Survey
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
* Your assessment is very important for improving the work of artificial intelligence, which forms the content of this project
The major types of organism-organism interactions Predator-prey interactions Dik Heg 1. Competition Interspecific competition (competition between different species) Intraspecific competition (competition within the same species) 2. Predation Interspecific predation (predator-prey interactions) Intraspecific predation (cannibalism, infanticide) 3. Cooperation Interspecific cooperation (mutualism, symbiosis) Intraspecific cooperation (kin selection, reciprocal altruism) 4. Parasitism Interspecific parasitism (host-parasite interactions, e.g. ectoparasites, endoparasites, viruses, pathogens) Intraspecific parasitism (within-species brood parasitism, e.g. egg dumping, sneaking) I. Prey detection: stages of predation and the antipredator defences Overview I. Prey detection 1. Encounter rarity, hiding, activity budget 2. Detection immobility, crypsis, confusion, sensory limits 3. Identification masquerade, confusion, Mullerian mimicry, Batesian mimicry, honest warning colorations 4. Approach (attack) 5. Subjugation (prevent escape) 6. Consumption II. Optimal foraging theory III. Predator-prey population dynamics Overview Prey detection 1 Example apostatic selection Prey selection Predators might select prey based upon: - Species - Size - Morph etc. etc. Apostatic selection: predator preys on more common type of prey Prey detection Prey detection Reasons for apostatic selection 1. Formation of search image It takes time to learn to detect and handle prey: known prey preferred 2. Optimal search rate hypothesis Optimal search speeds for prey 1 and 2 differ: first search for more abundant prey 1, until 1 < 2, than switch search rate strategy to 2 3. Search intensity hypothesis (´stare duration´) Prey species 2 might be more difficult to detect Search rate increase: trade-off between increase in encounter frequency and a reduction in detection probability Prey detection 2 Cryptic prey Cryptic prey: colour, morphology and/or chemical profile resembles a random sample of the background as perceived by the predator (at the time and place at which the prey is most vulnerable to predation) Argentine horned frog Prominent moth Sandgrouse chicks Leaflitter frog Prey detection Aposematic prey Aposematic colouration (or sounds, odours etc.): conspicuous colouration combined with noxiousness (incl. unpalatability) Predator training and hence protection more efficient when: 1. Higher density of the signal 2. Higher densities of the aposematic prey 3. Fewer or more similar aposematic colour patterns around 4. Clumped- or higher local densities of prey Prey detection 3 Aposematic prey Mimicry Mimicry: two species resemble each other (in morphology and often also in behaviour). How can natural selection favour a defence that operates during or after the subjugation and consumption phases of predation? Species 1 Species 2 Batesian mimicry 1. Kin selection (Fisher 1930) 2. Synergistic selection (Guilford 1985, Maynard-Smith 1989) 3. Combination with a common post-attack defence (Wicklund & Jarvi 1982) Müllerian mimicry Batesian: one palatable species resembles unpalatable species Müllerian: two unpalatable species resemble each other Mimicry ring: species complex of groups of Batesian and Müllerian mimics Prey detection Example: pairs of Batesian mimics Unpalatable swallowtail model species Palatable female races of Papilio memnon Prey detection Example: pairs of Müllerian mimics from Ecuador and Northern Peru Heliconius melpomene H. erato 4 1. H. cydno morphs have increased survival when they match locally abundant co-model species when transferred at low densities (a, c & d). 2. H. cydno morphs do not differ in survival when many H. cydno morphs are transferred leading to high densities of both models (b). Example: Müllerian mimics of Heliconius Populations A: `white` Müllerian mimics Populations B: `yellow` Müllerian mimics Control (normal transfer, A to A or B to B) Experiment (reciprocal transfer, A to B or B to A) H. cydno Co-models Kapan (2001). Nature 409: 338-340 Kapan (2001). Nature 409: 338-340 The diet-switch model II. Optimal foraging theory (McArthur & Pianka 1966) The aim of optimal foraging theory is to predict the foraging strategy to be expected under specified conditions Major assumptions: 1. Natural selection on foraging in the past = present 2. High net rate of energy intake = high fitness 3. Experimental setup = close to natural environment Specialist: only pursues profitable prey, but has to search more. Generalist: also eats more and less profitable prey, has to search less. E = energy content h = handling time prey s h s = search time prey ith prey = the next most-profitable prey type s h Optimal strategy: predator should also pursue ith prey when: Ei / hi > E / (s + h) Optimal foraging theory Optimal foraging theory 5 The diet-switch model The constrained diet model (McArthur & Pianka 1966) (Belovsky 1978-1984) Some general predictions from the model: 1. Predators with short handling times should be generalists. 2. Predators should be more generalistic in an unproductive environment (low density = large search time). 3. Predators should ignore insufficiently profitable ith prey irrespective of their density (formula is si independent). Example using linear programming with herbivore moose Alces alces: Consumption Energy value Sodium (gram) per gram per gram Aquatic plants A 3.8 kJ 0.003 gram Terrestrial plants T 4.25 kJ 0.000 gram How much A and T can and should the moose eat to survive? Optimal foraging theory Maintenance requirement: 14000 kJ < 3.8A + 4.25T Sodium requirement: 2.57 gram < 0.003A (minimum energy needed for survival per day) (Only aquatic plants contain sodium, gram needed per day) Optimal foraging theory Constraints, limitations, currencies to be optimized are very often individual (phenotypic/genotypic), population and/or species specific! Predator present Predator (gram of wet food per day which can be maximally processed, aquatic plants contain 5x more water) Number of fish Digestive limitation: 32900 gram > 20A + 4.04T No predator present % Vegetation cover where prey is feeding Optimal foraging theory largemouth bass Micropterus salmoides Prey bluegill sunfish Lepomis macrochirus Optimal foraging theory 6 The functional response The numerical response Functional response (Solomon 1949): the relationship between individual‘s consumption rate (P) and local food density (N) Numerical or aggregative response: the relationship between the local consumer density and local food density. Three types of functional responses (Holling 1959): Type 1 Type 2 Type 3 No handling time Max P = maximum throughput At large N: predator only handling prey Switching, search image, increased (handling) efficiency Optimal foraging theory Patch use: the marginal value theorem Interference: functional response and numerical response interact Given: prey distributed in patches (which might differ in quality) prey within patches are depleted by foragers forager pays travel cost to reach other patch This will lead to animals distributing themselves over the habitats or patches until fitness: Question: at what point should a forager leave a patch? Equal: Ideal free distribution Unequal: Ideal despotic distribution (dominants occupy best patches or have less negative effects from interference) Optimal foraging theory Optimal foraging theory 7 Patch use: the marginal value theorem III. Predator-prey population dynamics The basic dynamics of predator-prey and plant-herbivore systems: a tendency towards cycles. Why?? Fig. 10.1c Begon p.370 Optimal foraging theory Population dynamics The Lotka-Volterra Model The Lotka-Volterra Model Description of the changes in abundance of predators and prey: P = number of predators (or consumers) N = number of prey (or biomass) Predator numbers are assumed to decline exponentially through starvation in the absence of prey, q is the mortality rate: Without predators, exponential increase in N per time step t: dP = -qP dt dN = rN dt This is counteracted by predator birth, which depends on predator consumption a´PN and the efficiency f of converting this food into predator offspring: As predator or prey densities increase, more predators encounter prey (PN) and prey will be consumed with ´attack rate´ a´: dP = fa´PN - qP dt dN = rN – a´PN dt Population dynamics Population dynamics 8 The Lotka-Volterra Model Predator abundance (P) The Lotka-Volterra Model Prey abundance (N) Despite the underlying tendencies, predator-prey cycles are not always to be expected! Population dynamics Parasitoid present Population dynamics Example: effects of piscivore predator on group-living in the Lake Tanganyika cichlid Neolamprologus pulcher Piscivore: Lepidiolamprologus elongatus host Plodia interpunctella No parasitoid present parasitoid Venturia canescens Group-living fish: Neolamprologus pulcher Population dynamics 9 Lake Tanganyika 10 Cages (2x2m) Piscivore (control, medium, large) Survival (a) breeder males breeder females large helpers medium helpers small helpers 17 19 29 63 survival 42 medium helpers 23 20 22 0.8 24 48 95 45 40 0.6 large helpers 1.0 survival 1.0 92 47 0.4 control (b) medium 0.8 small helpers 0.6 0.4 large 5 predator treatment 10 15 20 25 number of adults 60 Reproduction (a) Feeding rate 50 Large helpers number of offspring 70 fry 60 juveniles 50 40 30 20 Medium helpers 10 0 Feeding rate (bites/minute) 40 30 20 10 0 60 (b) 50 40 30 20 n = 18 20 18 control medium large 18 20 18 control medium large 10 0 0 predator treatment 20 40 60 80 100 Mean distance from shelter (cm) 11