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One factor likely to regulate the abundance of monarchs
is their interaction with natural enemies. Many predators, parasites,
and parasitoids attack monarch eggs, larvae, and adults. Predation
occurs when one organism (a predator) kills and consumes another
organism (a prey) to obtain energy and nutrients. Predators
such as spiders and ants attack eggs and young larvae feeding on
milkweed, whereas birds and wasps have been observed preying on
adult monarch butterflies. Parasitoids are specialized
insects such as small flies and wasps that lay eggs on other insects.
Parsitoid larvae then eat their prey from the inside out, usually
emerging from the prey carcass as a pupa or adult. Smaller organisms
that live and multiply inside their hosts, taking nutrients and
resources, are called parasites. Parasites can
be unicellular microbes such as viruses and bacteria, or larger
organisms like mites and nematodes.
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Monarch
defenses and warning coloration
Many prey species have mechanisms to avoid predation,
including camouflaged coloration or bright eye-spots to confuse
predators. Bright coloration in insects and other animals (typically
yellow, orange, or red) can act as signal, warning other animals
that they are poisonous or distasteful. Such color patterns are
called "aposematic." When an animal attacks, eats, or
encounters such a brightly colored animal and gets stung, bitten,
or poisoned, it learns to associate these warning colors with a
bad experience. Monarchs have a chemical defense that is toxic to
many natural enemies - they can sequester poisonous compounds from
milkweed called cardenolides, or cardiac glycosides (Zalucki et
al. 1990, Ritland and Brower 1993, Brower et al. 1994, Frick and
Wink 1995). Thus, when an animal eats a monarch and gets sick, it
learns to avoid potential prey with similar coloration. However,
research has shown that these toxins break down over time in adult
monarchs, and by several weeks of age the butterflies are much more
palatable to predators (Fink and Brower 1981, Brower and Calvert
1985, Brower 1988, Alonso M. and Brower 1994, Sakai 1994). In addition,
the role of sequestered chemicals in defending monarchs against
parasitoids and pathogens has not been explored.
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Monarch
larvae and adults display bright warning coloration as
a signal to potential predators. |
Birds such as black-beaked orioles and black-headed
grosbeaks are common predators at monarch overwintering sites. These
species can eat large quantities of monarchs without getting poisoned.
This may result from the decay of toxins inside the monarchs
bodies during the many months of migration and overwintering, or
from the specific feeding behavior of the birds. Orioles slit open
the monarchs abdomens before feeding, avoiding most of the
toxin-rich cuticle. Grosbeaks, which eat the entire abdomen, can
tolerate higher levels of cardenolides in their digestive tracts.
Research has shown that predation by these two bird species accounts
for over 60% of the total monarch mortality during overwinter, and
together they kill 7-44% of the total monarchs in any one colony
(reference).
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Orioles
(L) are avian predators of monarchs overwintering in Mexico.
Many dead monarchs litter the forest floor in Mexico (R),
such as these victims of bird predation. |
Invertebrate predators such as ants, spiders, and
wasps attack monarch larvae on milkweed plants. Less is known regarding
invertebrate predation in monarchs, but wasps have been observed
feeding on monarch abdomens at a California overwintering site (D.
Frey, personal communication), and fire ants have been suggested
as a major predator of monarch larvae in Texas (Calvert 1996). Other
research suggests that wasp predators may be sensitive to the chemical
defenses of monarch larvae, and that wasps fed monarch larvae with
high cardenolide concentrations had lower reproductive potential
and more deformities in their nests (L.S. Rayor, personal communication)
than wasps that preyed upon less toxic caterpillars.
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An
assassin bug (L) pierces the cuticle of a monarch larvae
and draws out the inner fluids and tissues. Ants
(R) attack a fourth instar larva that crawled onto the
wrong leaf. |
A few species of parasitoids lay their eggs on monarch
larvae, including tachinid flies and braconid wasps. Tachinid fly
larvae feed on monarch caterpillars, but usually dont kill
their hosts until just before the caterpillars pupate. When a parasitized
caterpillar hangs upside down in the pre-pupal "J"-shape,
several fly maggots will emerge from the dorsal anterior end and
drop to the ground on long, gelatinous tendrils. Braconid wasp parasitism
is less common, but as many as 32 tiny adult wasps may emerge from
a single monarch carcass. Very little is known about variation
in rates of parasitism and predation by invertebrates throughout
the monarchs' range.
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Monarch
larva parasitized by tachinid flies (L). Parts of
its body turned brown and transparent shortly before fly
maggots emerged. Tachnid fly maggots (R) leave long
tendrils hanging from the body of their host. |
Parasites are small organisms that complete most or
all of their life cycle within a host, and many are capable of a
high degree of within-host replication. Not all parasites kill their
hosts, but parasites almost always have negative effects on host
survival and reproduction. Many parasites and pathogens are known
to attack insects, including viruses, bacteria, fungi, protozoans,
nematodes, and mites. Several viral and bacterial pathogens can
infect monarchs, including a nuclear polyhedrosis virus and Pseudomonas
bacteria (Brewer and Thomas 1966, Urquhart 1987). Protozoan
parasites such as Ophryocystis elektroscirrha and a microsporidian
Nosema species have also been identified in wild and captive
monarchs (McLaughlin and Myers 1970, Leong et al. 1992;1997, Altizer
and Oberhauser 1999, O. Taylor, personal communication). The infective
stages of most insect parasites must be consumed orally, although
some can invade though pores or membranous joints in the insect
cuticle. Many researchers are currently exploring the role of parasites
and infectious diseases in regulating insect population size (E.G.
Faeth and Simberloff 1981, Bowers et al. 1993, Jaenike 1998).
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A
fifth instar larva (L) showing signs of bacterial decay
shortly after death. The nuclear polyhedrosis virus
that attacks monarchs has been called the "black
death" (R) because infected larvae and pupae turn
black and disintegrate following infection. |
continue to Background
and Life Cycle of Ophryocystis elektroscirrha
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