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The Study of Butterflies
4. Defences and Defensive Behaviour
Peter Smetacek
In this part we look at some strategies adopted by butterflies
Peter Smetacek works on
the taxonomy and zoo- that enable them to avoid predators.
geography of Indian
Lepidoptera. He is also Butterflies are a fine example of how creatures that are perfectly
interested in exploring non-aggressive can survive in nature. Indeed, they are not only
the potential of Lepi- surviving, but are also thriving. These fragile-looking, shortlived
doptera as bio-indicators
creatures have been around for millions of years, much longer
of ecological functions
and groundwater. than humans, or even the genus Homo: 35 million year old
butterfly fossils from Europe and North America all bear the
distinguishing characters of modern-day butterfly families, so
ancestral butterflies must have evolved much earlier than that.
Part 1. The Naming of Indian But-
terflies, Resonance, Vol.5,
Butterflies are found almost everywhere that it is possible for a
No.6,pp.8-14, 2000. creature to survive, from the harsh and cold Arctic tundra to hot
Part 2. Flight, Fuels and Senses, deserts like the Thar and Sahara and, of course, in very large
Resonance, Vol.5, No.8, pp.4-12, numbers in equatorial rain forests.
2000.
Part 3. Intra-specific Variation,
No butterfly can bite, sting, cause an itch, squirt acid or spring
Resonance, Vol.6, No.5, pp.8-15,
2001.
similar unpleasant surprises on would-be attackers. This is one
of the major reasons why they are among the most attractive
insects to us. The other reasons are obviously their large size,
wonderful colours, and graceful flight. One would expect that
creatures other than humans would also find them attractive for
the same reasons, but with a sinister rather than appreciative
purpose in mind. If butterflies were entirely defenceless, they
would probably be more or less extinct by now. In reality, they
are thriving, so they must have a few dissuasive tricks up their
sleeves, or rather on their wings. These tricks are mainly passive
defences, of which butterflies have an impressive array and some
of the better known ones are discussed in this article.
Keywords
Defence, flight, settling and The adult stage of butterflies differs from the early stages in that
distastefulness; in butterflies. it is highly mobile. Due to this mobility, it has to face and outwit
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not only actively foraging predators such as birds but also
predators that lie in wait, such as praying mantises, web-spiders,
flower spiders, frogs and toads. As a result, no butterfly restricts
itself to a single line of defence.
Flight
Flight is the primary mode of progression of adult butterflies.
As with other modes of progression, it is also the primary means
of escape from dangerous situations. A large number of butter-
flies have mastered the intricacies of rapid flight and habitually
zip about. Others have a rather moderate or even slow flight, but
nearly all are capable of considerable bursts of speed over short
distances if they feel the need to escape. Few can keep up the
escape velocity for long and generally slow down as soon as they
feel themselves to be out of danger.
The ability to fly fast requires the development of powerful
flight muscles. These, in turn, require large amounts of energy
to operate efficiently, besides turning the possessor into a spe-
cially dainty morsel, with more flesh than bones (sorry, chitin).
The best examples of the followers of this model of passive
defence are perhaps the Skippers (Hesperiidae), which are Lepi-
doptera intermediate between moths and butterflies. Almost all
the 3000 species of Skippers found worldwide have a stout
thorax housing powerful flight muscles (Figure 1) and habitually
Figure 1. The broad thorax
fly at such a speed that they are quite difficult to follow with the
of this Skipper (Celaenor-
eye. They are very wary and rarely allow one to approach close. rhinus sp.) houses power-
The drawback of this otherwise very safe system is that the ful muscles that enable the
minority of species that are active throughout the daytime need butterfly to fly very fast.
to spend a great deal of their waking hours searching for and (Credit: Peter Smetacek)
sucking up nectar to feed their powerful muscles. The majority
of Skippers are active only for a few hours each day at dawn and
dusk, when they zip about in the gloaming to locate mates. It is
suspected that such species are also active at night (Box 1) but
nothing is known about this for certain.
Other groups that habitually fly fast are the Hairstreaks
(Lycaenidae) and some Nymphs (Nymphalidae). Like the Skip-
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pers, the Hairstreaks are rather small butterflies that fly so fast it
Box 1
is difficult to follow them with the eye. They are active during
It has been proposed that, the daytime and, like the diurnal Skippers, spend a good part of
originally, there were no their time locating food and feeding.
day-flying Lepidoptera.
However, as bats per- Among the Nymphs, the Rajahs (Charaxes spp.) and Nawabs
fected their sonar-based (Polyura spp.) are notably swift fliers, with a broad thorax. They
hunting system, pressure are capable of very fast flight but, because of their large size, they
grew on the night-flying are comparatively easier to follow with the eye than Skippers
Lepidoptera and some
and Hairstreaks. Like these two groups, they require large
species evolved to exploit
quantities of food but prefer rotting fruit or crabs, animal
the hours of daylight,
when predator pressure is
droppings and other foul substances to flowers.
comparatively less. In
The second division consists of those butterflies that fly rapidly
support of this proposi-
but not as fast as the Skippers and the Rajahs and Nawabs.
tion is the fact that, com-
pared with moths, there Nevertheless, such butterflies are capable of short bursts of
are relatively few butter- speed that can match those of the Skippers. Most of the Swal-
fly species. Nor does the lowtails (Papilionidae), Nymphs, Whites (Pieridae), Blues and
diversity of physical Coppers (Lycaenidae) fit in this category. Their flight is moder-
structures found among ately swift and even a hint of danger is enough to cause a burst of
butterflies match those speed that will leave most pursuers behind.
found among moths, sug-
gesting moths are a more This is the middle path in the trade-off between powerful
ancient group. Even to- muscles and the ability to live off relatively scarce food re-
day, predator pressure is sources. While members of the second division are avid feed-
much greater for night fly-
ers, they do not have as stout thoraxes as Skippers and Hair-
ing Lepidoptera, for
streaks. They spend a good part of their waking hours flying and
though birds fill the role
most are capable of sustained flight, unlike the very fast fliers
of predators during the
day, most birds capture that tend to settle frequently.
resting butterflies and
The third approach to safe flight is practised by the Tigers,
there are very few species
of birds worldwide that
Crows, Costers, Windmills, Roses, some Browns and some Whites
habitually capture flying that habitually fly slowly and are capable of only moderate
butterflies, unlike bats bursts of speed even when threatened. While the Tigers, Crows,
which routinely predate Roses, Windmills and Costers deliberately fly slowly to adver-
upon flying insects. tise their distastefulness, the Browns and Whites depend on
erratic flight to escape attacks (Figure 2). The hopping flight of
some Browns, which proceeds in a series of bounds or hops, with
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the wings closed over the thorax in the upper half of the hop,
enables them to move through dense vegetation where pursuit is
impossible. Many Browns feed on grasses or bamboos and they
are consequently commonest in areas where these plants grow.
They are the only butterflies capable of getting through bamboo
clumps undamaged and, when disturbed, will often fling them-
selves into the middle of a handy bush or bamboo clump where,
if followed, they will display equal dexterity in making their way
to the other side of the bush or clump, leaving the cause of Figure 2. Cabbage Whites
(Pieris spp.) have devel-
disturbance struggling to get through!
oped erratic flight to a fine
The Whites, especially Cabbage Whites, species of which are art. (Credit: Rajani Smetacek)
found throughout Europe, N Africa, N Asia, the Himalaya and
the Nilgiri and Palni hills, have mastered the art of erratic flight.
Seen from a distance, they appear to be progressing in a single
direction, but the lurches, dips, hops and spurts that make up
the flight of these creatures make it difficult to believe that the
creature is not thoroughly intoxicated. Such flight, though
ludicrous at one level, is a masterful feat at another, since it
enables the butterfly to dispense with powerful muscles and the
fuel required to power them, while permitting relatively safe
passage through hostile territories.
Certain species of birds, such as crows and seagulls, are known to
Erratic flight, though
enjoy utilising upcurrents of air, rising effortlessly with wings
ludicrous at one
outspread and diving down again from a height, to repeat the
level, is a masterful
process over and over again. At least two species of butterflies,
feat at another,
the common mime (Papilio clytia) and the Blue Apollo (Parnassius
since it enables the
hardwickei) do more or less the same thing. The former butterfly
butterfly to dispense
occurs at low elevation, occasionally ascending to 2250 m in the
with powerful
hills but commonest below 1000 m while the Apollo is most
muscles and the
often found above 3000 m in the Himalaya. Numbers of each
fuel required to
species congregate in some steep ravine or on a spur where there
power them, while
is a strong upward breeze and practise their aerobatics for hours
permitting relatively
at a time, sometimes jousting with others of the same species, at
safe passage
other times chasing other species of butterflies, but always
through hostile
returning to rise upward with outspread wings and dive down to
territories.
the bottom again, to repeat the process. These butterflies are
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The very existence usually males and the Mime is known to await mates on hilltops
of a species is and ridges, as several other butterfly species do.
proof of its
members ability to
Settling
survive.
After high speed travelling which is comparatively safe, there
comes a time when the insect has to settle, whether to eat, sleep,
bask or merely rest. This is the time all the slower predators wait
for, when they can creep up to, leap upon or otherwise lay jaws or
claws upon the elusive morsels. However, the very existence of a
species is proof of its members ability to survive and nothing
that has survived is going to allow itself to be an easy prey every
time it settles.
Most of the Swallowtails rarely settle for long during the day:
instead, they perch briefly upon a flower or hover over wet sand,
their wings trembling nervously, ready to carry them to safety at
a moments notice. If the weather turns chilly they settle down,
usually under a twig or leaf and hang there until a warmer
period. They also pass the night in this manner.
In very few cases, as in the Snow Apollos (Parnassius spp.) and
the Swordtails (Pathysa spp.), Swallowtails settle on a flower as
other butterflies do without hovering nervously about and, with
wings outspread, drink their fill of nectar. During the daytime,
many Skippers habitually settle under leaves with their wings
Figure 3. Most Blues settle outspread and pressed flat against the leaf, making them invis-
so that they cast almost no ible from all sides except below. Since they usually settle on low
shadow, like this white-
growing bushes and herbs, the view from below hardly matters.
spotted Hairstreak (Euaspa
ziha). (Credit: Peter Smetacek) While the majority of butterflies settle facing forward, some of
the Hairstreaks settle facing the direction they came from,
turning sometimes in mid-air just before they settle, at other
times turning around soon after settling. The strategy behind
this will be discussed in a future instalment of this series.
A rather curious habit has been observed among the Blues and
Hairstreaks (Lycaenidae). Soon after settling, they move about
to align themselves with the Sun in such a manner that they cast
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a shadow consisting of no more than a line (Figure 3). If the Sun
is at an angle to their perch, they will even lean over to get the
angle right. The reason is probably that, in Nature, predators
such as birds tend to recognise the silhouettes of their prey. A
shadow is often as good as a silhouette, hence the need to reduce
it to a minimum. Secondly, butterflies absorb heat from the
substratum. It is possible that one side in shadow and the other
in the Sun is not the ideal position for basking. However,
butterflies of almost all other families do not seem to mind
casting a shadow.
In the case of the Evening Browns (Melanitis spp), who resemble
a dry leaf on the underside (see Resonance, Vol. 6, No. 5, p.11,
2001), a common response when they are disturbed or feel
threatened is to dash off among the undergrowth and, settling
among dry leaves on the ground, to lean over until they are
nearly horizontal. By doing this, they look even more like dry
leaves than if they were to remain erect. When they are not
disturbed, though, they settle with the wings erect. The Banded
Satyrs (Aulocera spp) of the Himalaya also lean over sometimes
so that their wings almost touch the ground. They are crypti-
cally patterned on the underside. Being mainly residents of
grassy hillsides, they do not resemble dry leaves but their cryptic
pattern makes them quite difficult to spot when they settle
among tussocks of grass and gravelly paths. The Oakleaf butter-
flies (Kallima spp.), which are perhaps the best example of
camouflage are found in forested hilly areas over most of India.
They perfectly resemble the shape and colour of dry leaves and
are almost impossible to spot when they sit amid genuine dry The Oakleaf
leaves. They, however, do not lean over as do the Evening butterflies perfectly
Browns. When disturbed, they dash off and plunge into thick- resemble the shape
ets, settling with head downwards among dense foliage. the tip and colour of dry
of the lower or hindwing touching the twig on which the leaves and are
butterfly is perched so that one gets the impression that the leaf almost impossible to
is attached to the twig. Some other leaf mimicking butterflies spot when they sit
also do this when disturbed. Since few other butterflies do this, amid genuine dry
i.e. touching the twig they are settled upon with the tip of their leaves.
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What immortal hand hindwing, one is left with the suspicion that there is more to
or eye prompted the the whole business than meets the casual, or even discerning
development of a eye. Why, for one, do they touch the wingtip to the twig only
translucent window when they have been disturbed and are in hiding and not at
in the centre of the other times? How do the Oakleaf butterflies always head for a
upper or forewings, tree whose leaves they resemble when they need to hide?
that enables light to What immortal hand or eye prompted the development of a
pass through thereby translucent window in the centre of the upper or forewings,
breaking up the that enables light to pass through thereby breaking up the
outline of the outline of the butterfly? The genetic code, no doubt, is
butterfly? ultimately responsible for all this and much more, but, is there
a ghost in the machine?
Distastefulness
Of the 20,000 species of butterflies known to science, a rela-
tively small percentage are distasteful to varying degrees. This
means that they store chemical compounds in their tissue and/
or haemolymph (butterfly blood) that may be distasteful or
even downright poisonous for potential predators. Such but-
terflies are known as protected species.
The Crows and Tigers (Danainae) are well-known poisonous
butterflies that contain pyrrolizidine alkaloids. Birdwings and
Red-bodied Swallowtails (Ornithoptera, Troides, Atrophaneura,
Pachliopta) store aristolochic acids.
Earlier, it was believed that such butterflies obtained their
poisons by sequestering them from their larval foodplants, i.e.
the plants the caterpillars feed upon. Therefore, since the Red-
bodied Swallowtails feed on the pipe-vine of Aristolochia fam-
ily of plants (Aristolochiaceae), it follows that they contain
aristolochic acids. Similarly, the poisonous Tigers and Crows
feed on Asclepiads (Asclepiaceae) which are known to be
poisonous. African Costers (Acraeinae), of which two species
are found in India, feed mostly on passion vines (Passifloraceae),
a group of plants known to be poisonous. However, this rather
simple theory was turned on its head as a result of experiments
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with some Costers. It turned out that some members of this sub- Some butterflies
family, of which all members were considered distasteful, were actually
in fact not so, despite the fact that they fed on passion vines. manufacture the
Meanwhile, some among the most poisonous species in the poison they contain
group did not even feed on passion vines but on a plant not in their body tissue,
known to contain poisons. It was therefore concluded that some rather than merely
butterflies actually manufacture the poison they contain in their sequestering them
body tissue, rather than merely sequestering them from the from the larval
larval hostplant. hostplant.
In India, the Yellow Coster (Acraea vesta) of the Himalaya,
which is distasteful, feeds on Debregeasia bicolor and Boehemiria
platyphylla, both members of the nettle family that are relished
by cattle and do not contain poisons. The Common Jester
(Symbrenthia lilaea), which is relished by birds, also feeds on the
same plants. It seems likely from the above that the Yellow
Coster actually manufactures the poisons that protect it.
To complicate matters, experiments with a North American
butterfly, the Pipe-vine Swallowtail (Battus philenor) produced
some unusual results: members of the species that were fed on
leaves of the pipe-vine (Aristolochia) were distasteful while mem-
bers of the same species that were fed on knotweed (Polygonum)
were not distasteful. In this case, it seems evident that the
butterfly cannot actually manufacture poisons but depends on
its larval foodplant for them.
While the ability to store chemicals that are distasteful or even
poisonous confers some advantage to the butterfly, the effects of
the chemicals vary from group to group of predators: birds and
bats may find a butterfly nauseating, but a toad, praying mantis
or spider will hardly notice the difference between a protected or
unprotected species. In other words, the chemical defences are
effective only against certain potential predators and the butter-
Address for Correspondence
fly has to use other means to circumvent the remainder.
Peter Smetacek
Jones Estate
P.O. Bhimtal, Nainital
Uttaranchal 263 136, India.
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