The Ecology and Geography of Coevolution

Concepts + case studies

Bob Week - BIOL227 - Kiel University

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Interactive slides available at:

https://bobweek.github.io/teaching/coevolution_intro/lecture.html

Many uses of the word Coevolution

A magazine from the 70s

but …

The most important and coolest meaning is

Coevolution of two (or more) species due to ecological interactions

Interspecific Coevolution is a Massive Topic

Roadmap

Act I: What is coevolution?

  • Definitions
  • Representative forms of coevolution

Act II: The ecology of coevolution

  • Coevolutionary outcomes of ecological interactions
  • Ecological outcomes of coevolutionary dynamics (ie, eco-evo feedback!)
  • The community context

Act III: The geography of coevolution

  • Spatial structure of ecological interactions
  • Determinants of host-parasite local adaptation
  • The spatial signature of coevolution

Learning Outcomes

  • Define coevolution precisely (and say what it is not)
  • Recognize how coevolution can maintain genetic diversity
  • Explain how ecology and coevolution feedback on each other

Act I: What is coevolution?

Conditions for Coevolution

DEFINITION: Coevolution occurs when

  1. the interaction between a pair of species alters selection pressures on both species and

  2. both species exhibit an evolutionary response to the altered selection pressures

Review:

  • What is selection?
  • What is an evolutionary response to selection?

Read:

When is it coevolution?”, Janzen 1980

Selection Review (see Lande & Arnold, 1983)

DEFINITION: Selection occurs when fitness and phenotype covary

DEFINITION: A phenotype is any measurable attribute of an organism

DEFINITION: Fitness is the lifetime reproductive output of an individual

Adaptation the Evolutionary Response to selection

DEFINITION: Adaptation is the inherited/evolutionary resposne to selection

Coevolution is Co-adaptation

Coevolution both species adapt in response to selection caused by interaction

Examples of Coevolution: Character Displacement

  • Interaction type: resource competition along a continuous niche axis
  • Altered selection: divergent resource use/specialization (directional selection)
  • Coevolutionary response: both species evolve to reduce resource competition

Examples of Coevolution: Mutualistic Matching

  • Interaction type: mutualism w trait-matching (eg, plant-poll. phenology)
  • Altered selection: positive freq. dependent selection (stabilizing selection)
  • Coevolutionary response: loss of polymorphism (decr. genetic variation)

Examples of Coevolution: Trench Warfare

  • Interaction type: host-parasite w lock-key (eg, viral receptor binding)
  • Altered selection: negative freq. dependent selection (diversifying selection)
  • Coevolutionary response: maintenance of polymorphism (incr. genetic variation)

Examples of not Coev: Unilateral Matching

  • Interaction type: commensalism w trait-matching
  • Why not coev?: selection is not modified for host species

Examples of not Coev: Unilateral Displacement

  • Interaction type: resource competition along niche axis
  • Why not coev?: niche location not a heritable trait for blue species

Act II: The Ecology of Coevolution

Taxonomy of ecological interactions

A classic coevolutionary woozle

A classic coevolutionary woozle (debunked!)

A pollination-based counter example

  • Pollinator fly:
    • Moegistorhynchus longirostris
  • Flower:
    • Lapeirousia anceps

Question:

  • If mutualism leads to matching, why the long face?

The Phenotypic Interface of Interaction

The Phenotypic Interface of Interaction

(Evidence for) Selection Due to Interaction

  • Fly trait > flower trait:
    • fly gets more nectar
  • Flower trait > fly trait:
    • more flower pollen transferred

Question:

  • If both do better with greater traits, what would coevolution lead to?

A mutualistic arms race!

Q: what if one outpaces the other?

(Co)Evolution of Interactions

Imbalance of armaments and the breakdown of mutualism

But also…

Many Possibilities

(left) mutualism maintained (right) mutualism from parasitism

It gets weirder…

Community Context

Indirect Consequences of Coevolution

Fly visits Babiana thunbergii

Malachite Sunbirds pollinate B. thunbergii

Inirect Effects may play big role

  • Q-matrix = direct
  • T-matrix = sum of indirect

Indirect effects drive coevolution in mutualistic networks, Guimarães et al 2017

Long-Tongue Fly Families

A Geographic Mosaic

  • Different fly pollination guilds
  • Partial spatial overlap
  • Coevolutionary hot/cold spots

Act III: The Geography of Coevolution

Spatially Distributed Populations

Metapopulation Concept

Powdery Mildew Parasitizing Plantago lanceolata

Spatial Host-Parasite System