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The Ecology of Adaptive Radiation. QUICK. Think of the First Adaptive Radiation That Comes to Mind. Hawaiian Silverswords. Malagasy Vangids. Andean Espeletia. Ecological Opportunity.
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QUICK Think of the First Adaptive Radiation That Comes to Mind
Malagasy Vangids
Andean Espeletia
Ecological Opportunity “loosely defined as a wealth of evolutionarily accessible resources little used by competing taxa” (Schluter 2000, p.69).
Ecological Opportunity • Does it promote AR? • If so, how? • Is it required for AR? • Is it an operationally useful concept, or just a useful heuristic?
“Evolutionary divergence of members of a single phylogenetic lineageinto a variety of different adaptive forms.” Futuyma, Evolutionary Biology, 1998 Adaptive Radiation
AR Definitional Issues • Is pace of diversification part of the definition? In particular, does it have to be explosive? • How are ARs distinguished from non-ARs?
AR Definitional Issues • Is pace of diversification part of the definition? In particular, does it have to be explosive? • How are ARs distinguished from non-ARs? • Does it make sense to categorize a continuously distributed characteristic?
AR Definitional Issues • Is pace of diversification part of the definition? In particular, does it have to be explosive? • How are ARs distinguished from non-ARs? • Does it make sense to categorize a continuously distributed characteristic? • Is the number of species the appropriate metric?
AR Non-Adaptive Non-Radiation Non-Adaptive Radiation
What Prompts AR? • Classic idea (Simpson, Major Features of Evolution, 1954)—AR requires: • Geographical Access • colonization • Physical Access • mass extinction, appearance of new resource • Evolutionary Access • key innovation
“The evolution of a trait that allows a species to interact with the environment in an entirely new way” Key Innovation
“The evolution of a trait that allows a species to interact with the environment in an entirely new way” Key Innovation Note: 1. KI does not necessarily cause AR
“The evolution of a trait that allows a species to interact with the environment in an entirely new way” Key Innovation Note: 1. KI does not necessarily cause AR 2. KI refers to ecological diversity, not number of species
What Is the Evidence that Ecological Opportunity Prompts AR? • Biogeography: AR on islands • Paleontology: AR after mass extinction • Evolution: AR associated with KI • Phylogeny: Decline in diversification as radiation progresses
What is the Evidence that Ecological Opportunity Prompts AR? • Biogeography: AR on islands • Paleontology: AR after mass extinction • Evolution: AR associated with KI • Phylogeny: Decline in diversification as radiation progresses • Experimental evolution in lab
Ecological Opportunity • Does it promote AR? • If so, how? • Is it required for AR? • Is it an operational useful concept, or just a useful heuristic?
Adaptive Radiation Requires: • Adaptive Diversification • Speciation
Two Questions: • Is interspecific competition the only ecological interaction that can lead to AR? • Is speciation+diversification a one-step process or a two-step process?
Two Questions: • Is interspecific competition the only ecological interaction that can lead to AR? • Is speciation+diversification a one-step process or a two-step process?
Speciation Adaptive Differentiation 2 Steps
Speciation + Adaptive Differentiation 1 Step
Ecological Speciation Andrew P. Hendry. 2009. Ecological speciation! Or the lack thereof? Canadian Journal of Fisheries and Aquatic Sciences 66: 1383-1398.
Two Flavors of Ecological Speciation • Strong—in the presence of gene flow (sympatric/parapatric speciation) • Weak—divergence driven by natural selection regardless of geographic context
Strong Flavor: Speciation in presence of gene flow Speciation + Adaptive Differentiation 1 Step
Speciation in allopatry Adaptive Differentiation • 2-Step Models: • Does adaptive differentiation occur in allopatry or sympatry?
Why might NS favor different adaptations on different islands?
Nat Sel may be involved in speciation (weak ES); Competition not important Nat selnot part of speciation
Character Displacement Assumption: Body size correlates with resource use
Nat Selpart of speciation; Competition not important Nat selnot part of speciation
Nat Selpart of speciation; Competition not important Nat selnot part of speciation
Two Questions? • Is interspecific competition the only ecological interaction that can lead to AR? • Is speciation-diversification a one-step process or a two-step process?
Other Processes That Could Promote AR • Apparent competition/Competition for predator-free space (predation)
Apparent Competition • In Habitat A, Predator 1 preys on Prey 1 • More Prey 1 more Predator 1 • Predator 1 also opportunistically eats Prey 2 • More Predator 1 fewer Prey 2 • Hence, more Prey 1 fewer Prey 2 • “Apparent” competition • Promotes habitat divergence by Prey 2 to escape Prey 1’s predators
Other Processes That Could Promote AR • Apparent competition/Competition for predator-free space (predation) • Parasitism/Herbivory ecologically the same as above • Mutualism/Coevolution