Chapter - 1

INTRODUCTION
1.1: Morphology
Paper wasps are one of the most common wasps seen around homes and buildings. They
are 0.75 cm to 1 inch Long and generally reddish-orange to dark-brown. They often have
yellow body markings. Paper wasps have three castes, infertile female workers, which make
up most of the wasps on nests during the summer males, and queens. Males and new queens
are produced primarily in late summer and fall. Unlike yellow-jackets and hornets, the paper
wasp queen is not much larger than the worker wasps. Paper wasps build their nests from
chewed wood fibers. The comb, which hangs from a single filament, is usually oriented
downward and consists of a single tier of hexagonal-shaped cells. Nests are most frequently
seen under the eaves of houses but may also be found in attics, garages, storage sheds, barns,
on shrubbery, trees and a variety of protected sites. The typical mature paper wasp nest
contains 20 to 30 adults.
Anatomically there is a great deal of variation between different species of wasp. Like all
insects, wasps have a hard exoskeleton covering their 3 main body parts. These parts are
known as the head, metasoma and mesosoma. Wasps also have a connective region joining
the first and second segments of the mesosoma known as the petiole. Like all insects, wasps
have 3 sets of 2 legs. In addition to their compound eyes, wasps also have several simple eyes
known as ocelli. These are typically arranged in a triangular formation just forward of an area
of the head known as the vertex. It is possible to distinguish between certain wasp species
genders based on the number of divisions on their antennae. Male Yellow jacket wasps for
example have 13 divisions per antenna, while females have 12. Males can in some cases be
differentiated from females by virtue of the fact that the upper region of the male's mesosoma
(called the tergum) consists of an additional terga. The total number of terga is typically 6.
The difference between sterile female worker wasps and queens also varies between species
but generally the queen is noticeably larger than both males and other females. Wasps can be
differentiated from bees as bees have a flattened hind basitarsus. Unlike bees wasps generally
lack plumose hairs. They vary in the number and size of hairs they have between species.

1.1.1. Characteristics of Wasps:

The following characteristics are present in most wasps:
1. Two pairs of wings, An ovipositor or stinger (which is only present in females because it
derives from the ovipositor, a female sex organ).
2. Few or no hairs (in contrast to bees)
3. Nearly all wasps are terrestrial; only a few specialized parasitic groups are aquatic.
4. Predators or parasitoids, mostly on other terrestrial insects; some species of Pompilidae,
such as the tarantula hawk, specialize in using spiders as prey, and various parasitic wasps
use spiders or other arachnids as reproductive hosts.
5. Wasps are critically important in natural bio control. Almost every pest insect species has a
wasp species that is a predator or parasite upon it.
6. Parasitic wasps are also increasingly used in agricultural pest control as they have little
impact on crops. Wasps also constitute an important part of the food chain.

1.1.2. Feeding Habit of Wasps:

Generally wasps are parasites as larvae, and feed only on nectar as adults. Some wasps are
omnivorous but this is relatively uncommon, they feed on a variety of fallen fruit, nectar and
carrion. Many wasps are predatory, preying on other insects. Certain social wasp species,
such as yellow jackets, scavenge for dead insects to provide for their young. In turn the brood
provides sweet secretions for the adults.
In parasitic species the first meals are almost always provided from the animal the adult
wasp used as a host for its young. Adult male wasps sometimes visit flowers to obtain nectar
to feed on in much the same manner as honey bees. Occasionally, some species, such as
yellow jackets, invade honeybee nests and steal honey or brood.
Ropalidia marginata, the most common Indian social wasp, belongs to a crucial stage of
social evolution showing no obvious morphological caste differentiation but a behavioral
caste differentiation and a dominance hierarchy that appears to influence division of labour.
The nests consist of a single open comb that can sometimes have up to 500 cells and 10
2

pedicels. Nests are initiated and abandoned all round the year. Initiation is by 1-20
foundresses, 1-4 being the most common number. There is a great deal of variation in brood
development at times both within and between nests. Male progeny disappear from the nest
soon after emergence while daughters stay on at the parent nest for a mean period of about a
month. Small nests have a single egg layer while large nests have two or more females with
well developed ovaries that presumably lay eggs. Most nests are short-lived, small nests
being highly susceptible to failure. Large nests are less susceptible to failure but the
emergence of multiple egg layers reduces the average relatedness of workers to the brood
which presumably is the cause for large scale emigrations from these nests. An interaction of
ecological and social factors therefore appears to determine the growth of a nest .Yellow
paper wasps are beneficial insects. They feed their young numerous insects that ordinarily
damage shade trees and crops. They also kill countless houseflies and blow flies.
A few species of yellow-jackets, however, such as Vespula pensylvanica (western
yellowjacket), V. germanica (German wasp), and V. vulgaris (common yellow-jacket),
scavenge for meat and sweets and can become pests, especially at picnics and camp
grounds .Even though they may at times become pests, yellow-jackets and paper wasps are
highly beneficial. Do not control them unless their stings present a hazard.
Recent years have witnessed a great surge of interest in social hymenoptera because the
emergence of a considerable body of theoretical ideas (Hamilton 1964; Lin and Michener
1972 ; Alexander 1974) have raised hopes that herein lies the key to understanding the
evolution of social behaviour (West-Eberhard 1969,1975 ; Wilson 1971, 1975 ; Jeanne 1972,
1980 ; Michener 1974 ; Trivers and Hare 1976; Litte 1977, 1979, 1981 ; Starr 1979). Bees
and wasps are of special interest in this connection because they exemplify a series of stages
in the evolution of colonial work order. 539,540 Raghavendra Gadagkar et al sociality from
the completely solitary to the highly advanced eusocial species (see Evans and West-Eberhard
1970 ; Michener 1974 ; Wilson 1971).
Ropalidia marginata is the commonest social wasp of Peninsular India (Vander Vecht
1962). This species shows cooperative brood care, reproductive caste differentiation and
overlap of generations (Gadgil and Mahabal 1974 ; Gadagkar 1980 ; Gadagkar and Joshi
1982b, 1983a; Gadagkar unpublished observations) and hence can be called eusocial
according to the classification of Michener (1969).There is no obvious morphological
3

differentiation between egg layers and non egg layers (Gadgil and Mahabal 1974) and
division of labour is brought about by a dominance hierarchy among the females belonging to
a nest (Gadagkar 1980). Analysis of the time-activity budgets of adults on R. marginata nests
has in fact revealed the presence of a behavioral caste differentiation in this primitively
eusocial wasp (Gadagkar and Joshi 1982b, 1983a).Apart from these few recent studies there
is very little information in the literature about this interesting genus (Roubaud 1916 ; Carl
1934 ; Darchen 1976; Belvadi and Govindan 1981 ; Gadagkar and Joshi 1982a,c, 1983b).
Moreover, in addition to understanding reproductive differentiation and social organization,
information on the dynamics of initiation, growth and extinction of colonies is essential
before we even begin to speculate about the factors that might be responsible for the origin
and maintenance of sociality. We present in this paper the results of our observations on the
natural history as well as population ecology of Ropalidia marginata in Peninsular India.

1.2: Life Cycle

Colonies are founded in the spring by queens that spend the winter in sheltered hiding
places. Although early season queens may cooperate in founding a nest, by midsummer there
is usually only one active egg-laying queen per nest. New nests are constructed every year,
often in the same general location where nests were built the previous season Queen paper
wasps lay a single egg per nest cell. The newly hatched paper wasp larva is white and legless.
As it grows in size, it fills the nest cell. The queen, and later the workers, brings food to the
larva .Eventually, the larva matures, the cell is closed and pupation occurs. After pupation,
the adult wasp emerges by chewing through the paper cell cover. During late summer and
fall, males and queen paper wasps are produced. Males and females mate in the fall. The
males then die, and fertilized queens enter sheltered locations for hibernation.
Overwintering paper wasp queens may join together, sometimes in large groups. They
seem to prefer high structures, such as the peaked attics of houses, chimneys and tall
buildings. During the winter, they frequently enter homes and offices, especially during
warmer periods. Overwintering paper wasps are not aggressive and may be captured and
released outdoors or killed with a fly swatter. The yellow paper wasps have annual colonies.
The only colony members to overwinter are inseminated queens, which spend the winter in
protected locations, such as under bark, in stumps and logs, and within stacks of firewood.
The queens emerge during the first warm days of spring (as early as March and April), select
a nest site, and build a small paper nest in which they lay their eggs. When the eggs hatch, the
4

queen feeds the young larvae for up to 3weeks. Larvae then pupate, to emerge as smaller
infertile females called workers .Once the first five to seven workers appear, they begin
rearing and feeding the brood. The queen rarely is seen again outside the nest. The colony
then expands rapidly, and depending on the species, may consist of as many as 5,000 workers
and 15,000 cells in the nest for some yellow wasp species. Colonies attain maximum size in
August and September. Worker yellow wasps, then at their peak, become pestiferous. About
this time, new males and queens are produced from reproductive cells. These emerge and
mate. The males soon die, and the inseminated queens seek sheltered locations in which to
overwinter. The abandoned nests rapidly decompose and disintegrate during the winter. They
are not used again. In spring, the cycle starts over. The German wasp has become widespread
in Washington. The cycle of this yellow wasp is slightly later, with peak colony size
occurring in October and early November. This species has a propensity for nesting in
structures and has become a widespread urban pest. Recently, the European paper wasp
(Polistes dominulus) also has become widespread and abundant throughout Washington. It
readily nests on and within human-made structures and is particularly populous in urban and
suburban areas. The tendency of the German and European species to found nests in
association with people contributes to their pest status.

1.3: COLONIES OF Ropalidia marginata:

Colonies of the primitively eusocial wasp Ropalidia marginata consist of a single egg
layer (queen) and a number of nonegg-laying workers. Although the queen is a docile
individual, not at the top of the behavioral dominance hierarchy of the colony, she maintains
complete reproductive monopoly. If the queen is lost or removed, one and only one of the
workers [potential queen (PQ)] becomes hyper-aggressive and will become the next queen of
the colony. The PQ is almost never challenged because she first becomes hyper-aggressive
and then gradually loses her aggression, develops her ovaries, and starts laying eggs.
Although we are unable to identify the PQ when the queen is present, she appears to be a
cryptic heir designate. Here, we show that there is not just one heir designate but a long
reproductive queue and that PQs take over the role of egg-laying, successively, without overt
conflict, as the queen or previous PQs are removed. The dominance rank of an individual is
not a significant predictor of its position in the succession hierarchy. The age of an individual
is a significant predictor, but it is not a perfect predictor because PQs often bypass older
individuals to become successors. We suggest that such a pre-designated reproductive queue

that is implemented without overt conflict is adaptive in the tropics, where con-specific
usurpers from outside the colony, which can take advantage of the anarchy prevailing in a
queen-less colony and invade it, are likely to be present throughout the year. Reproductive
division of labor is the hallmark of insect societies. This is achieved by the differentiation of
colony members into reproductive (queen/king) and non-reproductive (worker) castes. In
insect societies such as those of ants, honeybees, swarm-founding wasps, and higher termites,
which are traditionally referred to as highly eusocial, caste determination is achieved by preimaginal physiological processes that channel individuals into distinct reproductive or worker
developmental pathways. Thus, caste of an individual is already determined at the time of
eclosion and remains irreversible. If the reproductive dies, the colony has to rear a new one
from the egg or early larval stage because the workers cannot change their caste in adulthood. In societies such as those of most social bees and wasps, which are traditionally
referred to as primitively eusocial, all adult colony members are nearly totipotent and
morphologically similar. The process of caste differentiation into reproductives and non
reproductives takes place largely in the adult stage and is based on social interactions among
colony members. This makes the castes flexible and often reversible, so that workers can
become queens upon the loss or death of the original queen. In addition to indirect fitness
gained as workers, individuals, thus, also have a finite probability of direct reproduction as
future queens. Similarly in wood-dwelling lower termites, workers that are still in their
nymphal stages can be totipotent and can develop into future reproductives. This is also
reminiscent of many cooperatively breeding birds and mammals in which adult helpers
temporarily forego reproduction and assist breeders to rear and protect their offspring. Such
helpers are also totipotent and can inherit breeding opportunities in the future. In most
primitively eusocial species, colony members display aggressive dominancesubordinate
interactions, on the basis of which they can be arranged in a linear dominance hierarchy. The
queens typically occupy the top (alpha) position in this dominance hierarchy and may be
sequentially replaced by individuals occupying successively lower positions in the dominance
hierarchy. Thus, there is usually a reproductive queue based on the behavioral dominance
hierarchy. In the primitively eusocial wasp Ropalidia marginata, on the other hand, queens
are remarkably meek and docile individuals that seldom, if ever, occupy the alpha position in
the aggression-based behavioral dominance hierarchy of their colonies, thus breaking the link
between behavioral dominance and reproductive dominance. Lost queens are replaced by one
of the workers, although this is not based on the position of the worker in the behavioraldominance hierarchy. Thus, it appears that dominancesubordinate interactions are not used
6

in this species to express or settle reproductive conflicts. Instead, these kinds of interactions
appeared to have been co-opted for use by intranidal workers to regulate foraging by
extranidal workers. Upon natural loss or experimental removal of the queen, one (and only
one) of the workers becomes hyper-aggressive, increasing her levels of dominance behavior
several fold relative to her own levels in the presence of the queen. If the queen is not
returned to the colony, this hyper-aggressive individual will become the next queen, losing
her dominance behavior by the time she begins to lay eggs in about a week. We, therefore,
refer to her as the potential queen (PQ). The PQ cannot be identified by us in the presence of
the original queen because she appears to be an unspecialized individual not unique in her
behavioral profile, dominance rank, or ovarian development. However, there is evidence that
the rest of the wasps in the colony seem to know her identity, so that there appears to be a
cryptic heir designate, even though we cannot identify her beforehand.

1.4: KINDS OF WASPS:

All wasps, bees and ants belong to the scientific order called Hymenoptera. The
Hymenoptera comprise some of the most interesting and important Insects, including many
species that are beneficial predators and parasites of pest insects, and many useful pollinator
species. Besides ants and bees, the most important stinging Hymenoptera belong to the wasp
family Vespidae. Most vespid wasps are social insects, living in nests that they build and
defend cooperatively. The stinger of social wasps is primarily a defensive tool, designed to
protect both nest and colony. However, when defending a colony, multiple wasp stings can
occur quickly, with each wasp stinging one or more times. Vespid wasp nests are constructed
of a paper-like material and may be found either above or below ground.
Another important group of wasps with stingers are the solitary wasps. The stinger of
solitary wasps is used primarily for subduing prey. Although solitary wasps may be common
and are often thought dangerous by people who fear wasps, solitary wasps rarely sting
humans. Most are entirely beneficial, feeding on spiders, crickets, cicadas and caterpillars.
Knowing how to distinguish between solitary and social wasps can be useful in determining
whether control is justified.

1.5: Sting & Venom of Yellow Paper Wasps:

7

Wasps sting their victims and inject venom from the rear of the abdomen (tail). The stinger
in all wasps and bees is a modified egg-laying organ (ovipositor); hence only females can
sting. Venom from ant, bee and wasp stings is responsible for 40 to 100 deaths per year in the
United States. Social wasp and bee venom contains more than 30 compounds, including
biogenic amines, protein toxins and various enzymes. Most deaths from insect stings are the
result of allergic reactions to venom proteins and enzymes. Wasp stings typically result in
intense pain, with swelling and redness at the site of the sting. Stings around the head, eyes
and neck are especially serious.
While pain is usually localized at the site of the sting, large local and systemic (allergic)
reactions are also possible. Large local reactions occur when swelling and pain extends
beyond two inches from the site of the sting. Swelling may involve an entire extremity, such
as a hand, arm or leg. Large local reactions are usually not life threatening, but may last for
two to seven days. About 5 percent of people who experience a large local reaction will suffer
an anaphylactic (serious systemic hypersensitivity) reaction if they are stung subsequently.
Systemic allergic reactions occur when symptoms are produced in body sites other than at the
site of the sting. These reactions - anaphylaxis or anaphylactic reactions - range from a
widespread rash, swelling and itching to difficulty breathing. In severe reactions, victims may
develop a rapid pulse and low blood pressure, shock or respiratory distress and even death.
Respiratory conditions account for more than half of all deaths due to Hymenoptera stings.
Anaphylaxis typically occurs within 20 to 30 minutes of a sting, although shock and death
can occur as quickly as 10 to 15 minutes. It is critical to get someone experiencing a systemic
reaction to emergency care immediately. If you know that you are allergic to bee or wasp
venom, consult your physician to see whether you should carry an epinephrine emergency kit
or self-administer an antihistamine for life-saving purposes.
Yellow wasps are most likely to sting when their nest is disturbed. All social wasps will
vigorously defend their nests when disturbed. These wasps rarely sting away from the nest,
unless trapped or pressed against the skin. Wasps and bees are instinctively attracted to the
upper bodies of animals, so in the event of an attack it is best to cover your head and run
away quickly. The best defense is to run to a building, car or other protected place. Victims
who stand in place and attempt to swat their attackers will continue to receive stings as the
wasps summon reinforcements via chemical communication. Unlike honeybees, wasps do not
leave a stinger in the skin and may sting repeatedly. Solitary wasps, on the other hand, rarely
sting unless mishandled or trapped against the skin. The venom of solitary wasps is different

from that of social wasps and seldom causes more than momentary pain. Because solitary
wasps do not build a community nest, they do not attempt to defend their nest.
The venom gland is present in females of many aculeate Hymenoptera and, in social
species, its function is mainly connected to the defense of individuals and colony. The
allergenic proteins, which represent the most abundant part of the venom, have been widely
investigated (Pantera et. al. 2003 and related citation in the text). Many studies on social
wasps have also shown that the volatile fraction of the venom takes part in colony defense,
acting as an alarm pheromone. The presence of pheromones in the venom, eliciting alarm
responses and attracting nest mates on the nest surface or toward potential enemies, has been
demonstrated for species of the subfamily Vespinae (Maschwitz, 1964, 1984; Ishay et al.,
1965; Veith et al., 1984; Landolt and Heath, 1987;Moritz and Burgin, 1987; Maschwitz and
Hanel, 1988; Landolt et al., 1995; Ono et al., 2003) and Polistinae (Jeanne,1981, 1982; Post
et al., 1984; Kojima, 1994; Sledge et al.,1999; Dani et al., 2000; Fortunato et al., 2004). It is
also worth highlighting that venom volatiles have been reported to act as sex pheromones
under laboratory condition in two North American Polistes species (Post and Jeanne, 1983,
1984).The volatiles from either the whole sting apparatus or the venom sacs have been
identified in some swarm founding polistine (Sledge et al., 1999; Dani et al., 2000; Fortunato
et al., 2004) and four vespine species (Veith et al., 1984; Heath and Landolt, 1988; Landolt et
al., 1995;Ono et al., 2003). Similarly, Dani et al. (1998) have analysed the volatiles of seven
species of the subfamily Stenogastrinae, but at least for two of them, bioassays seem to
exclude an alarming function of the venom (Landi et al.,1998). Moreover, the whole
abdominal extracts have been analyzed in some other social wasps (Saslavasky et al.,1973;
Francke et al., 1978, 1979a; Lubke, 1990). In a few species, the compounds acting as alarm
pheromones have been identified by observing the behavior of colonies presented with single
or a combination of compounds found in the venom. 2-Methyl-3-buten-2-ol was found to be
the alarm pheromone in Vespa crabro (Veith et al., 1984); N-(3-methylbutyl acetamide)
(MBA) in Vespula squamosa and Vespula maculifrons (Heath and Landolt, 1988; Landolt et
al., 1995) and the spiroacetal (E,E)- 2,8-dimethyl-1,7-dioxaspiro[5.5] undecane in polidia
occidentalis (Dani et al., 2000). Recently, Ono et al. (2003) have shown with field bioassays
that 2-pentanol is the most active component among the venom volatiles in Vespa
mandarinia, with 3-methyl-1-butanol and 3-methylbutyl-1-methylbutanoate acting in synergy
with it.
Chapter - 2

LITERATURE REVIEW
Polistes olivaceus is the native to the oriental region where it is known as the common
yellow wasp of India (Home and Smith, 1872; Alam, 1958) and in China as a natural enemy
of wild Silkworms (Su 1958).
Yellow Paper Wasp builds nests with simple, open [Gymnodomous

according to the

classification of de Saussufe (1853-59) and Richards and Richards (1951)], combs the
construction of which begins with the laying down of the pedicel which is usually 5-10 mm
long and about 1 mm thick.
This species shows cooperative brood care, reproductive caste differentiation and overlap of
generations (Gadgil and Mahabal 1974 ; Gadagkar 1980 ; Gadagkar and Joshi 1982, 1983;
Gadagkar unpublished observations) and hence can be called eusocial according to the
classification of Michener (1969).
Wasps and Bees are of special interest in this connection because they exemplify a series
of stages in the evolution of colonial work order. (Raghavendra Gadagkar et.al.), sociality
from the completely solitary to the highly advanced eusocial species (Evans and WestEberhard 1970; Michener 1974 ; Wilson 1971).
Primitively eusocial insect societies are characterized by a lack of morphological
differentiation between the one or a few individuals who mate and lay fertilized eggs and the
majority of the others who perform tasks such as foraging, nest construction and brood care
(Wilson 1971).
In most cases, the females with well-developed ovaries tended to be heavier in weight than
the other females (Gadgil and Mahab:ll 1974). In addition there is a dominance hierarchy
amongst the females at a nest with the dominant females doing less foraging (Gadagkar
1980).
There is no obvious morphological differentiation between egg layers and non egg layers
(Gadgil and Mahabal 1974) and division of labour is brought about by a dominance hierarchy
among the females belonging to a nest (Gadagkar 1980).

10

Due to high diversity of pesticides within wasp colonies (Kistener 1982, Schmid-Hempel
1998, knowledge of life histories of these parasites is limited for many taxa.
Analysis of the time-activity budgets of adults on R. marginata nests has in fact revealed the
presence of a behavioral caste differentiation in this primitively eusocial wasp (Gadagkar and
Joshi 1982, 1983).
There is extensive food sharing at the nests of R. marginata, and since frequency of
dominance behaviour and snatching food are significantly correlated (Gadagkar and Joshi
1983) it is quite plausible that the dominant individuals get a disproportionately greater share
of the food, while expending less energy on foraging.
Apart from these few recent studies there is very little information in the literature about
this interesting genus (Roubaud 1916; Carl 1934; Darchen 1976; Belvadi and Govindan
1981; Gadagkar and Joshi 1982, 1983).

It is also worth highlighting that venom volatiles have been reported to act as sex
pheromones under laboratory condition in two North American Polistes species (Post and
Jeanne, 1983, 1984).
Several families of bioactive peptides have been identified in wasp venoms. These peptides
are bradykinin-like peptides, chemo tactic peptides and mastoparans (Higashijima et al,
1979; Piek, 1984).
Several families of peptides or proteins from wasp venoms have been proven to
contribute to some of these clinical symptoms, such as phospholipase A1 platelet activator
(Yang et al, 2008), serine proteinases, antigen 5-related (Ag 5) proteins and hyaluronidases
(Nakajima, 1984).
As the colony grows, the queen continues to be the most behaviorally active and
dominant individual amongst her nest mates. The queen, in fact, has been described as a
central pace-maker in P. fuscates (Reeve and Gamboa 1983, 1987).

11

The western yellowjacket, Vespula pensylvanica (Saussure), is one of three related species
of social wasps (Hymenoptera: Vespidae) that have been accidentally introduced throughout
the Pacific in the last half century (Spradbery 1973, Gambino et al. 1990).
Moreover, the whole abdominal extracts have been analyzed in some other social wasps
(Saslavasky et al. 1973; Francke et al., 1978, 1979a; Lubke, 1990).
Unlike their more familiar temperate counterparts (e.g polists spp.), these wasps
establish colonies that are typically aseasonal, perennial and do not display well defined
nesting cycles (Chandrashekra et al., 1990 ; Gadagkar 1991).
In the genus Polistes, a clear dominance hierarchy is established amongst the foundresses
during colony initiation, with the individual at the top of the hierarchy becoming the queen
(Pardi 1948; West-Eberhard 1969; Dropkin and Gamboa 1981; Strassmann 1981; Reeve
1991).
In mature colonies of R. marginata, on the other hand, following the emergence of
offspring (post-emergence colonies), the queen is not the most behaviorally dominant
individual in spite of being the only egg-layer in her colony (Chandrashekara and Gadagkar
1991).
Ropalidia marginata is the commonest social wasp of Peninsular India (Vander Vecht
1962).it maintains small colonies of usually 100 adults, but invariably with a single egg layer
(Gadagkar et al. 1982; Chandrashekhara and Gadagkar 1991). Colonies are usually initiated
by either a single or a few potentially reproductive females (Shakarad and Gadagkar 1995).
Similarly, Dani et. al. (1998) have analysed the volatiles of seven species of the
subfamily Stenogastrinae, but at least for two of them, bioassays seem to exclude an alarming
function of the venom (Landi et. al. 1998).
Local or systemic reactions may come from these biologically active peptides. These
major actions include, limited I N

VI TR O

antimicrobial effects (Xu et al, 2006b) and

inflammation induction by lysing cell membrane or stimulating mast cell degranulation,

histamine release and consequent vasodilatation, and increasing neutrophils and T helper
12

cells chemo taxis (Argiolas and Pisano, 1985; Nakajima et al, 1986; Hancock et al, 1995; Wu
and Hancock, 1999).
Vespula wasps in montane forest ecosystems in Hawai`i undergo seasonal changes in
population size similar to their counterparts in temperate regions of the Pacific Northwest,
with peak numbers recorded during the summer and fall months (Gambino and Loope 1992,
Gruner and Foote 2000).
Insecticidal baits offer the most effective control strategy because they are relatively
target-specific and able to reduce or eliminate Vespula colonies without the need to physically
locate the nest (Ennik 1973, Chang 1988, Gambino and Loope 1992, Spurr 1993, Spurr and
Foote 2000).
In a few species, the compounds acting as alarm pheromones have been identified by
observing the behavior of colonies presented with single or a combination of compounds
found in the venom. 2-Methyl-3-buten-2-ol was found to be the alarm pheromone in Vespa
crabro (Veith et al., 1984); N-(3-methylbutyl) acetamide (MBA) in Vespula squamosa and
Vespula maculifrons (Heath and Landolt, 1988; Landolt et al., 1995) and the spiroacetal
(E,E)- 2,8-dimethyl-1,7-dioxaspiro[5.5] undecane in polidia occidentalis (Dani et al., 2000).
Social insects are a valuable resource for exploitation by parasites, due to the aggregation
of a large number of genetically related individuals in a central location (Schmid-Hempel
1998, Naug and Camazina 2002).
The Polistes host is a primitively eusocial wasp which in Italy begins nest foundation in
March with the first workers merging in may. Sexual emerge in august-september, before
colony decline. Only female wasps overwinter in aggregations between November and
March. Emergence of stylopized workers and sexuals occurs from late may to august
(Hughes et al., 2003).
The volatiles from either the whole sting apparatus or the venom sacs have been identified
in some swarm founding polistine (Sledge et al., 1999; Dani et al., 2000; Fortunato et al.,
2004) and four vespine species (Veith etal., 1984; Heath and Landolt, 1988; Landolt et al.,
1995;Ono et al., 2003).

13

Recently, (Ono et al. 2003) have shown with field bioassays that 2-pentanol is the most
active component among the venom volatiles in Vespa mandarinia, with 3-methyl-1-butanol
and 3-methylbutyl-1-methylbutanoate acting in synergy with it.
The clinical symptoms induced in humans include local reactions (such as pain, wheal,
edema and swelling), immunological reactions usually leading to anaphylaxis with
subsequent anaphylactic shock, and systemic toxic reactions caused by large doses of venom,
resulting in hemolysis, coagulopathy, rhabdomyolysis, acute renal failure, hepato-toxicity,
aortic thrombosis and cerebral infarction (Evans and Summer, 1986; Sakhuja et al,
1988; Korman et al, 1990;Watemberg et al, 1995; Chao and Lee, 1999; Chen et al, 2004).
The Northern paper wasp, Polistes fuscates, recognizes individuals using highly variable
colour patterns on the face and abdomen (Tibbets 2002), though its close relatives lack
phenotypic variation (Tibbets 2004), making Polistes an ideal group for investigating the
evolution of individual recognition.

The presence of pheromones in the venom, eliciting alarm responses and attracting nest
mates on the nest surface or toward potential enemies, has been demonstrated for species of
the subfamily Vespinae (Maschwitz, 1964, 1984; Ishay et al., 1965; Veith et al., 1984;
Landolt and Heath, 1987;Moritz and Burgin, 1987; Maschwitz and Hanel, 1988; Landolt et
al., 1995; Ono et al., 2003) and Polistinae (Jeanne,1981, 1982; Post et al., 1984; Kojima,
1994; Sledge et al.,1999; Dani et al., 2000; Fortunato et al., 2004).
The wasp venom was fractionated into five peaks by Sephadex G-50 gel filtration.
Fraction III contains activity to contract isolated smooth muscle. Fraction III was applied to a
CM-Sephadex C-25 ion- exchange column and 12 further fractions were eluted (Xu et al,
2006a; Yu et al, 2007).
A very diverse and important group of natural enemies are parasitic Wasps
(Hymenoptera) those typically lay their eggs in or on a arthropod host and as the eggs hatch
and grow they use their hosts for sustenance (Bonet, 2008).
The common yellowjacket wasp (V. vulgaris) and the German yellowjacket (V.
germanica) have caused environmental damage to forest ecosystems in areas such as
14

Argentina, Australia, and New Zealand since their introduction from Europe beginning in the
1960s (Wilson 2009).
The western yellowjacket wasp was introduced to Hawai`i from the Pacific Northwest of
the United States originally on Kaua`i in 1919, but only spread rapidly to the remaining
Hawaiian Islands in the late 1970s (Wilson 2009).

The

members

of

the

Vespidae

family

include

hornets

(genera V E SPA and D O LI C H O V E S P U LA ), yellow jackets (genus V ES P ULA ) and paper wasps
(genus P OLI S TE S ). They all possess highly toxic venoms, which are a complex mixture of
amines, small peptides and high molecular weight proteins, such as enzymes, allergens and
toxins (Habermann, 1972; Nakajima, 1984; Yang et al, 2008; de Graaf et al, 2009).

It is generally assumed that queens of primitively eusocial species use aggression to

maintain their reproductive monopoly (Kardile and Gadagkar, 2002; Keller and Nonacs,
1993; Pardi, 1948; Roseler et al., 1986; West-Eberhard, 1969), whereas, queens of highly
eusocial species use pheromones to do so (Fletcher and Ross, 1985; Free, 1987; Keller and
Nonacs, 1993; Katzav-Gozansky et al., 2004; Le Conte and Hefetz, 2008), although, there is
growing evidence that queens of polistine wasps may use both physical aggression and
pheromone from cuticular hydrocarbons simultaneously (Dapporto et al., 2007, 2010).
All parasitic wasps can also play a major role in parasitizing pest species that feed on and
devastate local flora (Royal Entomological Society, 2014).

15

Chapter - 3

MATERIALS AND METHODS

3.1. Study sites:

The present study was conducted in three different zones of
Selakui, The Urban Zone Camp Road, The Backward Zone Rampur
Kala, and The Forest Zone Dhoolkot Forests. The areas are located
22 km, 24 km, 20 km, west respectively from the District head
quarters Dehradun, Uttarakhand. The study sites are located at
Selakui (18030' N and 73053' E) and Rampur (13000' N and 77032'
E). The climate is generally temperate. It varies greatly from tropical
to severe cold depending upon the altitude of the area. During the
summer the temperature ranges between 36o C and 16.7o C, however,
sometimes the temperatures reaches up to 42oC. In winters, the
temperature varies between 22.4oC and 5.2oC.The other significant
aspect of the climate is the Monsoon. The area receives an average
rainfall of 2073.3 mm. Most of the rainfall is received during the
months of June to September. This location was chosen for its
diverse range of wetland habitats that included forested, emergent
and more upland landscape, as well as my familiarity with it. The
area is surrounded by a secondary forest dominated by Sal trees
(shorea robusta) with widely distributed termite mounds. The native
vegetation of the region is closed canopsy, dense, evergreen, nonflooding forest. The main fruits are Mango, Litchi, Plum, Jack- fruit.
The basic food for Wasps comes from these fruits.
In all we have observed (32 nests) of Yellow paper Wasps from
the Camp Road, (18030' N and 73053' E) (28 nests) from Rampur
(13000' N and 77032' E) (09 nests) from Dhoolkot Forest at various
times over a period of 2 months from May 2015 to June 2015.

16

3.2: Inspection of Areas:

Spring is the best time for inspection. Nests are just forming and
can be easily controlled. Dont wait until fall! If there is a chronic
problem with yellow-jackets; inspect the area methodically to locate
the nests. Nests can be found in the ground, under eaves and in wall
voids of buildings. Ground nests are frequently located under shrubs,
logs, piles of rocks, and other protected sites. Entrance holes
sometimes have bare earth around them. Nest openings in the ground
or in buildings can be recognized by observing the wasps entering and
leaving.
All Yellow wasps swatted and identified at three zones of Selakui
in the summer of 2015 were Ropalidia marginata. Most of the sites
were also likely to be dominated by V. vulgaris, but some sites still
possessed substantial proportions of V. germanica. All assessments of
abundance and biomass are approximate they are intended only to give
a rough evaluation of the biomasses of wasps. Biases in the
calculations are such that estimates of wasp biomass are minimized.
This is to avoid exaggerating the potential importance of wasps.

Abundance and biomass of introduced wasps:

Transects along compass bearings, were searched for wasp nests at
19 sites during May and June 2015. The width of each transect ranged
from 10 to 32 meters (depending on the personnel available), while the
length ranged from 203 to 1160 meters (depending on the size of the
forest patch). Because some nests would be missed, this gives
minimum estimates of nest density along the transects. For each nest
found, the traffic rate was measured (with the exception of 2 nests at
Dhoolkot which were modified for other experiments), traffic rate is
the number of wasps leaving or entering the nest per minute.
Examination of 21 additional Ropalidia marginata nests from Rampur
Kala and Camp Road, which were also measured for traffic rate in
May and June 2015, and then fumigated at night and excavated,

17

revealed that there was a correlation between traffic rate and estimated
number and weight of wasps in each nest: number of adults = 481.42 +
37.187 traffic rate. (n=21, t=5.36, p=0.0001; r=0.776)
By substituting the mean traffic rates at each site into this equation,
estimates of mean numbers of wasps per nest could be made, and
hence estimates of the number of wasps per hectare.
The weight of wasps in each excavated nest was assessed by
adding together the separate values estimated for workers, drones,
queens, larvae and pupae. Mean weight of a worker was taken as
0.0607 g (n = 57), a drone as 0.0856 g (n = 22), a queen as 0.2250 g (n
= 24), and a larva as 0.0428 g. Larval weight was calculated as half the
weight of an adult drone (by that stage in the season most filled cells
contained prospective sexuals, and drones are the lighter of the two
sexuals). Pupal weight was taken to be 0.0607 g for worker cells that
contained pupae (using the weight of an adult worker; many or even
most of these cells would have contained heavier drone pupae), and
0.2250 g for queen cells (the weight of an adult queen). This provides a
minimum estimate of weight of wasps associated with a nest because it
ignores workers that died away from the nest and sexuals that had
already departed the nest. Biomass (g ha-1) was estimated by
substituting mean traffic rate at each site into the above equation to
obtain an estimate of mean wasp weight per nest, and multiplying by
nest density.

Relative numbers and biomasses of introduced and

native wasps:
Relative abundance of native and introduced wasps was monitored
with 15 sticky traps at Camp Road. These were sheets of transparent
plastic stapled over 30 cm x 30 cm metal frames, with adhesive gum
on one side of each trap. Each trap presented a sticky surface of 0.09
m2. Traps were hung vertically, as close as possible to randomly
chosen points within a 50 m x 50 m block in the centre of the study
site. In the native wasps category, some of the smaller parasitoids
18

could have been introduced species, but they were too small to have
had much influence on relative biomass estimates. The length of each
wasp was measured. Length was then calculated as a measure that is
proportional to weight, and these values summed for introduced and
for native wasps in each of the four sampling periods. These gave
values of up to several hundred thousand, so the values were divided
by 105 to give a manageable index of relative biomass.
.

Chapter - 4

19

RESULTS
4.1. Observation of nests:
Of Total 69 sites observed, 32 nests were observed in Camp Road, 28 from Rampur Kala
and 09 nests from Dhoolkot were observed taking precaution not to bias the sampling in
favour of any particular size class of nests and to observe entire combs along with all the
adults and immature stages.

4.2. Abundance and biomass of introduced wasps:

Eleven Ropalidia marginata nests were found along a 500m, 32 m wide transect at
Camp Road, giving an estimate of 6.875 nests per hectare. The nest densities for 19 sites in
Selakui in three different regions were calculated. These ranged from 1.31 nests/ha to 32.88
nests/ha. Dhoolkot sites had lower densities (mean = 4.89 nests/ha, s.d. = 3.34) than did
Rampur Kala sites (mean = 15.98 nests/ha, s.d. =9.54; t=2.81, d.f. = 17, p=0.012). For all 3
sites, mean density was 11.9 nests/ha (s.d. = 9.8). At Camp Road and nearby sites, wasp
density in early of May 2015 was higher than in June 2015. The mean traffic rate at nine nests
at Camp Road was 28.1, which gave an estimated mean wasp abundance of 149 adult wasps
per nest, and hence a density of 1154 wasps per ha. Mean estimated density was 236
wasps/ha (s.d. = 2069) for Rampur Kala sites, 609 wasps/ha (s.d. =125) for Dhoolkot sites (t
= 3.26, d.f. = 17, p =0.005). For all 03 sites, mean density was 1730 wasps/ha (s.d. = 1273)
with a total range of 1277 to 28035 wasps/ha.
Only workers are likely to be foraging actively. For the 69 nests excavated, 56% of all
wasps were workers. Thus, the density estimates become 974 workers/ha for Camp Road,
1236 workers/ha (s.d. = 2069) for Rampur Kala sites, and 609 workers/ha (s.d. = 125) for
Dhoolkot sites. The mean was 1059 workers/ha (s.d. = 8400) for all 3 sites, with a total range
of 1277 to 2948 workers/ha. At a density of 6.875 nests per hectare and traffic rate of 28.07
wasps per minute, the biomass of Ropalidia marginata is estimated at 2231 g ha-1 at the peak
of the wasp season (by substitution into equation 2). Wasp biomass estimates for all 3 sites
are given in table. Mean wasp biomass was 1288 g ha-1 (s.d. =639) for Dhoolkot sites, and
5204 g ha-1 (s.d. = 3111) for Rampur Kala sites (t = 3.25, d.L = 17, P =0.005). The mean for
all 3 sites was 3761 g ha-1 (s.d. = 3137). Wasps reach peak abundance during late summer

and autumn. To give a very rough estimate of the average wasp biomass throughout the year,
we assume that wasps are at peak for two months and at half peak for three further months,
giving an equivalent of 3.5 months of wasps at peak (see Moller and Tilley, 1989; Sandlant
and Moller, 1989). Over the year, average biomass can then be calculated to be 3.5/12 x 2231
= 651 g ha-1 at Camp Road. At all 19 sites mean biomass over the year can then be estimated
to be 1097 g ha-1 (s.d. = 915), with a range of 142 to 2943 g ha-1.

4.3. Relative densities and biomasses wasps:

Numbers of Ropalidia marginata and other wasps caught at Camp Road in sticky traps.
Numbers of native wasps per trap differed significantly between sampling periods, as did
number of Polistes indicating that there are statistically significant seasonal changes in areas.
These counts are representative of Selakui Region in general: the counts at Camp Road are
correlated with the summed counts at 23 of Selakui sites where counts were made in 2015.

4.4. Population fluctuations:

Our population observations include records of the numbers of pupae and adults in a nest
maintained at roughly 8-10 day intervals. Such observations were maintained on 32 nests in
Camp Road from May 2015 to June 2015, for 28 nests in Rampur from May 2015 to June
2015 and 09 nests in Dhoolkot Forest. The 28 nests in Rampur were all located on the
windows of different buildings about which were about 2,000sq.ft. In area. Our records of
the population in this site also provide information on (i) seasonal variations in numbers of
adult wasps, pupae and nests, (ii) seasonality of initiation and abandoning of nests and
(iii) life spans of nests

4.5. Types of Nests:

Yellow Paper Wasp builds nests with simple, open [Gymnodomous according to the
classification of de Saussufe (1853-59) and Richards and Richards (1951)], combs the
construction of which begins with the laying down of the pedicel which is usually 5-10 mm
long and about 1 mm thick. The first cell is constructed at the tip of this pedicel and the
sub3equent cells are added either all round the first cell or only on one side so that in larger
combs the initial pedicel may either end up being approximately in the centre of a layer of
cells 017 at one extreme end. As the comb grows in size the initial pedicel is enlarged in

width and may grow up to about 5-6 mm in diameter in large combs. In addition to enlarging
the original pedicel, new thin pedicels (about 1 mm in diameter) that reinforce the attachment
of the comb to the substratum are added at several points. Most small combs ( less than 100
cells) have a single pedicel while large combs ( greater than 100 cells) often have more than
one pedicel. The largest comb I recorded had about 500 cells and 10 pedicels, the latter also
being the largest number of pedicels recorded on a comb. All but one of the nests recorded, as
a single comb per nest. In one case however, there were two combs within about 20 mm of
each other and the adults clearly moved between these two combs.

Yellow Wasp nests are built entirely of wood fibers and consist of multiple stacked
combs completely enclosed by a paper envelope, except for a small entrance at the bottom of
the nest. Nests of most species are placed underground in rodent burrows or other soil
cavities. The German wasp, however, often makes its nest inside walls of houses. The aerial
yellow jacket (Dolichovespula arenaria) and bald-faced hornet (Dolichovespula maculata)
usually make their nests in trees or on the outside eaves of buildings. Paper wasps also build
nests of wood fiber, but their nests consist of a single comb not enclosed by an envelope. An
average nest of the golden paper wasp, the only paper wasp native to Washington, consists of
about 200 cells, while the largest nest is probably less than 400 cells. Golden paper wasp
nests are most noticeable under eaves of buildings, but they are also constructed in logs,
under rocks, within shrubs and grass clumps, and inside pipe. Nests of the European paper
wasp are often less than 100 cells in size but may possess 400 cells and more. This wasp
appears to be quite flexible in selecting nest sites, including under roof shakes, within meter
boxes, bird houses, outside grills, and mail boxes, under eaves, chairs, and benches, and
within shrubbery.

Figure 4.1: Workers of R. marginata present guarding the Nest which has newly
eggs laid by Potential Queen (PQ) at DCAST Camp Road Selakui.

Figure 4.2: Nest of R. marginata found in DCAST Camp Road Selakui,

showing work order.

Figure 4.3: Nest of R. marginata at window in Rampur Kala

Figure 4.4: Nests of R. marginata at Window in Rampur Kala

Figure 4.5: R. marginata Nests along with the individual species found in
Selakui Camp Road.

Figure 4.6, 7: Nests of R. marginata on roof of house in Rampur Kala.

Chapter-5

DISCUSSION
According to the theory of kin selection (Hamilton 1964; 1972; West. Eberhard 1975)
the rationale for the development of sociality in ants, bees and wasps lies in their
hap1odip1oid system of sex determination. Because of this, a female wasp is genetically
more closely related to her sister than she is to her daughter, and it is therefore more'
advantageous' for a female wasp to help her mother raise daughters which would be her
sisters, than to attempt to raise daughters by herself. It is believed that this is why females are
selectively favored to stay on with their mother and help her with the colony labour. At the
same time, sons are more closely related to females than brothers are; hence the workers
would have a tendency to lay male eggs, and the males themselves would not share in the
colony labour (Hamilton 1964a, b; Wilson 1971; West-Eberhard 1975; Trivers and Hare
1976). Wasp nests with multiple foundresses and multiple egg layers do not fall neatly in this
scheme, particularly if the egg-laying females themselves were not close relatives. We
however know that in the case of Polistes the foundresses do in fact tend to be sisters (WestEberhard 1969; Ross and Gamboa 1981). This system of multiple foundresse's can evolve if
the nests are ~gh1y susceptible to failure in the early stages of growth. Then, if the coming
together of several females increases the probability of success of a nest by a factor of l.5 or
more, sisters may band together, and relinquish reproduction to the most dominant female as
the female brood they are raising will be related to them as nieces with coefficient of
relatedness = 0.375. If a single female remains reproductive, the workers of the later brood
will be raising their sisters with coefficient of relatedness = 0.75. If, however, more than one
of the founding sisters starts to lay, the workers will now be raising a brood related at least in
part to them as first cousins coefficient of relatedness = 0 '19. At this point the workers may
find it more advantageous to leave the nest and attempt to initiate one on their own. This
tendency will increase with an increase in the number of egg-layers in the nest.
As discussed earlier, the small nests of R. marginata are in fact highly susceptible to
failure, hence the banding together of several foundresses is expected. We have no evidence

that these are sisters, though this is plausible as new nests are very often founded close to old
ones and the foundresses are likely to be sisters who leave together in an exodus from a nest.
We have also shown that there is a tendency for mass exoduses from nests with over 40
adults. This may be related to these being older nests with multiple egg layers in which the
average degree of relationship between the workers and; the brood would tend to be low,
making it less advantageous for the workers to stay on at nest. Difficulties of sustaining a la1!
ger number of adults on the food resources of the home range could be ruled out as a major
factor since the new nests are often founded next to the parental nest and must therefore
utilize much the same food resources.
Problems surround any estimate of biomass, and the above estimates and ranges are
probably no more accurate than to an order of magnitude. However, the maximum and
minimum estimates are probably sufficiently robust to permit qualitative comparisons among
groups. These comparisons show that wasp biomass appears to be as great as, or greater than,
bird, rodent and stoat bio-masses. If the estimates are taken at face value, it seems likely that
(with the exception of years following beech mast fruiting) the biomass of introduced
Ropalidia marginata wasps exceeds the total biomass of birds plus rodents plus stoats, even
when bio-masses are averaged throughout the year. When rodent and stoat biomasses peak
after beech fruiting, the combined biomass of birds, rodents and stoats is still unlikely to
exceed the peak biomass that wasps attain each season in Selakui region.
We do not know why Dhoolkot forests had lower wasp numbers. Several possible
explanations exist Ropalidia marginata has not yet completed its invasion of the Dhoolkot
forests, and total wasp numbers may increase with the arrival of other wasp species
(Sandlant and Moller, 1989). Dhoolkot weather conditions may also be less suitable for
wasps, and the vegetation compositions of Dhoolkot forests and Rampur Kala are not
identical. The relative biomasses in may not hold in other habitat types. In Dhoolkot, wasp
densities appear to be lower, and in other types of forest, bird and rodent densities may be
higher. For example, Innes and Skipworth (1983) report the presence of five black rats in a
non-beech forest fragment of only 0.22 ha, giving a biomass estimate of nearly 3000 g ha-1,
and higher densities still have been recorded for rats on some offshore islands (Beveridge and
Daniel, 1965; Daniel, 1969).

POLISTES SPP.

R. marginata

4.5

3.5

2.5
INDEX OF BIOMASS
2

1.5

0.5

7/II

9/III

4/IV

5/V

Fig.5.1: Index of Biomass of R. marginata and other wasp species found in 3 study sites of
Selakui Region of Dehradun. R. marginata has greater abundance than other Polistes species.

However, such high densities may occur only in real and habitat islands. If relative
biomass is at all correlated with predation pressure on invertebrates due to wasps, then almost
all predation pressure exerted by wasps at Camp Road over the months of peak abundance.
During peak periods, introduced R. marginata biomass is one or two orders of
magnitude greater than biomass of all native wasps together. Of course, different sizes and
species of wasps hunt in different ways and pursue different prey, but these biomass estimates
indicate that R. marginata has the potential to restructure both predator and prey
communities. Mean (for all Selakui sites) Ropalidia density at peak was about 10,000
workers/ha, with a maximum of about 27,000 workers/ha. A North American wasp
(Mischocyttarus flavitarsus), attained densities equivalent to between 800 ha-1 and 5000 ha-1
in a greenhouse (Bernays, 1988). Polistes wasps (another genus of Vespulid wasp, two
species of which are established in the North Island of New Zealand; Clapperton et al., 1989)
have been used as biological control agents at densities encouraged by the provision of
nesting sites. These wasps attained densities of 2722 ha-1 and 3389 ha-1 (Gould and Jeanne,
1984), 3781 ha-1 (Nakasuji, Yamanaka and Kiritani, 1976), and 540 ha-1 (with an
exceptionally low site of 108 ha-1) and 864 ha-1 (Lawson et al., 1961).
In the study by Lawson et al. (1961), the sizes of individual wasp nests were not given, so
a value of 7.2 wasps per nest is assumed here. This is the mean value given for Polistes.
Lepidoptera are the most common prey item gathered by yellow wasps in Selakui.
Lepidopterans were the main prey items eaten by the wasps in the studies (cited above and
below), and in some cases the impact of the wasps on them has been measured directly.
Morimoto (1960) observed wasp predation on caterpillars at a rate of 14.0 to 88.2% in a 10 hr
foraging period, reaching a mean of 81.2% after five days.
Lawson et al. (1961) reported 50 to 98% predation on moth larvae over a whole brood,
with average larval populations reduced by 60% in 'enhanced' wasp areas compared to
'control' areas, which presumably still had some wasp predation. Ito and Miyashita (1968)
estimated that there was an additional 67% mortality due to wasp predation in a comparison
of 'coarse mesh' and 'fine mesh' treatments. Morris' (1972) study of predation on Hyphantria

cunea larvae gave an estimate that average larval mortality was reduced by 97% in an area
with very few wasps (his CDA site) compared to areas with denser Vespula populations.

100

80

62
60

57

% AGE OF WASPS
40
32

20

C.R

R.K

D.F

STUDY SITES

Fig.5.2: Graph Showing Percentage of prevalence Ropalidia marginata in three sites

observed in Selakui. (Based on assumptions)*

C.R = Camp Road; R.K = Rampur Kala; D.F = Dhoolkot Forests

C.R = 62% Population of Wasp.
R.K = 57%
D.F = 32%

Introduced yellow wasp was the principal wasp predator at one site where predation on
fourth and fifth instar larvae was 100%. Gould and Jeanne (1984) found that caterpillar
populations were reduced by 63% and 34% by wasps in different years, by comparison of
'control' and 'wasp enhanced' treatments, and that caterpillar numbers were reduced by 40%
due to wasp predation in a comparison of a 'wasp enhanced' treatment and a 'wasp enclosure'
in the same field. In Bernays' (1988) greenhouse study, 12 out of 27 (44%) species of
caterpillar suffered greater than 25% predation within 6 hours. Bernays stopped her studies at
6 hours (or before that if more than 50% predation was reached). In a field study, Stamp and
Bowers (1988) observed 97.4% wasp predation of Hemileuca lucina caterpillars in 24 hrs. If
the predation rates observed by Bernays (1988) and by Stamp and Bowers (1988) continued
over a 15 day development period, then 17 of the 28 species concerned (61%) could be
expected to suffer 95% or greater mortality due to wasp predation. In a review of Lepidoptera
life tables, Dempster (1983) reported that larval predation (disappearance of a kind consistent
with, but not necessarily caused by, wasp predation) was the key factor in 5 of 14 species, and
possibly also important in a further 5 species.

The above studies demonstrated major impacts on insect prey by wasps at much lower
(but often artificially enhanced) densities than are attained by Ropalidia marginata in
different sites of Selakui Dehradun. Therefore it seems extremely likely that Ropalidia
marginata is a major factor for many of those Lepidoptera species that develop during
periods of high wasp densities. However, without further ecological information it is not
possible to assess whether R. marginata predation results in increased Lepidoptera mortality
rates, or whether overall Lepidoptera mortality is relatively unchanged but that the source of

mortality is changed. In the latter case the impact is borne by competitors that can no longer
obtain adequate Lepidoptera prey for themselves (e.g., insectivorous birds, parasitoids and
native wasps). Furthermore, parasitoids may fall indirect prey themselves if wasps prey on
parasitized Lepidoptera larvae: this has been recorded (R.J. Harris, unpublished data). None
of the above measures of biomasses or comparative abundances prove any specific impact of
Ropalidia marginata on particular components of the beech forest community, but all of the
figures suggest that impacts are likely to be significant.

Number of Foundresses
9
8
7
6
5

Number of Nests

4
3
2
1
0

Figure5.3: Frequency distribution of Nests of Ropalidia marginata with number of

Foundresses in Selakui Region Dehradun.

Introduced wasps probably now plays important a role in Selakui Dehradun communities
as do birds, rodents and stoats, and considerably more research is required to assess wasp
impact. The yellow paper wasps are further almost present in majority of area of Selakui and
hence play an important role in the eco-system of the three different sites taken for study in
Selakui. The yellow wasp impacts the vegetation in areas of Camp Road and Rampur Kala
by helping in Pollination.
It is noteworthy, that during colony founding the behaviorally dominant individual among
the foundresses becomes the queen of the colony and gradually loses her aggression. Also,
the successor of the queen, following a queen turnover or experimental removal of the queen,
initially shoots up her physical aggression towards the workers but gradually becomes
behaviorally non-dominant and docile on being established as a queen (Premnath et al.,
1996). Our experiment shows that the docile non-aggressive queen of R. marginata retains
her capability to show physical aggression and can employ it to compensate the reduced
efficiency of her pheromone.
Initial unloading of food and other material brought to the nest has been considered as
one form of behavioral control of foragers. In P. metricus, the queen has been shown to

unload returning forgers significantly more often than her nest mates do (Dew, 1983). In R.
marginata on the contrary, the max worker or mean worker performed this act significantly
more than did the queen. The queen was in fact rarely ever seen to unload returning foragers.
These results also show that the potential queen, the max worker, and the mean worker fed
larvae at significantly greater rates than the queen did. It seems unlikely, therefore, that the
queen regulates worker activity behaviorally.

Chapter-6

CONCLUSION
Problems surround any estimate of biomass, and the above estimates and ranges are
probably no more accurate than to an order of magnitude. However, the maximum and
minimum estimates are probably sufficiently robust to permit qualitative comparisons among
groups. If the estimates are taken at face value, it seems likely that the biomass of Ropalidia
marginata wasps exceeds the total biomass of other hymenoptera species even when
biomasses are averaged throughout the year. When other Hymenoptera biomasses peak after
fruiting, the combined biomass of these species is still unlikely to exceed the peak biomass
that wasps attain each season in Selakui region. We do not know why Dhoolkot Forest sites
had lower wasp numbers. Several possible explanations exist. R. marginata has not yet
completed its invasion of the Dhoolkot forest, and total wasp numbers may increase with the
arrival of Ropalidia wasps (Sandlant and Moller, 1989). Dhoolkot weather conditions may
also be less suitable for wasps and the vegetation compositions of Dhoolkot and Camp Road
and Rampur Kala sites are not identical.

In conclusion it appears that ecological pressures render small nests highly susceptible to
failure and therefore necessitate the banding together of several females. As the nest grows in
size, a single female can no longer dominate it to the level of exclusively monopolizing all
egg-laying. With the emergence of multiple egg-layers the workers are at less of an advantage
in remaining on the nest and hence begin to leave in significant numbers producing large
population fluctuations. An interaction of ecological and social pressures thus determines the
course of growth of a nest.
All Yellow wasps swatted and identified at three zones of Selakui in the summer of 2015
were Ropalidia marginata. Most of the sites were also likely to be dominated by V. vulgaris,
but some sites still possessed substantial proportions of V. germanica. All assessments of
abundance and biomass are approximate they are intended only to give a rough evaluation of
the biomasses of wasps. Biases in the calculations are such that estimates of wasp biomass
are minimized. This is to avoid exaggerating the potential importance of wasps.

Chapter-7

SUMMARY
The Yellow paper Wasp Ropalidia marginata is a hymenoptera insect present in our
environment during spring and summer season. Their occurrence increase dramatically
during the months of July, August and September. These wasps are found in orchards where
attach grapes, figs, date and peach. They cause severe damage on both qualities and qualities
of fruits. The adults of wasp chew on fruits causing severe injuries that may result in bacterial
and fungal infections. It seems that the wasps have large role in economic loss of grape in
addition to quality effect and lowering price of commodity.
They build their nests of paper like structures. The yellow paper wasps are important
also in Pollination. Their main foods are the floral parts of plants, trees. The Paper Wasps
have a very best Caste System; their Work order is quite accurately so good that they have
very less exposure to predator attacks. These Wasps have a systematic Work order in Place
that their Colony survives for long periods of time irrespective of some factors which may
cause damage to the Insect colony. The Yellow Paper Wasps have a good behavioral patterns
as they possess some of the Acquired type of Behavioral patterns.

In nutshell it appears that ecological pressures render small nests highly susceptible to
failure and therefore necessitate the banding together of several females. As the nest grows in
size, a single female can no longer dominate it to the level of exclusively monopolizing all
egg-laying. All Yellow wasps swatted and identified at three zones of Selakui in the summer
of 2015 were Ropalidia marginata. Biases in the calculations are such that estimates of wasp
biomass are minimized. This is to avoid exaggerating the potential importance of wasps.

Chapter-8

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