Stano Pekár“Populační ekologie živočichů“  dN = Nr dt True predators - catch several animals and gain sustenance for their own fitness (spiders, lions) Parasitoids - consume about single host, free adults but larvae developing on or within a host, consuming it prior to pupation (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), stenophagous (few prey types)  oligophagous (more prey types)  polyphagous/euryphagous (many prey types) - not capable of consuming all prey types  type can based on taxonomy, size, sex, ontogenetic stage  predators choose most profitable prey - select prey items for which the gain is greatest (energy intake per time spent handling) Krebs (1978)  Ecological context – stenophagy vs. euryphagy/polyphagy  Evolutionary context – generalist vs. specialist Euryphagous generalist Stenophagous generalist Euryphagous specialist Stenophagous specialist Pekár & Toft (2015) Ecological dimension Evolutionary dimension  predators tend to specialise to a greater or lesser extent during evolution - monophagy evolved where prey is abundant and exerts pressures which demands adaptations (e.g. morphological, behavioural, metabolic) - polyphagy evolved where prey was unpredictable  true predators - majority are polyphagous  parasites - commonly monophagous due to intimate association with hosts, their life-cycle is tuned to that of their host  parasitoids - often monophagous but some are polyphagous presumably because adults are free living  herbivores - rather polyphagous, many insect herbivores are specialised as a result of adaptation to plant secondary metabolites (Drosophila pachea consumes rotten tissues of Senita cactus which contain poisonous alkaloids)  even polyphagous predators prefer certain prey - constant preference irrespective of prey density - switching to more common prey Thais preferred Mytilus edulis over M. californianus Murdoch & Oaten (1975) Murton et al. (1964) Seasonal shift in Columba  predation has positive effect on population of prey because reduce intraspecific competition - stabilise prey population dynamic  true predators and parasitoids reduce fitness of individual prey to „0“ - Mustela consumed mainly solitary and injured individuals, so it has little effect on the Ondatra population growth  caterpillars defoliate partially so that re-growth can occur, but cause reduction in fertility  parasites - reduce fitness partially, effect is correlated with the burden Negative effect of mite parasites on Hydrometra Lanciani (1975)  mortality of prey increases with the prey density due to predation Total response of a predator is composed of: - individual response to changing prey density  functional response - population response to changing desnity of prey  numerical response  Holling (1959) found that predation rate of individual predator increased with increasing prey density - defined three types of functional responses - more types were defined later 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 (due to handling) 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 develop search image (e.g. respond to kairomones)  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 or “search rate” Type I  consumption rate of a predator is unlimited  TH = 0 so Sa aHTH  HS TTT  STT  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 so 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 HS TTT  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 (P): 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( dPfaHP t P  d d  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 captured 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 microcosmos 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 refuge types stabilise the interaction Connell (1970)