuals.

Biological Species
Concept - BSC

Geographic variation: populations

of a species differ; there exists a
spectrum from slight to great
difference, e.g. human populations.

Geographic Variation
Admiral Butterfly
Northeastern US and Canada
Southeastern US
Pacific Coast

http://www.anoleannals.org/

Biological Species
Concept - BSC

Sibling species: these are reproductively

isolated populations that are difficult or
impossible to distinguish by morphological
features, but which are often recognized
by differences in ecology, behavior,
chromosomes and other characters.

Domain and Application of

BSC
All concepts have limitations

Difficulties defining species with BSC:

Testing allopatric populations.

Evaluating differences in fertility of
offspring.
Fossils cannot be tested.

Domain and Application of

BSC
Difficulties defining species with BSC:

Asexual reproducing organisms are

genetically isolated because they
cannot reproduce sexually.
Self-pollinated plants.
Plasmid-mediated horizontal gene
exchange between different species.

Domain and Application of

BSC
DOMAIN

The domain of the BSC is restricted

to sexual, outcrossing organisms,
and to short intervals of time.

E. g. Fossils cannot be tested;

asexual reproducing organisms are
genetically isolated because they
cannot reproduce sexually; selfpollinated plants, etc.

Domain and Application of

BSC
A second meaning of species is a
taxonomic category, just like genus
or family.

Borderline Cases
Interbreeding versus reproductive
isolation is not an either/or, all or
none situation.

Narrow hybrid zones exist where

genetically distinct populations meet
and interbreed to a limited extent,
but in which there exist partial
barriers to gene exchange.

Borderline Cases
The hybridizing entities are often
recognized as species but may be
called semispecies.

A collection of semispecies is a
superspecies.

Borderline Cases
The biological species concept is
sometimes difficult to apply in
botany.
There are many types of vegetative
propagation and self-fertilization in
plants.

Borderline Cases

Geographic variation in status

occurs when genetically different
populations appear to be
conspecific in certain geographic
regions, but to be different
species elsewhere.

Divergence processes in the ring species Ensatina eschscholtzii. a)

Ecomorphotypes in Ensatina defined by color pattern. b) Nei's (1972) genetic
distances based in 26 allozyme loci, showing increased differentiation
towards the terminus of the ring (red) and lower genetic divergence between
populations around the ring (black). c) Floristic provinces of California;
different colors refer to distinct plant associations.

Borderline Cases - Ring

Species
A connected series of neighboring populations, each of
which can interbreed with close related populations, but for
which there exist at least two "end" populations in the
series, too distantly related to interbreed. There is a
potential gene flow between each "linked" population. The
non-breeding, though genetically connected, "end"
populations may coexist in the same region, closing the "ring".
Ring species provide important evidence of evolution in that
they illustrate what happens over time as populations
genetically diverge, and are special because they represent
in living populations what normally happens over time
between long deceased ancestor populations and living
populations, in which the intermediates have become
extinct
Ring species "are only showing us in the spatial dimension
something that must always happen in the time dimension.
Richard Dawkins

Practical Difficulties
E. g. Testing allopatric populations;
evaluating differences in fertility of
offspring.

The greatest practical limitation of the BSC

lies in determining whether or not
geographically segregated (allopatric)
populations belong to the same species.
Geographic isolation is an extrinsic
reproductive barrier.

Practical Difficulties
Populations with intrinsic barriers to gene
exchange can undergo independent
evolutionary change, even if they should
become sympatric.

Range extension or colonization could well

bring presently isolated populations in
contact, so the evolutionary future of the
populations depends on whether or not they
have evolved reproductive isolation.

Allopatric populations have been

classified as species if their differences
in phenotype or in DNA sequence are
as great as those usually displayed by

When Species Concepts

Conflict
Allopatric populations that can be
distinguished by fixed characters are
species according to the PSC, but if the
diagnostic differences are slight, advocates
of the BSC may recognize the populations
as geographic variants of a single species.

In some cases, a local population of

widespread species evolves reproductive
isolation from other populations, which
remain reproductively compatible with one
another. Under the BSC, two species would
be recognized; under the PSC, the various
distinguishable populations of the
paraphyletic group might be named as
distinct species.

Figure 17.3 The phylogeny of some species and populations in the moth genus Greya

Barriers to Gene Flow

Gene flow between biological species is
largely or entirely prevented by
biological differences that have often
been called isolating mechanisms.

Other terms are isolating barriers or

barriers to gene flow.

Under BSC, speciation consists of the

evolution of biological barriers to
gene flow.

Barriers to Gene Flow

Barriers to gene flow between
two populations or species can
be classified in various ways:
Premating vs. postmating
Prezygotic vs. postzygotic
Intrinsic vs. extrinsic

Premating Prezygotic
Barriers
A. Ecological isolation: potential mates do
not meet.

Geographic isolation: Do not live in same

place, physically separated in space.
Temporal isolation: reproductive period
occurs at different time of the year (or of the
day).
Habitat isolation: live in the same locality but
in different habitats, e.g. primarily aquatic while
the other mostly terrestrial.
Immigrant inviability: Immigrants between
populations do not survive long enough to

Geographic Isolation
by Formation of a Geographic Barrier
(Vicariance)

Premating Prezygotic
Barriers
B. Potential mates meet but do not
mate.

Behavioral isolation: differences in

courtship or life style.
Pollinator isolation: different
pollinators respond to different
colors, scents or forms of flowers.

Figure 17.4 Oscillograms of the songs of three morphologically indistinguishable species of green
lacewings

Figure 17.6 Pollinator isolation in monkeyflowers

Postmating Prezygotic
Barriers
Mating between species occurs but the
fertilization of ova does not occur.

Mechanical isolation: copulation

occurs, but no transfer of male
gametes takes place because of failure
of mechanical fit of reproductive
structures.
Copulatory isolation: failure of
fertilization because of behavior during
copulation or because genitalia fail to
stimulate properly.

Postmating Prezygotic
Barriers

Mating between species occurs but the fertilization of ova

does not occur.
Mechanical isolation:
Copulatory isolation:

Gametic isolation:
gamete recognition is
based on the presence
of specific molecules
on the coats around
the egg, which adhere
only to
complementary
molecules in the
sperm.

Postzygotic Barriers
Postzygotic barriers prevent the
hybrid zygote from developing into a
viable, fertile adult.

Extrinsic: hybrid fitness depends on

context.

Ecological inviability: hybrids do not

have ecological niche in which they are
competitively equal to parent species.
Behavioral sterility: hybrids are less
successful than parent species in
obtaining mates.

Postzygotic Barriers
Intrinsic: hybrid fitness is low because
of problems that are relatively
independent of environmental context.

Reduced hybrid viability: hybrid

zygote dies in the early stages of
development or fails to
reach sexual maturity.
Reduced hybrid fertility: hybrid does
not produce functional gametes.
Hybrid breakdown: offspring of hybrids
fail to produce functional gametes or do

Possible Causes of Hybrid

Infertility
Reduced fertility of hybrids can be caused by:
Structural differences between the
chromosomes that cause segregation of some
aneuploid gametes during meiosis: unbalanced
number of chromosomes.
Different number of chromosomes in each
of the parent species. Hybrid gametes cant
undergo normal meiosis.

Differences between genes from the two

parents interact disharmoniously.

Possible Causes of Hybrid

Infertility
Haldanes Rule (Haldane, 1922):

Hybrid sterility and inviability is often

reduced to the heterogametic sex: male
in mammals and most insects; female in
birds and butterflies.

When in the offspring of two different

animal races one sex is absent, rare, or
sterile, that sex is the heterozygous
(heterogametic) sex.

Hybrid Breakdown
Hybrid breakdown occurs in the F2
generation and backcross offspring,
between species and among different
geographic populations of the same species

The common interpretation of this

phenomenon is that the F1 generation
produced various combinations of alleles
that were disharmonious.

Alleles at different loci within the same

population have presumably been selected
to form harmonious combinations. They are

F1 hybrid is not sterile but offspring are feeble

and/or sterile.