Coevolution and Mutualisms Species interactions may be mutually beneficial In some cases, these interactions may lead to coevolution among participants What is coevolution? Evolutionary change in one species causes evolutionary change in the other Species 1 Five points about coevolution and specialization 1). Outcomes unlikely to be equal 2). Coevolution may occur in one population, but not another 3). Population-level variation in degree of specialization Species 2 (Thompson 1994. The Coevolutionary Process) Five points about coevolution and specialization 4). Variation in response leads to geographic mosaic of coevolution 5). Temporal variation in mosaic as well. Coevolution difficult to document Many interactions are highly specialized but not coevolutionary Evolutionary response may be to generalized type of organism: diffuse coevolution. May also be specifically between 2 species- “narrow sense” (Thompson 1994. The Coevolutionary Process) 1 Example: rabbit control in Australia European rabbits introduced into Australia, spread rapidly Gene-for-gene mechanisms of coevolution 1. single-allele complementary system Introduction of myxoma virus Coevolved relationship between resistance and virulence Mutualisms 2. elicitor-receptor model 3 types of mutualisms •Three basic types of mutualistic interactions 1. Trophic (obligatory) •May or may not involve coevolution 2. Defensive (both facultative and obligatory) •Obligate versus facultative mutualism •Cheaters Trophic mutualisms 3.Dispersive (some obligatory, most not) Defensive Mutualisms Copyright: Hilton Pond Center 2 Dispersive Mutualisms Pollination versus seed dispersal Pollination systems sometimes coevolved, seed dispersal systems are not Very great degree of specialization in some pollinator systems Seed dispersal: mutualistic but not coevolved Why not coevolved? No specialization Sites suitable for seeds not predictable No “payment on delivery” Pollination: mutualistic, not always coevolved Target specificity Need for predictability Advantages to non-specialization (Wheelwright and Orians 1982) “How to be a fig” A stunning example of coevolution… Two paths to pollination (and seed dispersal) Deception Photo by Ken Gardiner Reward Photo by Babs and Bert Wells 3 Trade-off in mutualisms How do you maximize your benefit for your cost? The plant perspective… Mutualisms: how do they work?? •Theory poorly developed •Apparent evolutionary instability •Cheaters The animal perspective… (Ferriere et al. 2002) Specialization and phylogeny Specialization may already exist“preadaptation” Phylogeny can often explain preadaptations and intermediate steps Implications?.... Summary •Mutualisms offer benefits to both participants •Mutualisms have enormous impact on natural communities •Coevolution can occur in mutualisms but appears rare, hard to demonstrate •Mutualisms represent a cost-benefit balance between participants Ungraded homework Assignment What are two ways plants can attract pollinators, and what is the cost or risk associated with each strategy? 4