Lecture Notes in CROPPROT105 - Beneficial Arthropods and Micro-organism

Introduction to Beneficial
Arthropods and Microorganisms

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Lecture Notes in CROPPROT105 - Beneficial Arthropods and Micro-organism

The biological world is heterogenous and offers numerous differing habitats.

Habitat refers to area that can be used by a particular organism to support survival and/or
reproduction. It can be studied at different spatial scales, ranging from microhabitats which
contain factors an individual requires to conduct a particular activity (e.g. foraging or
reproduction), to macrohabitats which contain all the factors required to support a
population of a given organism (Stamps, 2008). An organism, however, will select and
occupy those areas that fall within certain tolerance limits with the maximal population near
the optimal preferences. These habitats include aquatic environment and terrestrial
environment.

The maintenance of the population dynamics equilibrium within particular upper

and lower limits over a period of a complex combination of the environment factors affecting
upon the population is called natural control.

The rebounding of the population in returning to the characteristic mean density

after periods of positive or negative excess is most important. Consequence to this result to a
particular species continued to increase or decrease in number. This is called Balance of
Nature.

The Balance of Nature is the result of natural regulative processes in the

environment of every living organism and it assures that a species will neither decline in
numbers to extinction nor increase to infinite density. Regardless of whether a species is
abundant or scarce, the average characteristic density of its population in a given habitat is
constant. Any given species in a community with few exceptions is attacked and fed on by
one or more such natural enemies, and indication of the tremendous potential for biological
control.

All organisms are capable of increasing in numbers through process of reproduction.

Most insects are notable for their high potential rate of numerical increase because of their
relatively very high fecundities and short life cycles. But the fact is that most organisms
including rapidly breeding insects do not increase over successive generations or for
prolonged period. On the contrary, they increase only periodically and to a limited extent, a
consequence of Natural controls presents in their environments (Solomon, 1949).

The natural control generally limits the number of insects. Such checks to
numerical growth include limited resources (Food, Space, and Shelter) periodically occurring
inclement (severe) weather or other hazards (Heat, Cold, Wind, Rain, and Drought),
competition from themselves (intraspecific competition) or from other kind of animals
(interspecific competition) and natural enemies (Predator, parasites and pathogens).

Two Major Components of Natural Control of Population Numbers

1. Density - independent factors – the effect on the proportion of organisms affected
by this factor is constant for any population density, or the factor does not depend on
population density. In other words, these affect all individuals equally, regardless of
the overall population density.
 Physical (e.g. Temperature, Humidity, Air movement, Exposure to heat)
 Biological (e.g. Host Suitability, Food Quality)
2. Density - dependent factors –the effect on the proportion of organisms affected
varies with density, or the effect of the factor depends on population density
 Non - reciprocal (e.g. Some Foods, Space territoriality)
 Reciprocal (Parasites, predators, pathogen, herbivores, some food sources

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1.2 Review of Ecological Concepts and Principles – Basis of Biological

Control
Defining Ecology

The term “ecology” comes from the two Greek words, “oikos” meaning house or
household, and “logos”, which refers to “the study of” some particular topic. Literally
translated then, ecology means the study of households, in this case, the households of
nature. German zoologist Ernst Haeckel, who is credited with coining the word in 1870,
defined it as “the body of knowledge concerning the economy of nature – the investigation of
the total relations of the animal both to its inorganic and its organic environment.”
“Ecology is the study of the relationships of organisms to their environment and to
one another” (Brewer, 1994).
“Ecology is the scientific study of the interactions that determine the distribution and
abundance of organisms” (Krebs, 1972).
“Ecology is the study of the structure and function of ecosystems” (Odum, 1963).
“Ecology is a scientific natural history” (English ecologist Charles Elton, 1927).
Natural history – the observations and descriptions of the behavior and adaptations of
organisms.

The transition to “scientific” natural history was Darwin’s theory of evolution.

Specifically, his concept of natural selection provided a mechanism to explain how
populations of organisms change, adapt, evolve, to an ever-changing environment.

1. Everything is connected.
In addition to the definitions
2. Everything must go somewhere.
of ecology, we have here Four
3. Nature knows best.
Laws of Ecology according to
4. There is no such thing as a free lunch.
Barry Commoner (1971)

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Lecture Notes in CROPPROT105 - Beneficial Arthropods and Micro-organism

Ecological basis of biological control

• Biological control is a manifestation of the association of different interdependent species

in nature.
• Ecological concepts such as nature of population and communities, balance of nature and
natural control of numbers are important basis of biological control.
• When species exist as a group of like individuals which interbreed, reproduce and die, and
occupying a definite area, it is called a population.
• An assemblage of different population in a given area constitutes a community.
• In communities, trophic or nutritional interactions are distinguished.

Types of Trophic Interaction

1. Phytophagy (herbivory)– consumption of plants by herbivorous animal

2. Carnivory – consumption of animals by another animal
3. Entomophagy – consumption of insects by another insect.

Trophic Relationships
The most obvious functional relationship linking plants and animals together in any
ecosystem was food based. Feeding or trophic relationships delineated who ate whom in
order to obtain the energy and nutrients necessary for survival. Hence, any community of
organisms could be organized on the basis of the following trophic levels:

1. Producers – Those organisms capable of producing their own food, primarily by

fixing energy from the sun via photosynthesis. These autotrophs (self-feeders), most
of which are plants, then serve as the primary energy source for the rest of the
biosphere.
2. Herbivores (primary consumers) – Those organisms obtaining their energy
directly from plants.
3. Primary Carnivores (secondary consumers) – Those organisms obtaining their
energy from herbivores.
4. Secondary Carnivores (tertiary consumers) – Those organisms obtaining their
energy from other carnivores. While one could conceivably continue “stacking up”
carnivores in this fashion indefinitely, most ecosystems rarely exceed 4 or 5 trophic
levels.
5. Decomposers or Detritivores - These organisms obtain their food from dead
plants and animals. Through their actions, the building blocks of life are returned to
the environment in elemental form to be used again. While often not considered a
distinct trophic level, they are indispensable members of the biotic community.

• Food chains are usually observed when a given plant is fed upon by defoliating insects,
which in turn is fed upon by bird species. Below are examples of food chains.

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Lecture Notes in CROPPROT105 - Beneficial Arthropods and Micro-organism

(pinterest.com) (byjus.com)

• When food chains are linked together in complex communities, this is called food web.

• Species interacting with other species form a distinct community. Below are examples of
food webs.

(projects.ncsu.edu) (igbiology.blogspot.com)

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Ecological Relationships

Species Interactions
An organism’s trophic relationships, and its relative importance in the flow of energy
and the cycling of nutrients, are important aspects of its role, or niche in the community.
Another key element in describing an organism’s niche is the way it interacts with other
species within its community. What kinds of interactions exist among species? One way of
answering this question is to determine the effect that one species has on another’s ability to
survive and reproduce. In the table below, types of interactions are listed along with their
effects on the two species involved. In a predator-prey interaction, for instance a plus
indicates that species one (the predator) benefits from the interaction. For species two (the
prey), the negative sign signifies a negative impact on its population. Note that this
particular type of interaction would also include herbivores eating plants and host/parasite
interactions.

Name Description Effect

Neutralism The state of being neutral 0/0
Competition Organisms of two species use the same limited resource -/-
and have a negative impact on each other.
Amensalism A symbiotic relationship between organisms in which one 0/-
species is harmed or inhibited and the other species is
unaffected.
Predation A member of one species, predator, eats all or part of the +/-
body of a member of another species, prey.
Herbivory A special case of predation in which the prey species is a +/-
plant.
Mutualism A long-term, close association between two species in +/+
which both partners benefit. Example: pollination of
plants by honeybees.
Commensalism A long-term, close association between two species in +/0
which one benefits and the other is unaffected. Example:
mites attach themselves to larger flying insects for
transportation.
Protocooperation The first in time association of organisms working +/+
together for common benefit.
Parasitism A long-term, close association between two species in +/-
which one benefits and the other is harmed. For example,
roundworms are parasites of mammals, including
humans, cats, and dogs.

Competition is defined as the use of a limited resource by two or more individuals,

either of the same species (intraspecific competition) or different species (interspecific
competition). Competition is negative for both because use or defense of a resource by one
(individual or species) always reduces availability of that resource for any other.
For much of the past 130 years, most ecologists believed that the old dictum “nature,
red in tooth and claw” succinctly described the dominant forces shaping and controlling the
natural world. We discovered that the concepts of trophic interaction and energy flow were
pivotal in developing an ecological framework. They also influenced thinking about the way
in which biological communities were structured. Predation and competition, (killing and
fighting for resources) were seen as the key to understanding how communities were
organized. They were also viewed as important determinants of population size and stability
of natural systems. As one famous study put it, “Are populations limited primarily by what

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they eat or by what eats them?” (Hairston, Smith and Slobodkin, 1960). As a result, the
ecological literature is especially rich in articles detailing predator-prey interactions and
competition. Over the years, observation, theory and experiments have clearly demonstrated
that both do play important roles in structuring biological communities. More recently,
ecologists have turned their attention to other types of interactions. Within the last several
decades, they have begun to elucidate the importance of win-win interactions, such as
symbiotic relationships among plants and their pollinators and the mycorrhizal association
of fungi with plant roots, to similarly shape community structure.

Species-Community Relationships
Interspecific (between species) interactions often have a significant effect on the
number of species present in a community and their relative abundance. Evolutionary
ecologists believe, for example, that much of the great diversity in life we see has come about
through competitive exclusion. The Competitive Exclusion Principle states that two
or more species cannot coexist on a single limited resource. Competition thus
leads to one of two scenarios. Either one species will “out compete” the other(s) and gain sole
possession of that resource or natural selection will, over time, select for those individuals
that exploit different resources, thereby avoiding competition. Ultimately, characteristics of
species diverge sufficiently to allow for coexistence with each species occupying a unique
niche in the community.

In some habitats, ecologists have identified keystone species, species whose

addition or removal may lead to major changes in community structure.

Think of examples in the Philippines of effects of removing a keystone species. What

happened?

Population Dynamics
As illustrated in the preceding section, species interactions can have profound effects
on the numbers of individuals in a given population. Obviously, many other environmental
factors, both abiotic and biotic, affect population size. Ultimately, we can track changes in
population density, the number of individuals per unit area, as a resultant of four factors:
 Natality – the production of new individuals through either sexual or asexual
reproduction
 Mortality – loss of individuals through death
 Immigration – new individuals moving into a population
 Emigration – residents moving out of a population. All the fancier models of
population growth (which we happily won’t go into) are based on this simple equation:

N(t+1) = Nt + B + I – D – E

In part, the job of population biologists and wildlife managers involves assigning
values to each of these four variables to better predict future population trends. Intrinsic
(internal) factors affecting population growth include its sex and age distribution,
age-specific fecundity (rate at which an individual produces offspring), and social
structure. A population comprised mostly of young, pre-reproductive individuals is going
to have a different growth pattern than one that has a high percentage of older, post-
reproductive individuals. Certain populations, such as some species of salamanders, need a
critical minimal number of individuals in order for successful breeding to occur. Territorial
species, such as many of our songbirds, will behaviorally limit the number of breeding pairs
allowed in a given habitat.

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There are also many extrinsic (external) factors affecting population growth,
including competition, predation, disease, pollution, hunting, and carrying
capacity of the environment.

Carrying Capacity
Carrying capacity is a very important ecological concept. It is defined as the
maximum number of individuals of a given species that a habitat can sustain
indefinitely. When habitat quality improves, its carrying capacity increases. If habitat
declines so does the carrying capacity. This is why both the quantity and quality of wildlife
habitat is so critical to maintaining wildlife populations and why wildlife managers “manage”
habitat, not wildlife.

Two growth forms: Exponential (first graph) and logistic (second graph)
Courtesy of E.O. Wilson and W.H. Bossert, 1971.

Population Growth Models

For many species (most invertebrates), populations may exhibit exponential

growth (first graph). After starting slowly, numbers begin to accelerate rapidly, increasing
at an ever-increasing rate, mimicking the way money grows in an account earning compound
interest. Populations often continue to grow exponentially until a sudden change in
environmental conditions causes them to “crash.” An insect population growing
exponentially throughout the spring and summer may be brought to a sudden halt by the
first cold snap. This is density-independent growth; the growth rate of the population is
independent of the population density. Other species, especially long-lived vertebrates, may
exhibit a logistic growth pattern, as idealized in the second graph. Their populations show
the effects of increasing environmental resistance. The greater the population size, the more
the environment “pushes” against further growth. As numbers of individuals approach the
habitat’s carrying capacity (K), the population growth rate gradually slows until, at K, it
becomes zero, thus stabilizing the population at carrying capacity. This density-dependent
growth is the result of both intrinsic factors (greater social stress leads to lower natality rates,
lower survivorship rates, higher emigration rates, etc.) and extrinsic factors (increased
predation, disease).
No species in nature follows either pattern exactly or indefinitely. The logistic growth
model, for example, assumes that populations are capable of immediately changing their
growth rates in response to environmental resistance. The reality is that there is usually a lag
effect, or time delay. A population at carrying capacity may continue to grow for some time
before environmental factors leading to zero population growth take effect. Thus, rather than
leveling out at K, a population may overshoot it, leading to over-exploitation of resources and

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an eventual decline below the original carrying capacity. The long-term result may either be
a population that oscillates around K in an increasing tighter pattern (damped oscillation) or
one that exhibits more or less regular periodic cycles of abundance. Periodic fluctuations in
numbers have often been documented in northern habitats such as the classic 10-year cycle
exhibited by snowshoe hare, lynx, ptarmigan and ruffed grouse and the 4-year cycles
exhibited by various species of voles and lemmings.

1.3. Review of Basic Definition of Pests and their Characteristics

Concept of Pest
Pest is derived from French word ‘Peste’ and Latin term ‘Pestis’ meaning plague or
contagious disease. The following are some of the definitions of pests:
 Pest is any animal which is noxious, destructive or troublesome to man or his interests
 A pest is any organism which occurs in large numbers and conflict with man’s welfare,
convenience and profit
 A pest is an organism which harms man or his property significantly or is likely to do
so (Woods, 1976)
 Insects are pests when they are sufficiently numerous to cause economic damage
(Debacli, 1964)
 Pests are organisms which impose burdens on human population by causing
- injury to crop plants, forests and ornamentals
- annoyance, injury and death to humans and domesticated animals
- destruction or value depreciation of stored products.
- pests include insects, nematodes, mites, snails, slugs, etc. and vertebrates like rats,
birds, etc.

Categories of Pests
The pests are categorized based on the following:

1. Based on occurrence
 Regular pest – pest that frequently occurs and with close association on crop. e.g.
Rice stem borer, eggplant fruit borer
 Occasional pest – pest that infrequently occurs and with no close association on
crop. e.g. Caseworm on rice, Mango stem borer
 Seasonal pest – pest that occurs during a particular season every year. e.g. Red hairy
caterpillar on peanut, Mango hoppers
 Persistent pest – pest that occurs on the crop throughout the year and is difficult to
control e.g. Chili thrips, mealybugs on guava
 Sporadic pest – pest that occurs in isolated localities during some period. e.g.
Coconut slug caterpillar

2. Based on level of infestation

 Epidemic pest - sudden outbreak of a pest in a severe form in a region at a particular
time e.g. Corn planthopper (Stenocranus pacificus) that infested corn in Mindanao
 Endemic pest - occurrence of the pest in a low level in few pockets, regularly and
confined to particular area e.g. Aphids in sitao that are confined in shoots and young
pods

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3. Based on importance
 Agricultural pests – pests that infest crops and stored agricultural products. E.g.
insect pests such as rice black bugs, rice bugs, eggplant fruit borers, weevils, and many
more; golden apple snail; rats; rice birds (maya); mites
 Forest pests – pests that infest forest trees. E.g. long-horned beetles, mites
 Household pests – pests found in our homes that attack furniture, appliances,
buildings and clothing such as termites, cockroach, powder post beetles, cloth moths,
rats
 Medical pests – pests that infest humans such as lice, mosquitoes, bedbugs, fleas,
black flies
 Aesthetic pests -pests that infest museum collections, cut flowers and lawns
 Veterinary pests – pests that infest domestic animals such as mites, ticks, deer flies,
horse flies

Parameters of insect population levels

1. General equilibrium position (GEP) - The average density of a population over a
long period of time, around which the pest population over a long period of time
tends to fluctuate due to biotic and abiotic factors and in the absence of
permanent environmental changes.
2. Economic threshold level (ETL) - Population density at which control measure
should be implemented to prevent an increasing pest population from reaching
the ETL.
3. Economic injury level (EIL) - The lowest population density that will cause
economic damage
4. Damage boundary (DB) - The lowest level of damage which can be measured.
ETL is always less than EIL. Provides sufficient time for control measures.

Pest Categories According to EIL, GEP AND DB

1. Key pests
 Most severe and damaging pests
 GEP lies above EIL always
 Spray temporarily bring population below EIL
 These are persistent pests
 The environment must be changed to bring GEP below EIL. e.g. Cotton bollworm,
Diamondback moth
2. Major pests
 GEP lies very close to EIL or coincides with EIL
 Economic damage can be prevented by timely and repeated sprays. e.g. Cotton jassid,
Rice stem borer
3. Minor pests/Occasional pests
 GEP is below the EIL usually
 Rarely they cross EIL
 Can be controlled by spraying e.g. Cotton stainers, Rice hispa, Ash weevils
4. Sporadic pests
 GEP generally below EIL
 Sometimes it crosses EIL and cause severe loss in some places/periods. e.g. White
grub, Hairy caterpillar
5. Potential pests
 They are not pests at present.
 GEP always less than EIL.
 If environment changed may cause economic loss.

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Causes of Pest Outbreak

Activity of human beings which upsets the biotic balance of ecosystem is the prime
cause for pest outbreak. The following are some human interventions that cause pest
outbreak:
1. Deforestation and bringing under cultivation
 Pest feeding on forest trees are forced to feed on cropped
 Biomass/unit area more in forests than agricultural land
 Weather factors also altered - Affects insect development
2. Destruction of natural enemies
 Due to excess use of insecticides, natural enemies are killed
 This affects the natural control mechanism and pest outbreak occurs, e.g. Synthetic
pyrethroid insecticides kill natural enemies
3. Intensive and extensive cultivation
 Intensive monoculture leads to multiplication of pests
 Extensive cultivation of susceptible variety in large area
- No competition for food
- multiplication increases e.g. Stemborers in rice and sugarcane
4. Introduction of new varieties and crops.
 Varieties with favorable physiological and morphological factors cause
multiplication of insects. e.g. Succulent, dwarf rice varieties favor leaffolder
5. Improved agronomic practices
 Increased Nitrogen fertilizer
 Closed planting of rice cause high leaffolder incidence
 Brown planthopper (BPH) and leaffolder increases granular insecticides
 Possess phytotoxic effect on rice
6. Introduction of new pest in new environment
 Pest multiplies due to absence of natural enemies in new area
 Apple wooly aphid Eriosoma lanigerum multiplied fast due to absence of Aphelinus
mali (Parasitoid)
7. Accidental introduction of pests from foreign countries (through air/sea ports). Some
examples of pests accidentally introduced in the Philippines are as follows:

Gypsy moth – Brown marmorated stink bugs- Potato cyst nematode -

Lymantria dispar Halyomorpha halys Globodera pallida
(groweg.com) (extensionunh.edu) (potatopro.com)

Argentine ants – Coconut mealybug - Coconut scale insect-

Linepithema humile Nipaecoccus nipae Aspidiotus rigidus
(antark.net) (entnemdept.ufl.edu) (pestnet.org)

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Mango pulp weevil-

Coconut leaf beetle – African giant snail –
Sternochetus frigidus
Brontispa longissima Achatina fulica
(topbest.ph)
(coconut protectors.com) (shutterstock.com)

8. Large scale storage of food grains

 Serve as reservoir for stored grain pests (such as weevils)
 Urbanization - changes ecological balance
 Rats found in underground drainage

Resurgence
Resurgence is the tremendous increase in pest population brought about by
insecticides despite good initial reduction in pest population at the time of treatment. Below
are some examples of insecticides and the insect pests that resurged.
 Deltamethrin, Quinalphos, Phorate - resurgence of brown planthopper (BPH) in
rice
 Synthetic pyrethroids – resurgence of whitefly in cotton
 Carbofuran – resurgence of leaffolder in rice

Losses caused by pests

Crop loss from all factors is equivalent to 500 billion US $ annually worldwide. The
crop loss according to the organism is shown below.
 Insect pests - 15.6% loss of production
 Plant pathogens - 13.3%
 Weeds - 13.2%

1.4. Factors that Determine the Existence of an Organism in an

Ecosystem

Key Regulating Factors

1. Abiotic Factor
A natural abiotic control is any condition of the environment that inhibits the
growth of organism populations and cannot be altered by man.

a. Weather
 Temperature. Temperature affect the activity of an organism. In insects, for
example, warmer weather conditions increase its metabolic rate, while cooler
temperatures slow it. The insect’s activity also generally parallels temperature
levels, the result being that in many species the individual may consume greater
amount of food, reproduce more and may experience more generations in a given
warm season. In addition, the insect’s ability to fly may sometimes be diminished by
lower temperatures; consequently, the distribution of species is reduced, as are the
opportunities to find both food sources and mates. Sometimes, extremely low
temperatures may freeze and kill insects.

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 Moisture. Dry weather may drastically affect populations by restricting the insect’s
ability to retain water, a critical consideration to survival. Desiccation may either kill an
insect by reducing its metabolic activity, render it more susceptible to disease or affect
its behavior.
 Winds. Winds may affect populations either by helping them to move to new food
sources and breeding areas, or by driving them away. In addition, wind increase the
evaporation rate and may be a significant contributor to insect desiccation.
 Sunlight. For many species there a specific cutoff daylength under which the insect
may experience the onset of diapause (a slowdown of metabolism and
development). Other light-related reactions include polymorphism (the existence
of a number of distinct insect forms at a given stage), lower fecundity (or
reproduction potential) and certain aspects of behavior (e.g. oviposition).
 Extreme weather such as hurricanes, torrential downpours and droughts
increases the risk for species extinction, especially in biodiverse ecosystems such as
coral reefs and tropical rainforests.

b. Climate
 Changing climate affects ecosystems in a variety of ways. For instance, warming
may force species to migrate to higher latitudes or higher elevations where
temperatures are more conducive to their survival. Similarly, as sea level rises,
saltwater intrusion into a freshwater system may force some key species to relocate
or die, thus removing predators or prey that are critical in the existing food chain.
 Projected warming could greatly increase the rate of species extinctions, especially
in sensitive regions.

c. Shelter availability
 An organism’s shelter protects it from predators and weather.
 Shelter also provides a space for eating, sleeping, hunting, and raising a family.
 Shelters come in many forms. A single tree, for example, can provide sheltered
habitats for many different organisms. For a caterpillar, shelter might be the
underside of a leaf. For a mushroom fungus, shelter might be the cool, damp area
near tree roots. For a bald eagle, shelter may be a high perch to make a nest and
watch for food.
 Unavailability of shelter will result in competition for space and the possible
extinction of an organism

d. Geographic barriers or topography

 Geographic barriers such as deserts, mountains, oceans, lakes and rivers may
prevent some species to move from place to place in search of food, mate or
escaping from enemies
 These obstacles directly prevent species from dispersal, especially in the cases of
oceans for terrestrial species, land masses for marine species, and transitions
between freshwater and marine systems for many aquatic organisms.

2. Biotic factors
The living components which affect the survival of organisms such as insects in the
environment are called biotic. These include food, competitors, and natural enemies
(Wilson et al., 1984).
a. Food
Plant feeding insects in the community are called phytophagous, whereas
those species that feed on other insects are called entomophagous. In general,
phytophagous insects are considered injurious and entomophagous insects, beneficial.

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Insects may be further classified by their feeding habits. Polyphagous insects feed
on a wide variety of plant or animal species. Oligophagous insects are somewhat
restricted in their eating habits, usually feeding on genera within a plant family.
Monophagous insects usually restrict their feeding to only closely related species
within a single genus. For example, the armyworm which feeds only on species within
the grass family including grasses, wheat, oats, barley and corn is oligophagous,
whereas the European corn borer is polyphagous, feeding not only on grasses, but also
on legumes, solanaceous plants and many other families. The spotted alfalfa aphid, on
the other hand, is monophagous, feeding only on alfalfa or closely related species.
These feeding habits may favor those polyphagous compared to monophagous when
there is food shortage.

b. Competitors
Competition, a major regulating factor, occurs between individuals of the same
species (intra-specific competition) as well as between species (inter-specific
competition). Competition between individuals for food can become very intense as the
available food supply diminishes. This adds stress to community as a whole. For
example, a given food supply may be sufficient to support a population of insects to
complete their development, but an increase in the population may exhaust the food
supply and destroy the population before any individuals reach maturity. However, the
latter rarely occurs because of biological variability between individuals which allows
some to feed faster, or develop faster and survive at the expense of the slower
individuals. This intraspecific competition thus brings about the selection of better
adapted individuals. There can also be competition for mates, space and habitat, all of
which can limit the development of insect population.
When different species compete (interspecific competition) with each other for
food and space a small species which needs less food to complete its development,
usually has an advantage over a larger species. Likewise, a species which completes its
development in a short time usually will have an advantage over a species requiring a
longer period of time to complete development. A polyphagous species has an
advantage over a monophagous species because it may be able to move to a different
food source.

c. Natural enemies
Other insects and animals which are natural enemies of an insect species
contribute to the regulation of its numbers. However, through natural selection a
balance has usually been developed. Predators do not always get their prey, nor do
parasites find the last host. It has been found that the reaction time of many predators
and their prey is about the same so that half of the time the predator gets its prey and
about half the time the prey escapes. Likewise, a parasite has difficulty locating the last
few individuals, and may itself be reduced to low numbers through starvation. Thus,
host species are able to rebuild their populations. This preserve both predator and prey
species, and is important if a biological control program is to have lasting effectiveness.
Extreme efficiency on the part of predators and parasites, eliminating all prey and
hosts would eventually cause the predators and parasites to perish, too.

Population Increase in Absence of Regulatory Factors

a. Exponential growth (also known as a J-curve) - occurs when there is no limit to

population size. Ex. Houseflies. Assuming houseflies have an average generation time
from egg to adult of about 10 days and each female can lay about 120 eggs, the
resulting offspring in just three months from this single female would number more

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than 325 trillion individuals. If we were to line up all of the resulting individuals
lengthwise, the resulting line of flies (assuming an average housefly length of 7mm)
would encircle the equator 757,000 times. This type of unchecked population increase
is known as exponential growth as shown below.

Population growth showing exponential curve

b. Logistic Curve (also known as an S-shaped curve) - shows the effect of a limiting
factor (in this case the carrying capacity of the environment). Population growth is
slow initially then it enters into a point where growth is rising rapidly.

Eventually the population stops increasing and reaches its maximum level or
“carrying capacity”. The maximum population size that can be reached is based on the
availability of light, in the case of plants, or food, shelter, etc. Most population never
approach the “carrying capacity” but instead remain at lower levels because of the
regulating effects of both abiotic and biotic factors.

Note that populations do not typically remain at a steady state continually but
instead tend to fluctuate or oscillate around some characteristic density.

Population growth showing logistic curve

Factors Governing Populations at Maximum and Minimum

The major factors that regulate populations act differently with regards to how it
exerts control over a population. For example, biotic factors interacting within a
population (i.e. intra-specific competition) work together to maintain populations
below the “carrying capacity”.
When populations become too large, the individuals of the same species begin to
compete for the same resources such as food, shelter, egg-laying sites, etc. This
interaction between members of the same species tends to be the most important
factor maintaining population below the “carrying capacity”.

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Conversely, harsh abiotic factors such as hurricanes, unusual freezing conditions,

high temperatures, shifts in climate, and change in sheltering conditions can all act to
reduce populations to very low or maximal levels, levels where the populations may be
eradicated from a particular area.

Factors Governing Populations Below Carrying Capacity

The regulations of populations below the carrying capacity is influence by the

biological control agents (parasites, pathogens, and predators) as well as competition
between species with similar environmental requirements (i.e. inter-specific
competition). While other factors, most notably abiotic factors, may influence the
fluctuations.

Processes Responsible for Causing Variation in Numbers (Malthus and

Darwin)
1) Limited food supply
2) Disease
3) Competition
Two basic elements which influence the rate of the three variables:
Birth rate, dispersal rates and death rate (Clark et al. 1967)

1) Species characteristics
 Vigor – vigorous or healthy species can produce healthy offspring,
can move from place to place in search for food, mate or shelter or
escape from enemies. This may result to high birth rate, high
dispersal rate and low death rate.
 Fecundity – fecund species can produce more offspring. This result to
high birth rate.
 Behavior – cannibalistic behavior, for example, a mother eats her
eggs or offspring. This results to low birth rate.
 Competitive ability – competitively superior species drives the other
species to extinction or migration.
 Genetic factors – long-lived (K-organisms) such as forest trees or
short-lived organisms (r-organisms) such as agricultural crops, for
example may result to low death rate or high death rate.
2) Environmental influences
 Weather
 Natural disasters
 Pollution
Birth rate of a population is the ratio of births or oviposition during a short
period to the total number of individuals in the population at the beginning of
the period. This is determined primarily by innate characteristics of the
subject species with only slight input from environmental characteristics

Dispersal rates determined by a more or less coequal interplay by species

characteristics and environmental influences interacting upon individuals in
the population and causing some to emigrate and some to stay

Death rate is the ratio of the number of deaths during the given period to the
number of animals at the beginning. This is influenced most heavily by
environmental processes with modification by species characteristics.

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Three kinds of Mortality Processes (Howard and Fiske, 1911 as cited by

Coppel and Mertins, 2012) as observed in Gypsy moth or Brown tail moth, N.
phaerrhoea (Donovan):

a) Facultative agencies which destroy a larger proportion of the population as

abundance increases. Facultative agencies are called density –dependent such as
climatic conditions (Thompson, Uvarov, and Andrewartha and Birch).

b) Catastrophic agencies which are totally independent of rarity and abundance

in their effects. Catastrophic agencies are density –independent such as
competitors, parasitoids, predators and contagious diseases (Malthus, Howard
and Fiske, Nickelson, and Smith).

c) Agencies, including birds and other predators which act in a manner opposite
to the facultative agencies, destroying a certain gross number of individuals each
year, regardless of abundance, as a part of their diversified diet. Usually referred
to as inversely density-dependent – having little impact on the determination of
average population density (Smith, 1935)

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