Chapter 1.

Nature of
Biological Control Agents
ANGELYN C. RODULLO
Instructor 1
Lesson 1. Parasitoids, Pathogens &
Predator of Insect & Vertebrate
Pests
 BIOLOGICAL CONTROL
• An approach that uses the natural enemies to
suppress/manage pests.
• Form of enhancing natural defenses to achieve a
desired effect.
• Raising & releasing one insect to have it attack
another, almost like a “living insecticide
Durable biological control program depends on
two main strategies:
1.Using ecological farm design to make your
farm more attractive to biological control
“agents”
2.Introducing beneficial agents onto your
farm
As natural occurring enemies would do,
introduced beneficials may:
 Produce antibiotics parasitize target
organisms
 Form physical/chemical barriers to
infection out compete plant pathogens for
niches
 Simply help the plant grow better, masking
symptoms where disease is present
PARASITOIDS
PARASITOIDS
 PARASITOIDS
 Parasitic insect that lives in/eventually kills a larger host
insect (other arthropods).
 Most parasitoids (parasitic insects that kill their hosts) live
freely & independently as adults; they are lethal &
independent only in their immature stages.
 They attack hosts larger than themselves, eating most/all
their host’s bodies before pupating inside/outside them.
 When parasitoids emerges from its pupa as an adult, it
usually nourishes itself on honeydew, nectar/pollen
 PARASITOIDS
 Some adults make meals of their host’s body fluids & others requires
additional water.
 Adult female parasitoids quickly seek out more victims in which to lay
their host killing eggs. With their uncanny ability to locate even
sparsely populated hosts using chemical cues, parasitoid adults are
much more efficient than predators at ferreting out their quarry.
 Diff. parasitoids can victimize different stages of the same hosts,
although specific parasitoids usually themselves to one stage.
 Thus parasitoids are classified as egg parasitoids, larval (nymphal)
parasitoids or adult parasitoids.
 PARASITOIDS
 Parasitoids can be classified also as either ectoparsites or endoparasites
depending, respectively, on whether they feed externally on their hosts/develop
inside them. Their cycle is commonly short, ranging from 10 days to four weeks.
 Most beneficial insect parasitoids are wasps or flies, although some rove beetles
(salagubang) & other insects may have life stages that are parasitoids.
 Insect parasitoids only attack a particular life stage of one/several related
species.
 Immature parasitoids develops on/within a pest, feeding on body fluids & organs,
eventually leaving the host to pupate or emerging as an adult.
 Life cycle of the pest & parasitoid can coincide/ that of the pest may be altered
by parasitoid to accommodate its development.
 Characteristics of Parasitoids
a.Parasitoids usually destroy their hosts during development.
b. The parasitoid's host is usually in the same taxonomic class
(Insecta).
c. Parasitoids are large relative to their hosts.
d. Parasitoid adults are free living while only the immature stages are
parasitic.
e. Parasitoids develop on only one host individual during the immature
stages.
f. With respect to population dynamics, parasitoids are similar to
predatory insects.
 Mode of Development of Parasitoids
1. (With respect to the host)

1.Endoparasitoid (internal): usually in

situations where the host is exposed.
2.Ectoparasitoid (external): usually in
situations where host lives within
protected location (e.i. Leafminers, in
cocoons, under scale covers).
 Mode of Development of Parasitoids
2.(With respect to numbers of immature per individual
host)

1.Solitary Parasitoid
2.Gregarious Parasitoid
 Mode of Development of Parasitoids
3. (With respect the host stage)

1.EGG
2.LARVAE
3.PUPA
4.ADULT
5.COMBINATIONS OF THE ABOVE (I.E., Egg-larval
parasitoid)
Mode of Development of Parasitoids
3. (With respect the host stage)
 Mode of Development of Parasitoids
4. (With respect to affect on host)

1. Idiobionts: host development arrested or

terminated upon parasitization (e.g., egg
parasitoids)
2. Koinobionts: Host continues to develop
following parazitizations (e.g., larval-pupal
parasitoids)
EGG PARASITOIDS: Larval Parasitoids
Trichogramma wasps
 Mode of Development of Parasitoids
5. (With respect to other parasitoid species)

1. Primary parasitoids
2. Secondary parasitoids
(Hyperparasitism)
3. Tertiary parasitoids
(Hyperparasitism)
 Mode of Development of Parasitoids
6. (Competition among immature parasitoid stages)

1. Intraspecific competition: Superparasitism

2. Interspecific competition: Multiple
parasitism
 TYPES OF HYPERPARASITISM

a. Direct: The situation when a secondary

parasitoid searches out and deposits its egg
in or on the body of its parasitic host (a
primary parasitoid), which may or may not
be contained in the body of a living
phytophagous insect.
 TYPES OF HYPERPARASITISM

b. Indirect: The situation when a secondary

parasitoid searches out and deposits its egg
in the body of an unparasitized, nonparasitic
host. The egg usually remains undeveloped
until the nonparasitic host is subsequently
parasitized by a primary parasitoid, which
then serves as a host for the secondary
parasitoid.
 TYPES OF HYPERPARASITISM

c.Facultative: Situation in which secondary

parasitoid may develop as a primary
parasitoid under the right conditions.
d. Obligatory: Situation in which
secondary parasitoid can only develop
within or on another primary parasite.
 TYPES OF HYPERPARASITISM

e. Autoparasitism: Situation in which the male of the species

develops as a hyperparasite (sometimes of the female of the
same species) and the female develops as a primary parasite.
These species may be referred to as “heteronomous” because
male and female develop on different hosts.
f. Cleptoparasitism: Not true hyperparasitism. Situation in
which parasitoid preferentially attacks hosts already parasitized
by another parasitoid species and then competes with first
parasitoid for nutrients in the host. The cleptoparasite usually
wins the competition.
Mode of Reproduction in Parasitic Hymenoptera
(Ants,wasps,hornets,sawflies,bees)
 Mode of Reproduction in Parasitic Hymenoptera

a. All hymenoptera display the phenomenon of haploid

parthenogenesis. This means that the unfertilized egg
undergoes parthenogenetic development to produce normal,
viable adult male which is haploid. All fertilized eggs develop
into diploid females.
b. There are variations on basic pattern which cause
extremely important differences in the mode of reproduction.
c. The variations may be divided into 3 types of reproduction:
 Mode of Reproduction in Parasitic Hymenoptera

1. Arrhenotoky: This is the basic mode. Unfertilized eggs produce males and
fertilized eggs produce females. May be referred to as "biparental" species. Male
or female offspring can be produced through internal or external control of
fertilization in some species.
2. Deuterotoky: Unmated females produce both male and female (mostly)
progeny. Males produced are biologically and ecologically nonfunctional. Such
species are called "uniparental". A diploid condition is produced in the female
progeny through various cytogenetic mechanisms.
3.Thelyotoky: Only female progeny are produced. Males are unknown. Also called
"uniparental". Males can be produced when female adults are reared under
extreme temperature conditions or if female adults are feed antibiotics to kill
endosymbionts (e.g., rickettsia in the genus Wolbachia) that make them sterile
through cytoplasmic incompatibility
 ARRHENOTOKY

It occurs in members of the

Insect Order Hymenoptera
(bees, ants,wasps) and the
Thysanoptera (thrips). Also in
spider mites, Hemiptera,
Coleoptera (bark beetles)
Honey bees produce
malesv(drones) by
parthogenesis;Arrhenotoky.
 Thelytoky

form of
parthenogenesis
wherein female
offspring develops
from an unfertilized
egg. E.i Aphids
Physiological and Behavioral Aspects of
Reproduction in Parasitoids
 a. Premating Period

• A premating period after emerging from the pupal

stage is generally not a necessary characteristic of the
parasitic life. In most parasitoids mating is almost
immediate after emergence of pupa.
• This is aided in nature by the males of parasitic
hymenoptera having a shorter developmental period
than females which allows them to precede the
females to the adult stage by 1 or 2 days.
 b. Mating Habits

 In many species
courtship behaviour
has been recorded
(e.g., Aphytis species).
In some species
females tend to ignore
males after mating
occurs.
 c. Preoviposition Period.

• This includes the interval between emergence of the adult

female and deposition of the first egg. In some species it
does not occur, in some it is facultative, and in some it is
obligatory. Occurrence is usually associated with nutritional
requirements of the adult female.
• With respect to egg production females can be divided into
2 groups:
1. Proovigenic
2. Synovigenic
 1. Proovigenic

• Those species of hymenoptera which reach the

adult stage with a complete complement of
ripe eggs which they deposit in a short time.
No more eggs are produced during the
parasitoid’s life.
 2. Synovigenic

• Those species (most parasitoids) of

hymenoptera which continue to produce eggs
throughout the adult stages. Production of
eggs is dependent on the nutrition of the adult
female rather than on the metabolites
retained from the immature stages.
• a. Adult nutrition. A source of protein is required for continuous production of
the eggs. This protein may be provided by aphid honeydew, plant nectaries, and
host feeding.
• b. Host feeding and host mutilation. Wounds made by the parasitoid’s ovipositor
give rise to host body fluids which the parasitoids feeds upon. Actual “feeding
tubes” may be formed by secretions from the ovipositor. Host mutilation may
have been the first step in the evolutionary development of host feeding. Host
feeding usually renders host unsuitable for development of immature parasitoids.
• c. Oviposorption. When the synovigenic female does not obtain proteinaceous
food or is unable to find hosts for a period of time, the ripe eggs in the ovarioles
are absorbed. Eggs can be produced at a later time when food or hosts are found.
The observation of reproductive material is correlated with a high searching
capacity.
d. Behavior in Host Selection
1. Host habitat finding.
2. Host finding.
3. Host acceptance.
4. Host suitability.
 Developmental Stages of Parasitoids

a. Egg types. Forms of eggs in entomophagous parasites are not uniform. Many
adaptive modifications are superimposed on those differences due to phylogeny.
b. Embryology. Usually the parasitoids deposits partially or completely incubated
eggs in, on, or apart from the host and the larvae upon hatching commence to feed.
These are monoembryonic eggs. A special exception to this polyembryony.
c. Polyembryony: The production of many parasite larvae from a single egg. The
polar nuclei give rise to an embryonic membrane called the “trophamnion”. The
trophamnion surrounds the embryonic area of the egg and functions in a nutritive
fashion by extracting and concentrating nutrients from the host haemolymph to the
embryonic area. The embryonic area divides into small groups of cells (morulae)
within the trophamnion which lengthens into a chain-like structure. When the
trophamnion breaks up the morulae form separate embryos. As many as 1500
embryos may be produced in Copidosoma.
Braconidae: a family of Parasitoid wasps
Encyrtidae: large family parasitic wasps;
Superfamily; Chalcidoidea
Platygasteridae exclusively parasitoids wasps, mostly verysmall, black, and
shining, w/geniculate antellate antennae that have an eight-segmented
flagellum. The wings may have lack venation, though they may have slight
fringes of setae.
Dryinidae solitary wasps; the larvae are parasitoids
of the nymphs and adults of Auchenorrhyncha
 Characteristics of polyembryonic species are:

1. Oviposition into egg or young larvae of host with parasitoid

maturity obtained in mature host larvae or pupae;
2. Exceptionally large numbers of progeny (1000 t0 3000)
develop within the host;
3. Simultaneous development and emergence of progeny
from a host individual; and
4. Major proportion of the broods consist of only 1 sex or
mixed broods with widely varying sex ratios.
 d. First-Instar Larvae

• The most distinctive parasitic stage in the life cycle is the primary or
first instar. Subsequent larval stages are usually grub-like without any
conspicuous structures. The main types of first instar larvae
recognized among the parasitoids or egg predators are:
1. Triungulin 2. Planidium 3. Sacciform 4. Teleaform
5. Mymamariform 6. Cyclopiform 7. Eucoiliform 8.Mandibulate
9. Microtype 10. Muscoidiform 11.Vesiculate 12. Caudate
13. Hymenopteriform 14. Chrysidiform 15. Agriotypiform
e. Planidium larvae: First-instar larvae that have elongated setae arising
from the thoracic or caudal regions to assist them in moving to their
hosts after hatching from the egg.
f. Triungulin larvae: First-instar larvae that have segmented legs that
assist them in moving to their hosts after them hatching from the egg.
g. Intermediate and Mature Larvae. The larva usually exhibits a change
of form other than size as it develops through its various instars. The
form change may be subtle, but because hypermetamorphosis is the
general rule among endoparasitic larvae, there usually are conspicuous
modifications between certain instars. Generally the larva loses any
bizarre first-instar characteristics as it develops toward maturity.
h. Pre-pupae. Stage when the last larval instar ceases to feed prior to
pupation and exhibits little movement.
1. During this time the protocdaeum (hind gut) links up with the mid
gut. These have been separated during development to avoid fecal
contamination of parasitoid’s environment.
2. Connection of mid and hind gut allows excretion of material. Fecal
matter is referred to as meconia.
i. Pupae. Most parasitic larvae pupate within the host remains. They
may pupate within the host cocoon or puparium or in a mine or tunnel
produced by a host.
 Ovipositional Sites and Developmental
Characteristics
A. Eggs or larvae deposited before apart from the host.
1. Eggs hatch or larvae deposited before host contact.
2. Eggs hatch after ingestion by host.
B. Eggs or larvae deposited on or near host.
1. Larvae develop externally on host.
a. Among eggs (egg predators)
b. In hosts concealed in cocoons, puparia, mines, galleries, galls, etc., or under
scale covers
c. On exposed hosts.
2. Larvae develop internally in host.
 Ovipositional Sites and Developmental
Characteristics
C. Eggs, or more rarely larvae, deposited in host.
1. Deposited in host egg.
a. Development completed in host
b. Development completed on other eggs as egg predators
c. Development completed in larva, prepupa, or pupa of host
2. Deposited in host larva or hemimetabolous insects.
a. Development completed in host larva or in nymphs
b. Development completed in host pupa
c. Development completed in homopterous nypmhs or adults
3. Deposited in host pupa.
4. Deposited in host adult.
D. Oviposition site correlated with sex of the
egg
 COMMON PARASITOIDS

Order Family Host or Prey Mode of Attack

Diptera (True flies) Tachinidae Beetles, butterflies, moths Internal
Nemestrinidae Locusts, beetles Internal
Phoridae Ants, caterpillars, termites, flies, others Internal
Crytochaetidae Scale insects Internal
Hymenoptera (Ants, Chalcididae Flies and butterflies (larvae and pupae) Internal/external
bees and wasps) Encyrtidae Aphids, scales, mealybugs, whiteflies Internal
Eulophidae Aphids, gall midges, sawflies and mealybugs Internal/external
Pteromalidae Flies including houseflies and stable flies Internal
Pteromalida Boll weevils Internal
Aphelinidae Whiteflies, scales, mealybugs, aphids Internal/external
Trichogrammatidae Moth eggs Internal
Mymaridae True bugs, flies, beatles, leafhopper eggs Internal
Scelionidae Eggs of true bags and moths Internal
Ichneumonidae Larvae or pupae of beetles, caterpillar, wasps Internal/external
Braconidae Larvae of beetles, caterpillars, sawflies Internal (mostly)
DIPTERAN FLIES
• The tachinid Compsilura concinnata, for example, has
been successfully reared from more than 100 host
species and three different host orders. Members of
other Diptera families — such as big-headed flies in
the Pipunculidae family, which are endoparasites of
leafhoppers and planthoppers, and the small-headed
Acroceridae, which only target spiders — are
generally more specialized. However, some attack
hosts from several families or subfamilies.
Major Groups of Dipteran (Fly) Parasitoids

Family Described Species # Primary Hosts/Prey: Gastropods

(Sciomyzidae) (100) (snails/slugs)
 Major Groups of Dipteran (Fly) Parasitoids

Family:Nemestrinidae
Primary Hosts/Prey: Acrididae
Described Species #: 300
 Major Groups of Dipteran (Fly) Parasitoids

Family:Bombiliidae Primarily Hymenoptera,

Described Species #: 5000 Coleoptera, Diptera
 Major Groups of Dipteran (Fly) Parasitoids

Familiy:Pipunculidae Primary Hosts/Prey:

Described Species #: 1000 Homoptera: Auchenorrhyncha
Major Groups of Dipteran (Fly) Parasitoids

Family: Conopidae Primary Hosts/Prey: Hymenoptera:

Described Specie #: 800 Adults of bumblebees & wasps
 Primary Hosts/Prey: Lepidopetra,
Family{ Sarcophagidae Orthoptera, Homoptera, Coleoptera,
Desribed Species #: 750 Gastropoda + others
Tachinidae Lepidoptera, Orthopetra, Homoptera,
8200+ Coleoptera, Hemiptera, Diptera + many others
Pyrgotidae
350 Coleoptera:Scarabaeidae
Acroceridae
500 Arachnida:Aranae
 Hymenoptera, Diptera, Coloeptera,
Phoridae Lepidoptera, Isoptera, Diplopoda +
300 others
Rhinophoridae
90 Isopoda
Calliphoridae Earthworms,Gastropods (Snails)
 CHALCID WASPS

 They are often underestimated in their nos. and

effectiveness.
 They can be seen tapping leaf surfaces with their
antennae in search of their host’s scents but their
presence is most commonly revealed by the
sickly /dead hosts they leave in their wake.
 The parasitize a great nos. of pests, & different
species attack different stages of the same host.
 The following six families have proven especially
useful in managing pests.
 FAIRYFLIES (Mymaridae)

 Between 0.008 & 0.04 inches (0.2–1 mm) long, these

smallest of world’s insects can fly through the eye of
needle
 Back of the wings contain distinctive long hairs.
 They parasitize the eggs of other insects — commonly
flies, beetles, booklice and leafhoppers.
 Genus Anaphes, play crucial roles in biological control. The
introduced egg parasite A. flavipes, for example, is one of
two parasites that have been established for cereal leaf
beetle management in small grains. In pesticide-free
California vineyards with ground vegetation, the tiny
Anagrus epos wasp can make a big dent in grape leafhopper
densities
 Trichogramma wasps (Trichogrammatidae)

• Most widely released natural enemies.

• Tiny female wasps generally less than 0.04 inch
(1 mm) long — lays an egg inside a recently laid
host egg, which blackens as the larva develops.
• Host range of many Trichogramma wasps spans
numerous species and families of insects.
• Moths, butterflies, beetles, flies, wasps and true
bugs are all frequent victims.
• Some Trichogramma wasps even use their wings in
a rowing motion to reach aquatic hosts
 Eulophid wasps (Eulophidae)
 More than 600 in North America, making theirs one of the
largest chalcid families
 About 0.04 to 0.12 inches (1–3 mm) long, they are often
brilliant metallic blue or green
 Some species of eulophids are mite predators while others
attack spider egg cases, scale insects and thrips. Most
eulophids, however, parasitize flies, other wasps or the
larvae or pupae of beetles or moths. Leaf-mining and wood-
boring insects are frequent hosts. Eulophids destroy many
major crop pests.
 In the Midwest alone, Sympiesis marylandensis is an
important parasite of spotted tentiform leafminer in apples.
Diglyphus isaea — available commercially — is a primary
parasite of agromyzid leafminers in greenhouses. Edovum
puttleri attacks the eggs of Colorado potato beetles.
Finally, Pediobius foveolatus — introduced from India and
also available commercially — parasitizes Mexican bean
beetle larvae. Pteromalid wasps (Pteromalidae).
 Eulophid wasps (Eulophidae)
 This large family of wasps assaults many types of
insects, including the larvae of moths, flies, beetles
and wasps. Several pteromalids target scale insects
and mealybugs and some even act as
“hyperparasitoids” — parasitizing other parasites
within their hosts.
 In the upper Midwest, Pteromalus puparum is a key
enemy of imported cabbageworm pupae, each of
which can involuntarily host more than 200
Pteromalus offspring.
 Anisopteromalus calandrae, which attacks the larvae
of beetles that infest stored grain, impressed
scientists several decades ago with its ability to
suppress 96 percent of rice weevils in wheat spillage
in small rooms. A. calandrae can now be purchased
for release in grain storage and handling facilities.
 Encyrtid wasps (Encyrtidae)
 Responsible for much of the classical biological control of scale
insects and mealybugs in fruit trees, this important family of
natural enemies encompasses about 400 species in the United
States and Canada
 Its extensive host range includes soft scales, armored scales,
mealybugs and the eggs or larvae of insects in about 15 families
of beetles, 10 families of flies and 20 families of moths and
butterflies.
 Several commercially available encyrtids now help manage scale
and mealybugs in greenhouses: Leptomastix dactylopii, for
example, parasitizes citrus mealybug, while Metaphycus
helvolus attacks black, hemispherical, Nigra citricola, brown
soft and other soft scales. Other noteworthy encyrtids include
Ooencyrtus kuwanae, an introduced parasite of gypsy moth eggs,
and Copidosoma floridanum, a native parasite of cabbage looper
larvae.
 Aphelinid wasps (Aphelinidae)

 The effectiveness of aphelinids in managing

scale insects has earned them one of the
best reputations in biological control. They
also destroy mealybugs, whiteflies, aphids
and other families of Homoptera. Aphelinus
varipes parasitizes greenbugs, A. mali
targets the woolly apple aphid, and
members of the genus Eretmocerus attack
silverleaf whitefly. Encar sia formosa, in
commercial use since the 1920s, is now
released into greenhouses worldwide; it kills
almost 100 greenhouse whitefly nymphs
during its 12-day life span.