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EXPLOITATION + Species 2 (predator P) Species 1 (victim V) - Getting into the mind of the predator: What constrains consumption? Inside the predator’s daily planner: 6:00 am 8:00 am 12:00 pm 1:00 pm 3:00 pm 3:20 pm 6:00 pm 7:00 pm 7:10 pm Get up and clean. Re-assert dominance in family group. Begin searching for victims. First victim spotted. Pursuit begins. Abandon pursuit. Victim got away. Second victim spotted. Victim subdued and killed. Begin eating. Victim eaten and sufficiently digested. Return home. Arrive home. Try to mate. Rest. The point is: 1. Time is limited. 2. There are other things to do besides activities related to eating (e.g. social interactions, mating rituals, grooming, sleeping). 3. Eating-related activities consist of: • • • • • searching for prey pursuing prey subduing the prey eating the prey digesting (may not always exclude other activities) The time constraints of foraging other essential activities foraging Foraging time The time constraints of foraging Foraging time other essential activities Handling time search handling Search time The time constraints of foraging Foraging time other essential activities Handling time eating pursuing & subduing Search time search eating pursuit & time subdue time Different species will allocate foraging time differently: Filter feeder: Sit & wait predator (spider) eating waiting eating subduing Time allocation also depends on victim density and predator status: Well-fed mammalian predator: Starving mammalian Predator (victims at low dnsity): eating eating pursuing & subduing searching pursuing searching & subduing Building a theory of predation: (After C.S. Holling) ts th Total search time per day Total handing time per day tmax t s th C.S. (Buzz) Holling Total foraging time is fixed (or cannot exceed a certain limit). 1) Define the per-predator capture rate as the number of victims captured (n) per time spent searching (ts): n ts 2) Capture rate is a function of victim density (V). Define a as capture efficiency. n aV ts 3) Every captured victim requires a certain time for “processing”. th hn t t s th n ts aV th hn n t hn aV n aV t 1 ahV n aV t 1 ahV 0.1 Capture rate 0.09 0.08 0.07 0.06 0.05 0.04 Capture rate limited by prey density and capture efficiency 0.03 0.02 0.01 0 0 50 Capture rate limited by predator’s handling time. 100 Prey density (V) 150 Damselfly nymph (Thompson 1975) Asymptote: 1/h Decreasing prey size The larger the prey, the greater the handling time. (Thompson 1975) Three Functional Responses (of predators with respect to prey abundance): Holling Type I: Consumption per predator depends only on capture efficiency: no handling time constraint. Holling Type II: Predator is constrained by handling time. Holling Type III: Predator is constrained by handling time but also changes foraging behavior when victim density is low. Per predator consumption rate Type I (filter feeders) Type II (predator with significant handling time limitations) Type I: n aV t Type II n aV t 1 ahV Type III (predator who pays less attention to victims at low density) victim density Type III n (aV ) 2 t 1 (ahV ) 2 Holling Type I functional response: Type I functional response Daphnia path Daphnia (Filter feeder on microscopic freshwater organism) Thin algae suspension culture Thick algae suspension culture Holling Type II functional response: Slug eating grass Cattle grazing in sagebrush grassland Holling Type III functional response: Paper wasp, a generalist predator, eating shield beetle larvae: The wasp learns to hunt for other prey, when the beetle larvae becomes scarce. We have seen that there are four possible outcomes for two-species competition. What are the possible outcomes for a predator-prey system? Gause’s Predation Experiments: Didinium nasutum eats Paramecium caudatum: Gause’s Predation Experiments: 1) Paramecium in oat medium: logistic growth. 2) Paramecium with Didinium in oat medium: extinction of both. 3) Paramecium with Didinium in oat medium with sediment: extinction of Didinium.