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Consumption of one organism (the prey) by

another organism (the predator), in which
the prey is alive when the predator first
attack it

Taxonomic classifications of predators:

Carnivores, herbivores and omnivores

Functional classifications of predators:

xiii.True predators

• Kill their prey more or less immediately

after attacking them
• Over the course of their lifetime they kill
several or many different prey individuals
ii. Grazers

• attack large numbers of prey during their

lifetime, but they remove only a part of each
prey individual rather than the whole

• their effect on a prey individual is typically


• their attacks are rarely lethal in the short

term; never predictably lethal

iii. Parasites

• consume parts of their prey (their host)

• their attacks are typically harmful but

rarely lethal in the short term

• their attacks are concentrated on one or a

very few individuals during the course of
their life
iv. Parasitoids

• a hymenopterous or dipterous insect which

develops as a larva inside a host, eventually
killing it

• they belong to the order Hymenoptera &


• they are free living as adult, but they lay

their eggs in other insects

• the larval parasitoids then develops inside

its host individual, which is itself usually a

• they kill their host during the pupal stage

• often just one parasitoid develops from

each host
Peristenus diogneutis is a brown parasitic wasp about 3 mm
long (Day 1997). A native parasitoid of Lygus rugulipennis in
northern Europe, P. diogneutis only attacks plant-pest leaf
bugs such as Lygus. After mating, a female wasp will use her
ovipositor to inject a small egg into a Lygus nymph. Once the
egg is deposited in a host, it will emerge as a larva in 5 to 7
days (Day 1997). The newly emerged wasp larvae will feed
on the internal tissue of the dying Lygus for 7 to 10 days at
which time it pupates and drops to the soil below (Day 1996).
• parasitoids are intimately associated with
a single host individual, they do not
cause immediate death of the host, but
their eventual lethality is inevitable

• the rate of predation is determined by the

rate at which adult females oviposit

• The Effects of Herbivory on Individual


A.1. Plant compensation

• Herbivory can reduce self-shading

• Herbivory can lead to the mobilization of

stored carbohydrates

• Herbivory can lead to an altered pattern

of photosynthate (the energy-rich organic
molecules produced during
photosynthesis) distribution – to balance
the root/shoot ratio
• herbivory can lead to an increase in unit
leaf rate

• herbivory can lead to a reduced death rate

of plants part – there is compensatory
regrowth of defoliated plants when buds
that would otherwise remain dormant are
stimulated to develop

A.2. Herbivory and plant survival

• repeated defoliation can kill plants

• many seedlings are killed by herbivores

• smaller plants bear fewer seeds

• herbivory can lead to delayed flowering

• herbivory destroy reproductive structures

(flowers, fruits, seeds) directly
•in some cases, ‘herbivory’ of reproductive
tissues are mutualistic (animals that
consume pollen ; vertebrate fruit-eaters –
enhance its ability to germinate)

B. The effect of predation on a prey


• predatory attacks are often directed at the

weakest prey

• the impact of predation may also be

limited by compensatory reactions among
the survivors – the result of reduced
intraspecific competition

• predation reduced the density of a

population but net recruitment &
productivity of the survivors increase
because of reductions in intraspecific
Experiments in Predation
Gause, using the ciliates Paramecium and Didinium,
could not create a real system that would mimic
predator-prey cycles predicted by Lotka-Volterra.
However, the system could be maintained either by
occasional re-introduction of prey. In other words,
open system required to maintain predator-prey.
• prey populations reduced by their predator
will experience a compensatory decline in
the depressant effects of intraspecific

• the effects of parasitoids on their host, &

the effects of true predators on their prey
are straightforward – the fitness of their
host/prey is reduced to zero

• the effects of parasites on the hosts are

often not drastic – the parasite must have
some adverse effect on its host’s fitness

• predation-rate in parasites can be taken as

the rate at which host tissue & energy is
diverted from hosts to parasites
C. The effects of predation-rate on predator

• for parasitoids, every host successfully

attacked by a parasitoid represents an
incremental increase in parasitoid fitness

• for predators, herbivores & parasites, they

require a certain quantity of prey tissue for
basic maintenance; only when their intake
exceeds this thresholds that increases in
predation-rate lead to increased benefits to
the predator

• Food quality:
-The chemical composition & its
accessibility via digestion in herbivores

- toxic or digestibility-reducing secondary

D. The behaviour of predators

• predator can be classified based on their

diet width:

v. Monophagous (feeding on a single prey

vi. Oligophagous (few prey types)
vii.Polyphagous (many prey types)

• Specialists (broadly, monophagous &


• Generalists (polyphagous)

• Parasitoids – typically specialized & are

often monophagous

• Herbivores – grazing & ‘predatory’

herbivores are polyphagous

• Parasitic herbivores are often specialized

2 basic points to understand the patterns of
diets width among predators:

iv. Predators choose profitable prey

• Natural selection; evolution favors those

individuals with the highest fitness;

• Profitable prey items are the greatest gain

(in terms of energy intake per unit time
handling prey)

ii. Predators & their prey are likely to have


• Animals & plants have evolved in

response to selection pressures
originating from the other animals &
plants in their environment
• A continuous selection pressures on prey
to avoid death or fitness reduction at the
hands of predators
• continuous selection pressures on
predators to increase their fitness by
exploiting their prey more effectively

• coevolution provides an added force in

the restriction of diet width

Time & timing

• levels of specialization need not involve

morphological adaptation, but the result of
differing degrees of temporal coincidence

• some animals switch their food-

preference seasonally

• the effects of differing lengths of life cycle

among predators & their prey

• some exhibit a pattern of abundance that

reflects the yearly changes in
environmental quality
B.1. Functional responses: consumption
rate & food density

• Consumption rate rises with prey density

but gradually decelerates, until a plateau
is reached at which consumption rate
remains constant irrespective of prey

• A consumer has to devote a certain

handling time to each prey item it
consumes (pursuing, subduing &
consuming the prey item, and then
preparing itself for further search)

• Characterized by a handling time & an

attack rate
Numbers eaten


40 80

Prey density

Functional response of the predators in

relation to the prey density

• common in herbivore - plant, predator-

prey & parasitoid - host interaction
Pe = the no. of prey items eaten by a

a’ = searching efficiency or attack rate of

the predator
Ts = searching time
N = the density of prey items
Th = handling time

Pe = a’TsN

Ts = T – ThPe
(ThPe = the total time spent handling

Pe = a’(T - ThPe)N

Pe (1 + a’ThN) = a’NT
Pe = a’NT

1 + a’ThN


Handling time
Attack rate

0.5 0.1

1.0 3.0
Mean prey size

Attack rate & handling time for these functional

• Handling-time is determined by:

iii. The time spent pursuing & subduing an

individual prey
iv. The time spent eating (or ovipositing in)
each prey
v. The time spent resting or cleaning or
fulfilling any other essential function (like
digestion) prior to the act of predation

• Searching efficiency (attack rate) will

depend on:

ix. The max distance at which a predator

can initiate an attack on a prey
x. The proportion of these attacks that are
xi. The speed of movement of the predator
& prey (their rate of encounter)
xii.The ‘interest’ taken by a predator in
obtaining prey as opposed to fulfilling
other essential activities
Mathematical Models
2. Host-parasitoid models

Ht: the number of hosts

Pt: the number of parasitoids (in generation
r: the intrinsic rate of natural increase of the
c: the conversion rate of hosts into
parasitoids (the mean no. of parasitoids
emerging from each host)
Ha: the number of hosts actually attacked by
parasitoids (in generation t)
Ht+1 = er(Ht – Ha)
Pt+1 = cHa
(ignoring intraspecific competition &
assume that each host can support only
one parasitoid (c = 1)):
Pt+1 = Ha

Et: the no. of host-parasitoid encounters (or

interaction) in generation t

A: the proportion of the hosts encountered

by any one parasitoid (parasitoid’s
searching efficiency)

Et = AHtPt
Et = APt

• encounters occur randomly (Poisson
distribution; the proportion not encountered
at all, po:

po = exp Et

Thus, the proportion that is encountered is
1 – po, and the no. attacked is:
Ha = Ht (1 – po) = Ht 1 - exp

Or, Ha = Ht 1 – exp (- APt)

Substituting this expression for Ha into

equation Ht+1 = er(Ht – Ha) :

Ht+1 = Htexp(r - APt)

Pt+1 = Ht 1 – exp (- APt)

Nicholson-Bailey model of the host-
parasitoid interaction:

c) Parasitoid numbers are determined

solely by the rate of random encounters
with hosts
d) Host numbers would grow exponentially
but are removed by random encounter
with parasitoid