The

origin of species part I

Campbell chapter 24
How are new species formed?
Ursus maritmus Ursus arctos Ursus americanus
Biological species concept
Eurasian beaver North American beaver

Castor fiber Castor canadensis

Morphological species concept

Pumpkinseed
(Lepomis gibbosus)

Bluegill (Lepomis macrochirus)

Phylogenetic species concept
What is a species?
In the broadest sense, a species is the smallest evolutionary
independent unit.
Two units (i.e. populations) are evolutionary independent from one
another when mutations, drift and selection operate in each
population separately. For these evolutionary mechanisms to be
occurring separately, gene flow between populations must be very
limited or absent.
Species concepts are sets of rules used to established that two entities
are evolutionarily independent i.e. they are different species
Species concepts
Biological species: a group of populations whose members have the
potential to interbreed in nature and produce fertile offspring
Rationale: if two groups cannot reproduce it is because they are evolutionarily
independent
Limitations: Impossible to apply the BSC to fossil groups and to organisms with
asexual reproduction
Morphological species: Species identified based on morphological
differences
Rationale: if two groups are morphologically different it is because they are
evolutionarily independent
Most commonly used concept. Useful for fossils and asexually reproducing
organisms
Limitations: Based on somewhat arbitrary criteria
Species concepts
Phylogenetic species: smallest group of individuals sharing a common ancestor
Rationale: traits can only distinguish populations on a phylogeny if the population
has been isolated in terms of genes flow and has diverged genetically
Relatedness is usually established from similarity in DNA sequences
Useful for fossils (if DNA can be obtained) and asexually reproducing organisms
Limitations: Challenging to establish how much differences is necessary to consider
two groups different species. Different type of DNA sequences can give different
conclusions

8
Reproductive isolation
Reproductive isolation
Reproductive isolation: the existence of barriers preventing the
members of two populations to produce viable, fertile offsprings. In
short bagenetic rriers prevent gene flow (remember, gene flow
increases the similarity among populations)
Reproductive isolation is necessary to achieve evolutionary
independence and thus for speciation to occur.
Reproductive barriers
Prezygotic barriers: prevent mating or fertilization between two species.
Ecological isolation occurs when ecological factors (e.g. habitat or reproductive
season) prevents hybridization
Behavioural isoloation occurs when the signals used for mate recognition and mate
choice are not shared between two populations
Mechanical isolation exist when two populations are morphologically incapable of
mating.
Gametic isolation occurs when the gamete of males and females from two
populations are incompatible.
Habitat barriers
Postzygotic barriers prevent the formation of viable hybrids after fertilization occurred.
Post zygotic barriers can be manifested in three ways:
Reduced hybrid viability
Reduced hybrid fertility
Hybrid breakdown
How reproductive barriers evolve?:
Allopatric speciation
Allopatric speciation:
evidence
Allopatric speciation: evidence
Seeing speciation happening is impossible (except for small organisms with
fast generation time) but we can see the signature of speciation in the DNA of
organisms

The Isthmus of Panama completely separate the Caribbean sea from the
Pacific Ocean

Many pairs of morphological species have been recognized with one

Caribbean species and one Pacific species.

Genetic analyses revealed that the species in one pair are more closely related
to each other than each of them is to any of the other species found on the
same side of the isthmus.

For instance Pacific species B is more closely related to Caribbean species B

(from which it is geographically isolated) than to any other of the Pacific
species.

This pattern is predicted if geographical isolation caused by the emergence of

the isthmus of Panama is responsible for the speciation.

Moreover, geological evidence indicate that the Isthmus of Panama formed

between 10 and 3 million years ag and molecular estimate of divergence
between species in each pair range from 9 to 3 million years ago.
Allopatric speciation: reproductive isolation
increases with distance
A little stats detour: Linear regression
The trend of a relationship between two continuous variable is given by the slope of
equation relating X to Y. A slope of 0 (i.e. a flat horizontal line) indicates no
relationship.
The P-value provided by a linear regression tells us the probability that the slope is
different from 0 or, in other words, the probability of being wrong if we reject the
the null hypothesis that there is no relationship between X and Y
A linear regression calculates a R2 value which varies between 0 and 1. This value
tells us how much of the variation in Y is explained by X.
For example a R2 of 0.55 means that 55% of the variation observed in Y can be
explained by X.
Always keep in mind that if X is causing Y, we expect a significant correlation
between X and Y BUT the existence of a correlation between X and Y does not show
that X is causing Y.
However, if we dont find a correlation between X and Y we can be confident that X
does not cause Y.
Allopatric speciation: isolation as a by-product of
selection: field data
Allopatric speciation: isolation as a by-product
of selection: experimental data
Allopatric speciation: isolation as a by-product
of selection: experimental data
Sympatric speciation by habitat isolation: the
apple maggot
In Figure 6
taxonomic
race is u
identify th
flies that r
on apples
maggot fli
reproduce
hawthorns