Strategies for
Survival
Page 3
1 - Mate recognition
2 - Camouflage & Disguise
3 - Aposematic
& Diematic
colouration
4
- Mimicry
5
- Sexual dimorphism
6 - Roosting behaviour
7 - Seasonal dimorphism
8 - Chemical warfare
Aposematic
colouration
Altinote dicaeus callianira - it's
distinct pattern advertises it's unpalatability to avian predators
Aposematic colouration is a term used to describe colours and / or
patterns that act as a warning to predators that a potential prey
species is unpalatable, toxic or dangerous.
Various studies have shown that all vertebrates including
insectivorous birds associate greens and blues with safety, and
inherently regard red, orange, yellow and white as signs of danger.
It is also widely accepted that patterns incorporating stripes or
spots draw attention to objects. Consequently it is no surprise to
find that toxic or unpalatable butterflies have evolved colour
schemes that reflect these facts, in order to "label" themselves as
being unpleasant to eat, and thereby dissuade birds from attacking
them. Equally it is unsurprising that a significant number of
palatable species have evolved which mimic the patterns of toxic
species, in order to trick birds into leaving them alone.
Birds can
remember the colours and patterns of butterflies, and associate
them with pleasurable or unpleasant experiences. If a bird pecks at
a toxic butterfly it finds the taste very unpleasant, and is likely
to suffer consequences including vomiting, nausea and visual
disturbance. Experiments with various insectivorous birds have shown
that if they suffer this experience they then avoid eating similarly
coloured butterflies for several hours or days. Periodically they
re-sample aposematically coloured butterflies, thereby reaffirming
their unpalatability.
Diematic patterns
Many
butterflies and moths are
marked with conspicuous ocelli
- "false eyes" that can scare off a predator, or at least deter it
long enough for the insect to make it's escape. The ocelli make
the insect appear larger and scarier to predators. In many cases
the ocelli have the effect of simulating the face of a small
mammal or reptile.
Markings that have the effect of startling or frightening
potential predators are known as diematic patterns. There are
hundreds of examples among the Lepidoptera, including
Automeris Bullseye moths,
Smerinthus Eyed Hawkmoths,
Mesosemia Metalmarks and the Peacock
butterfly Inachis io.
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In a study by Stockholm University, the ocelli of 20 Peacock
butterflies were blanked out with a marker pen. When exposed
to blue tits, 13 of them were attacked and eaten. A
control group of 34 Peacocks with intact ocelli fared
much better - only a single butterfly was attacked.
It can be concluded that in 97 percent of encounters with
blue tits, the ocelli are effective as a deterrent, and
the butterfly will escape unharmed.
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Inachis io
-
very
beautiful to human eyes, but frightening to a
small bird.
The neotropical
Bullseye moths such as
Automeris liberia, keep their
ocelli hidden most of the time beneath their cryptically patterned
"dead leaf" forewings. If disturbed they immediately drop to the
ground and twitch rhythmically, drawing maximum attention to the
ocelli. The appearance is quite fearsome, and more than enough to
deter a bird from attacking.
Automeris liberia, Ecuador
( image
©
Steve Ife )
Diematic mimicry is quite a common form of defence in
caterpillars as well as in adult butterflies and moths. The larvae
of many Swallowtail species including
Papilio polymnester and Papilio
troilus have a pair of false eye-spots on the thoracic
segments. Many hawkmoth larvae such as
Deilephila elpenor and Hippotion
celerio employ the same strategy.
When alarmed the larvae of these species puff up the thoracic
segments causing the eye-spots to expand. This is considered to be
a form of diematic defence in which the larvae are mimicking the
heads of snakes.
Snake mimicry is also found in adult Atlas moths in the genera
Attacus and
Rothschildia. In these species the forewing apex is lobed,
and bears markings corresponding to the eyes and mouth of a snake.
The illusion is enhanced by rhythmic movements of the wings which
draw attention to the snake-head markings.

Snake-head markings on apex of
Giant Atlas moth
Attacus atlas
( image
©
Gan
Cheong Weei )
Decoys to distract
birds
Often the frightening effect of diematic markings is only temporary.
Having gotten over the initial shock a bird may resume it's attack.
In such circumstances the ocelli on the butterfly's wings take on a
secondary defence role, diverting
and confusing the predator :
When a bird
attacks a butterfly it naturally focuses it's aim on an obvious
target area.
The presence of a decoy target such as a false eye diverts the attack away from the
butterfly's body, towards the wing borders. Usually this results
in nothing more than a small chunk being pecked out of the wing, and
the insect escapes virtually unharmed.
It is not unusual to find ocelli-marked butterflies which have had
chunks pecked out
of their wings by birds. Butterflies are able to fly and go about their lives with quite
large pieces
missing from their wings, but an attack on their body would be
fatal.
Sometimes the ocelli are huge, as in the neotropical Owl butterfly
Caligo
teucer, but even the small ocelli on
species such as the Gatekeeper Pyronia tithonus are enough to divert a bird attack away
from the butterfly's body.

Caligo
teucer,
Peru
The Zebra Teaser
Arawacus separata and many other
butterflies in the subfamily Theclinae have wings that are
marked with bright streaks which lead the eye away from their head, and
towards antennae-like tails on their hindwings. In many species there is also a red or black spot near the tail,
which simulates an eye.
The overall effect is to create the illusion of a "false head", and to give the butterfly a
back-to-front appearance. This is further enhanced by the
butterfly's habit of immediately turning to face the other way as
soon as it lands on a flower or leaf. It is also likely to dip it's
real head, and raise the false head. Periodically, it oscillates the
hindwings, causing the false-antennae tails to wriggle.
An attacking bird always tries to anticipate the escape route of it's
prey, so it aims it's attack at a point fractionally in front of the
head. The false head fools the bird into aiming behind the butterfly
instead. The butterfly then darts off in the opposite direction to
that which the bird expects, and makes it's escape.
Arawacus
separata
-
the
stripes give a back-to-front appearance, and false antennae act
as decoys
Patterns to confuse
Many butterflies have patterns which at first
glance seem to serve no purpose. What for example would be the point
of a striking and easily remembered pattern such as the chequered
appearance of the Swallowtail
Papilio machaon ?
Observing butterfly behaviour often
reveals the answer to such riddles. The Swallowtail normally
rests with it's wings closed, but if it is disturbed it suddenly
flicks them open in exactly the same manner as adopted by the
Peacock and other ocelli-equipped species such as Bullseye
Silkmoths or Eyed Hawkmoths. Furthermore, when alarmed the butterfly
often moves the outspread wings in a jerky and almost threatening
motion, as if to deliberately draw attention to itself.
It seems likely therefore that the pattern
acts either to make the butterfly appear too large to eat, or that
it simply confuses the bird - causing it's eyes
to wander all over the pattern while the bird tries to fathom out
what it all means - Is it edible ? Is it dangerous ? Is it small
enough to eat ? Which bit of it should I aim my beak at ? The bird
may be so confused that it decides to abandon the attack, or the
attack may simply be delayed long enough to allow the butterfly to
escape.
The purpose of the crimson ocelli on the hindwings remains a
mystery. They are not big enough to be scary, and their positioning
close to the vulnerable body effectively rules out any possibility
that they act as decoys or targets.

The Swallowtail Papilio machaon flicks
it's wings open in a threatening manner if alarmed
Signalling danger to other butterflies
Warning colouration is generally assumed to be
targeted directly at birds and other predators, but in the case of
Panacea prola from Peru the purpose is quite
different :
Large groups of males gather to feed on
mineralised moisture on riverbanks. They bask when feeding,
displaying their metallic blue uppersides, which help other passing
males to locate them and home in on the feeding grounds.
A group of butterflies on the ground could easily
be attacked by birds however, so the butterflies need to employ a
defence strategy. Any kind of minor disturbance causes one or two
butterflies to get twitchy, at which point they start fanning their
wings nervously so that the bright red undersides are visible to the other
butterflies.
Increased levels of nervousness cause them to fan
their wings faster, which attracts the attention of further basking
males, which join in the fanning activity. This "red
for danger" signalling is a very
effective alarm system. It quickly alerts all
Panacea in the vicinity to any
perceived threat, and allows them time to take
evasive action before the threat becomes severe.
Panacea prola
basks
while feeding on mineralised water on a Peruvian riverbank.
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Panacea prola
group wing-fanning to warn each other of potential danger. |
Flash
colouration
Many butterflies have a
"now you see me, now you don't" alternating display of a bright
upperside and sombre underside. This is known as flash colouration.
An example is the
South American
Morpho
helenor which has a
brilliant iridescent blue upperside that makes it highly visible to
predators as well as to potential mates. If alarmed, the butterfly
will immediately land, snapping it's wings shut so that only the dark
brown underside is visible. After landing however there is always
the possibility that it might be spotted at rest by a pursuing bird,
so then the secondary decoy-ocelli defence may help the butterfly to
escape by diverting the birds beak away from the butterfly's body
and towards the wing edges.
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Morpho
helenor |
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