MODULARIZACE VÝUKY EVOLUČNÍ A EKOLOGICKÉ BIOLOGIE CZ.1.07/2.2.00/15.0204 True predators - consume several animals and gain sustenance for their own fitness (spiders, lions) Parasitoids - free adults but larvae developing on or within a host, consuming it prior to pupation, consume about single host (Hymenoptera, Diptera) Parasites - live in close association with a host, gain sustenance from the host, but often do not cause mortality (Acari, Trematodes) Herbivores - feed on plants, may totally consume plants (seedeaters) or partially (aphids, cows) monophagous (single prey type) oligophagous (few prey types) polyphagous/euryphagous (many prey types) - not capable of consuming all prey types predators choose most profitable prey - select prey items for which the gain is greatest (energy intake per time spent handling) Krebs (1978) mortality of prey increases with the prey density due to predation Total response of a predator - increasing consumption rate of individual predators → functional response - increasing consumption of popualtion of predators → numerical response Holling (1959) found that predation rate increased with increasing prey population density - defined three types of functional responses Type I number of captured prey is proportional to density - prey mortality is constant less common found in passive predators (web-building spiders) the handling time exerts its effect suddenly Daphnia feeding on Saccharomyces - above 105 cells Daphnia is unable to swallow all food Rigler (1961) Type II predators cause maximum mortality at low prey density as prey density increases, search becomes trivial and handling takes up increasing portion of the time saturation of predation at high densities - prey mortality declines with density Thompson (1975) Ischnura eating Daphnia Type III when attack rate increases or handling time decreases with increasing density predators respond to kairomones predators develop search image polyphagous predators switch to the most abundant prey - prey mortality increases then declines Notonecta switched from Cleon to Asellus based on its abundance Lawton et al. (1974) T .. total time TS .. searching time - searching for prey TH .. handling time - handling prey (chasing, killing, eating, digesting) H .. prey density Ha .. number of captured prey a .. capture efficiency, “area of discovery”, or “search rate” Type I consumption rate of a predator is unlimited TH = 0 Sa aHTH = HS TTT += Type II consumption rate of a predator is limited because even if no time is needed for search, predator still needs to spend time on prey handling TH > 0 predator captures Ha prey during T Th .. time spent on handling 1 prey at low density predator spends most of the time searching, at high density on prey handling haH THT = aH H TaHTH a SSa =→= aH H THTTT a haSH +=+= h a aHT aHT H + = 1 n h n a HaT aTH H + = 1 Type III consumption increases at low densities and decreases at higher densities n .. rate of increased consumption at higher densities if n = 1 → Type II a .. rate of increase at low densities H Ha T/Th a 0 n Increase of predator population may result from: increased rate of reproduction - the more prey is consumed the more energy can predator allocate to reproduction - delayed response parasitoids - one host is sufficient predators, herbivores, parasites - certain quantity of prey tissue is required for basic maintenance = lower threshold Growth rate in Linyphia Turnbull (1962) conversion of prey into predator numbers f .. conversion efficiency d .. mortality of predators Ivlev (1955) model V .. amount of prey a .. search rate f .. conversion efficiency d .. mortality of predators H rp 0 dear fV −−= − )1( dPfaHPr −= attraction of predators to prey aggregations - immediate response - aggregated distribution makes search of predators more profitable instead of concentration on profitable patches perspective predators and prey may play “hide-and-seek” Huffaker (1958): Typhlodromus fed upon Eotetranychus that fed upon oranges - Eotetranychus maintained fluctuating density - addition of Typhlodromus led to extinction of both Experimental setup Eotetranychus population dynamic Predator-prey dynamic making environment patchy - by placing Vaseline barriers - facilitating dispersal by adding sticks each patch was unstable but whole cosmos was stable - patch with prey only → rapid increase of prey - patches with predators only → rapid death of predator - patches with both → predator consumed prey Sustained oscillations of the predator-prey systemAltered experimental setup Refuge For fixed proportion of prey - certain proportion of Ephestia caterpillars buried deep enough in flour are not attacked by Venturia with short ovipositors For fixed number of prey - adult Balanus occur in the upper zone where Thais can not get during short high tide thus consumes only juveniles - a fixed number of Balanus is protected from predation irrespective of Thais density both refuges stabilise the interaction Connell (1970)