Coevolution

COEVOLUTION
 Coevolution

Qu es coevolucin?

Coevolucin es aquel proceso por el cual dos o ms organismos ejercen presin de seleccin mutua y
sincrnica (en tiempo geolgico) que resulta en adaptaciones especficas recproca.

Janzen 1980

Requisitos del proceso de coevolucin

1) Especificidad: la evolucin de cada rasgo en una especie es debida a presiones selectivas de otros
rasgos de las otras especies del sistema.

2) Reciprocidad: los rasgos en ambos participantes del sistema evolucionan conjuntamente.

3) Simultaneidad: los rasgos en ambos participantes del sistema evolucionan al mismo tiempo.

2
R1 = h S (2 ) 1 1

2
R2 = h S 2 (1 ) 2
i is the genotypic distribution of species i
 Coevolution

What it is coevolution

Species A evolves an Species B evolves in

adaptation in response to response to the adaptation
species B of species A

What it isnt coevolution

Species B evolves in
Species A has some trait
response to that trait in
unrelated to species B
species A
 Coevolution

El proceso coevolutivo puede generar

1) Coadaptacin: ajuste microevolutivo recprocos de unos organismos a otros (Micro-

coevolucin).

2) Coespeciacin: cladognesis recproca como fruto de la interaccin (Macro-coevolution).

 Coevolution

MicroCoevolution

The outcome of the coevolutionary process largely depends on:

1. The type of ecological interaction. The interactions differ in the form of reciprocal selection

2. The genetic mediation of the interaction

Competitive interactions lead to divergence

Antagonistic interactions lead to cycles or escalation

Mutualistic interactions lead to convergence

D. Futuyma y M. Slatkin 1983. Coevolution. Sinaur

 Coevolution

MicroCoevolution

Coevolution in competitive interactions

Reciprocal Selection:
1

0.8 The fitness of Species 1 individuals is decreased by

0.6 interacting with Species 2
0.4
The fitness of Species 2 individuals is decreased by
0.2
interacting with Species 1
0
Frequency

0 0.5 1 1.5 2 2.5 3

Reciprocal selection favors traits in each species that

Time
reduce the efficacy or frequency of the interaction

0.8 Coevolutionary dynamics:

0.6

0.4 Divergence in traits mediating the interaction

0.2
(i.e., character displacement)
0
0 0.5 1 1.5 2 2.5 3

Phenotype
(e.g., beak size)
 Coevolution

MicroCoevolution

Coevolution in antagonistic interactions

1 Reciprocal Selection:
0.8
The fitness of victim individuals is increased by not
Frequency

0.6 interacting
0.4 The fitness of exploiter individuals is increased by
0.2 interacting
Exploiter Victim
0 Reciprocal selection favors victim traits that decrease
0 0.5 1 1.5 2 2.5 3 the efficacy or frequency of interaction, but exploiter
traits that increase the efficacy or frequency of the
Phenotype
interaction
(e.g., running speed)

Coevolutionary escalation Reciprocal selection favors increased (or decreased) phenotypes in both
victim and exploiter. Selection is directional

Coevolutionary matching Reciprocal selection favors exploiters that match the phenotype of the victim,
but victims that mismatch the phenotype of the exploiter. Selection is time-delayed negative frequency-
dependent
 Coevolution

MicroCoevolution

Coevolution in antagonistic interactions: Coevolutionary escalation

Time

1 1 1
Frequency

0.8 0.8 0.8

0.6 0.6 0.6

0.4 0.4 0.4

0.2 0.2 0.2

0 0 0
0 0.5 1 1.5 2 2.5 3 0 0.5 1 1.5 2 2.5 3 0 0.5 1 1.5 2 2.5 3

Phenotype
Coevolutionary dynamics:

Without cost: Endless escalation of phenotypes

With cost: Phenotypic cycles

Probability of attack 1 1
0.75
Examples: 0.5
0.25
Concentration of plant defensive compounds 0 Large
Large
Concentration of insect detoxification enzymes
Parasite trait zP Host trait zH

Small Small
 Coevolution

MicroCoevolution

Coevolution in antagonistic interactions: Coevolutionary matching

1 1
Coevolutionary dynamics:
-
Trait means (zi)

0.8
Exploiter 0.8
Victim
0.6 0.6 Phenotypes cycle endlessly
0.4 0.4
Exploiter adapts to common victim phenotypes
0.2 0.2

0 0 Should produce an advantage for rare victim phenotypes

1000 2000 3000 4000 5000 1000 2000 3000 4000 5000

Generate coevolving polymorphism

Generation

Probability of attack
Example:
1
Plant flowering time
0.75
Insect emergence time
0.5
0.25 0.25
Lice matching feather barb size 0 0 Large
Large

Parasite trait zP Host trait zH

Small Small
 Coevolution

MicroCoevolution

Coevolution in mutualistic interactions

Reciprocal Selection:

The fitness of Species 1 individuals is increased by

1 interacting with Species 2 individuals
0.8
The fitness of Species 2 individuals is increased by
0.6 interacting with Species 1 individuals
0.4
Reciprocal selection favors traits in both species that
0.2
increase the efficacy or frequency of the interaction.
0
Frequency

0 0.5 1 1.5 2 2.5 3 Selection is positive frequency-dependent

Time
1 Coevolutionary dynamics:
0.8
Coevolving complementarity in symbiotic
0.6 interactions.
0.4
Coevolutionaty convergence in free-living
0.2 interactions.
0
0 0.5 1 1.5 2 2.5 3

Phenotype
(e.g., Timing)
 Coevolucin

Modelos Genticos microevolutivos

Gen-for-Gen Coevolution

Matching allele
 Coevolucin

Coevolucin multispecifica o difusa

1) La interaccin con varios organismos esta genticamente correlacionada.

2) La presencia/ausencia de otro organismo interactuante afecta a la

interaccin con nuestro organismo focal.

3) El impacto de un organismo en el fitness de su pareja se ve afectado por

la presencia/ausencia de un tercer organismo
 Coevolution

Geographic Mosaic Theory of Coevolution (GMTC)

a) Most species are formed by a collection of genetic and ecologically differentiated populations
inserted in a complex landscape.

b) The ecological interactions and community context varies spatially, among populations

The coevolutionary dynamics is driven by three main components

1) Geographic selection mosaics: Natural selection on interspecific interactions

varies among populations.

2) Coevolutionary hotspots: Interactions are subject to reciprocal selection only

within some local communities.

3) Trait remixing: The genetic structure of coevolving species changes through new
mutations, gene flow across landscapes, random genetic drift, and extinction of local
populations.

http://bio.research.ucsc.edu/people/thompson
 Coevolution

Geographic Mosaic Theory of Coevolution (GMTC)

This theory visualizes the landscape as a mosaic of coevolutionary hotspots, populations

where reciprocal selection is strong and coevolution is ongoing, embedded in a broader matrix
of coevolutionary coldspots, where local selection is weak, non-reciprocal or where only one of
the participants occurs

http://bio.research.ucsc.edu/people/thompson
 Coevolution

Geographic Mosaic Theory of Coevolution (GMTC)

 Coevolution

Geographic Mosaic Theory of Coevolution (GMTC)

 Coevolution

Un ejemplo
 Coevolution

Modelos macro-coevolutivos

1 Escape y radiacin: fenmenos de especiacin sincrnicos en ambos

participantes. Especiacin ocurre cuando no hay interaccin

2 Coespeciacin: Existe especiacin simultnea

3 Coevolucin diversificadora: La propia interaccin induce especiacin

como consecuencia de adaptacin local recproca

4 Rastreo filogentico: Un participante especia en funcin de la

diversificacin del otro participante

5 Escalada: El ambiente selectivo se hace ms severo, induciendo cambios

conjuntos en el fenotipo. Pero no hay proceso coevolutivo.

Page, M. 2003. Tangled Trees: Phylogeny, Cospeciation, and Coevolution. University of Chicago Press
 Coevolution

Modelos macro-coevolutivos
 Coevolution

Escape and radiation coevolution (Ehrlich and Raven 1964)

Step 1: A new mutation arises within a host (plant) lineage which confers resistance to
parasitism
 Coevolution

Escape and radiation coevolution (Ehrlich and Raven 1964)

Step 2: The resistant host lineage diversifies rapidly in the absence of parasitism
 Coevolution

Escape and radiation coevolution (Ehrlich and Raven 1964)

Step 3: A new mutation arises in the parasite allowing it to feed on the previously
resistant host lineages. Rapid parasite diversification follows.
 Coevolution

Escape and radiation coevolution (Ehrlich and Raven 1964)

The result is rapid bursts of diversification, but NOT cospeciation

 Coevolution

Escape and radiation coevolution (Ehrlich and Raven 1964)

Esperamos congruencia filogentica

 Coevolution

Coespeciacin

The parallel evolution of two associated taxa (such as a host and a symbiont), such that speciation events in
the two taxa are coupled
 Coevolution

Host-shift and phylogenetic tracking in antagonism and mutualism

Coevolution
Coevolution
 Coevolution

Why poor congruence of fig and fig wasp

phylogenies?

-Multiple wasp species pollinating one fig species

-Host switching
-Evidence for frequent hybridization in figs but in
wasps (inbreed)

Diversification of this mutualism has not occurred

through strict-sense coadaptation and
cospeciation between pairs of fig and wasp
species