Abstract

The Banded Swallowtail butterfly, Papilio demolion demolion, is characterized by a single light green
band that extends from the forewing apex to the hindwing mid dorsum. It has been hypothesized that
colored band patterns in several butterfly species could have an aposematic or camouflage function but
this has not been scientifically proven in the case of the Banded Swallowtail. In this experiment, we have
tested whether the band in the Banded Swallowtail butterfly functions as an aposematic signal, or
disruptive coloration, or both. Paper models which realistically imitate the wing patterns and colours of
the Banded Swallowtail were created. Paper models which are variations from the wild type, where the
band pattern is distorted, and models with the original colours converted to greyscale, were also created.
These models were then placed in the field, at 3 different locations, each with a live mealworm attached
to it. We measured the predation rates on the paper models and discovered that wild type models were the
least predated on, followed by models with green distorted band patterns, followed by grey scaled models
with the bands still intact, and then lastly, the grey scaled models with distorted band patterns were
predated on the most. We conclude that the band must contain antipredator qualities, and some of these
qualities might have been aposematism and disruptive coloration.

Introduction
Animals employ a wide variety of techniques to protect themselves from potential predators, and one such
technique is the use of body coloration. Protective body coloration comes in various forms and some of
these include aposematism and camouflage. Within camouflage, we have disruptive camouflage and
crypsis. The former normally involves colored patterns placed on the body outlines of the animal, thus
interfering with the correct identification of the animal’s shape, rendering it less detectable by predators
(Stevens and Merilaita, 2011) . The aim of crypsis is also to avoid detection, where animals use their body
colors to blend into the environment as best as they can (Stevens and Merilaita, 2011). Mimicry on the
other hand, is where palatable insects evolve over time to look more similar to noxious or poisonous
counterparts so that predators will not consume them (Stevens and Merilaita, 2011). Finally, there is
aposematism, which is normally a bright colored signal which informs potential predators that the animal
is probably poisonous, thus discouraging predators from consuming it (Santos et al., 2003). It is important
to note that an animal might utilize more than one type of body coloration to best protect itself from
predators. This is best observed in the case of Lataste’s Viper, where scientists have come to conclude
that although aposematic qualities in the species are most apparent, the species also possesses the qualities
of disruptive coloration, and a combination of these two qualities are at work to protect the species from
predators (Niskanen and Mappes, 2005).

As with most antipredator traits, an understanding of the evolution of aposematic coloration gives further
insight as to how this trait has changed and maintained its effectiveness over the course of time. It has
been postulated that over time, predators of aposematic species have come to learn that the bright
aposematic colors are often associated with distastefulness or toxicity in the prey (Santos et al., 2003).
Predators waste little time and effort in concerning themselves with prey possessing such colors, allowing
such prey to survive and the aposematic color trait to be passed down across many generations. There are
two criteria to be fulfilled for aposematism to be passed down across generations - i) the aposematic
signal must be obvious, so much so that the predator avoids the particular prey after as few encounters as
possible, promoting rapid learning (Harvey and Paxton, 1981); ii) the prey possessing the signal must also
be unpalatable (Harvey and Paxton, 1981). However, there is also much uncertainty surrounding the
evolution of aposematism. A study suggests that aposematism in general is likely to have evolved
gradually over time, where mutations over a range of weak signals could accumulate over time under
neutral selection to produce strong signals (Lindstrom et al., 1999). Another study argues that predators
learn to avoid aposematic prey only after killing a few of them, and as a result rare aposematic forms will
always be selected against, making the passing down of aposematic genes an impossible task (Wiklund
and Jarvi, 1982). As such, kin selection is thus proposed as the solution. The fundamental basis of this
theory is built upon the notion that aposematic prey are killed when tasted by predators, but this could
well be an assumption, as the prey could be spat out by the predator upon the first unpleasant taste,
allowing the prey to survive (Wiklund and Jarvi, 1982). It has been shown that most predators such as
birds, reptiles and amphibians often release distasteful prey insects unharmed after tasting them,
effectively contradicting the kin selection theory (Wiklund and Jarvi, 1982).

Several factors determine the effectiveness of an aposematic signal. It is important to note that different
colors vary in their effectiveness to deter different predators (Svadova et al., 2008). In a particular study
done on firebugs and birds, yellow and red colored firebugs were exposed to birds for an equal amount of
time (Svadova et al., 2008). Birds that learnt to avoid red colored firebugs did not apply this learning to
the yellow colored firebugs, resulting in more yellow firebugs being predated, and showing that red was a
more effective signal color than yellow (Svadova et al., 2008). All of the firebugs used in this experiment
were non-toxic and not poisonous, thus removing the possibility that the birds could have learnt to avoid
certain toxic firebugs throughout the course of the experiment. Perhaps this differing effectiveness of
various colors is especially apparent in avian predators, since birds possess UV-A sensitivity and
tetrachromatic color vision (Bennett and Thery, 2007). Among other things, high chromatic contrast
between prey and background, and high brightness or luminance contrast also help to determine the
effectiveness of the aposematic signal (Prudic et al., 2006).

The effectiveness of aposematism is also determined by how certain predators perceive their prey.
Different animals see things differently. For example, there is an innate visual biasedness present in most
birds. In an experiment once again involving birds and firebugs, larger numbers of birds avoided firebugs
with aposematic coloration compared to to firebugs of the normal brown variety from the start of the
experiment (Exnerova et al., 2007). As such, there is evidence to show that birds might prefer or not
prefer certain colors, and this biasedness might have been present from their birth. This leads to the
question as to whether other predators could have some sort of visual biasedness as well, or might
perceive prey differently from each other. Lizards, which are common butterfly predators, have been
shown to be able to recognize a very wide range of colors, and this includes recognition of aposematically
colored insects (McGovern et al., 1984). However, we have amphibian predators and praying mantids
which make predatory decisions mostly based on prey movement, with prey color being low on their
priorities (Hatle and Salazar, 2000). As such, it can be observed that the effectiveness of aposematism
will vary across different groups of predators.

Natural selection is a crucial factor in deciding which anti-predator traits are passed down, and
camouflage is one such trait (Schaefer and Stobbe, 2006). Camouflaged individuals which are less likely
to be found, are thus more likely to survive and reproduce, propagating its genes in the next generation
(Evans and Schmidt, 1990). Disruptive coloration is a form of camouflage, where the scattering of
colored patterns on an animal’s body breaks apart its outline (Cuthill et al., 2005). The evolution of this
particular trait, and whether or not it is successfully passed down from one generation to the next, is
highly dependent on changes in visual perception of key predators and also changes in the immediate
background (Beatson, 1976).

Disruptive coloration is most effective when the high-contrast markings are placed on the animal’s
outline, but more current studies have now shown that these markings have even greater effectiveness
when placed further away from the body outline, creating effective surface disruption (Stevens et al.,
2009; Cuthill et al., 2006). Any patterns and colorations on the animal should serve to promote
concealment, and highly contrasting colors should serve this purpose greatly (Cuthill et al., 2005).
Through detailed studies on cuttlefishes, it has been discovered that size, contrast and density of patterns
affect the disruptive strength of the patterns in varying amounts (Chiao et al., 2007). These studies also
tend to suggest that Weber Contrast (color contrast against the background) of the patterns, combined
with their size make for the best disruptive signals (Chiao et al., 2007). However, studies have also
emerged to show that disruptive coloration can be just as effective regardless of the background. A study
stated that moths which were different from their background but sported disruptive patterns were
significantly less predated on than cryptic moths which looked similar to their backgrounds (Schaefer and
Stobbe, 2006).

Colour patterns of butterflies can serve various functions other than predator deterrence such as sexual
selection. Combinations of coloration aspects in butterflies have been studied in Heliconius erato
butterflies. It is believed that the color patterns in these butterflies are important for both predator
deterrence and mate selection (Finkbeiner et al., 2014). They are predominantly black in color, with wild
splashes of red and white on various sections of the forewing. The results seem to suggest that the color
patterns on the butterfly seem to contribute quite equally to aposematic and mate attraction purposes,
showing that colored patterns can serve different functions at the same time. Papilio butterflies are among
the flashiest and most vibrant of butterflies, usually sporting bright colors and patterns on top of a black
background (Kunte, 2008). One of the distinct characteristics of the Banded Swallowtail is that it
possesses is a single straight horizontal light-green band that stretches across the butterfly. We
hypothesize that this band could have qualities of disruptive coloration, or aposematism or a mix of both.
We also ask if there could be other elements of crypsis present, where the butterfly is using the green
color of the band to blend into its surroundings.

More recent studies show that disruptive patterns can be just as, or if not more effective when placed not
at the outlines of the body, but on the middle sections (Cuthill et al., 2006). This suggests that even
though the band pattern on the Banded Swallowtail is found in the middle of its body, it still has a good
chance of possessing disruptive properties. There are also two important aspects of effective camouflage.
Any patterns and colorations on the animal should serve to promote concealment, and highly contrasting
colors should serve this purpose greatly (Cuthill et al., 2005). Once again, since the bright greenish band
of the Banded Swallowtail lies on top of a black background, this conforms with the earlier mentioned
notion, providing a little more evidence for disruptive coloration in the butterfly.

This project aims to discover if there is also a combination of coloration aspects at work in the Banded
Swallowtail butterfly, and whether aposematism and disruptive coloration are working hand in hand to
deter predation. In order to test the true functions of the colored patterns found on the Banded
Swallowtail, we have utilized paper models bearing different modifications to the original wing patterns
and colorations. Four different types of models were produced, and these included models of the wild
type, a grayscaled version of the wild type, a version with wing patterns greatly distorted, and finally one
last version which was a grayscaled version of the distorted model. We then placed these models in the
field to observe predation on them. Artificial paper models have been used in other studies involving
predation and mate preferences to great effect, showing that that these models are effective imitations of
their real counterparts (Diggins et al., 2004). Through this study, we hope to elucidate the function of the
colored band of the Banded Swallowtail butterfly. We have modified the band in several ways, distorting
the original horizontal straight band structure and also removing the original light green colour. Through
this, we aim to discover whether it has aposematic qualities, or functions as disruptive camouflage
helping to break up the outline of the butterfly, or possesses a combination of both of these qualities.

Materials and Methods

(a) Experimental specimens

Preserved specimens of the Papilio demolion demolion (Banded Swallowtail butterfly) species were
obtained via Ebay from Andreas Muller, an insect collector located at Oberhaft 33, A-5221 Lochen,
Austria. Upon obtaining the specimens, the wings were cut off and spectral measurements were done on
the forewing sections.

(b) Preparation of paper butterfly models

When the Banded Swallowtail is at its resting position, its wings are outstretched and extend fully (Tan,
2012). The models were produced to best imitate the butterfly at its natural resting position. A high
definition photograph of P. demolion was edited with Photoshop CC 2014 so that its size would match
that of a real specimen. The same program was used to modify the wing colors and patterns on the
forewing sections. Each image was then mirrored and duplicated so that the left and right wings would be
identical to one another. After the images and designs were finalized, they were printed on a, HP Deskjet
2540 printer with HP61 ink, on HP printer paper. The models were then soaked and covered with paraffin
wax to render them resistant towards bad weather conditions. In order to ensure that the colors of the
models were similar to real butterfly specimens, spectral measurements of the finalized waxed models
were taken using an Ocean Optics USD2000 fiber optic spectrometer and compared with those of the real
specimen (Figure 1a, 1b). All measurements were taken with the axis of the illuminating and detecting
fibre directed at a 90° angle to the plane of the wing at a distance of 2mm using a deuterium-halogen
tungsten lamp (DH-2000, Ocean Optics) as a standardised light source and calibrated using a white Ocean
Optics WS-1 reflectance standard.

We produced 4 different type of paper models: models looking similar to the wild type (Figure 2),
grayscaled versions of the wild type (Figure 2), models with the wing patterns distorted (Figure 2), and
finally grayscaled versions of the distorted models (Figure 2). Using predation levels on the wild type
model as the baseline, we tested whether varying colors and wing patterns would lead to lower or higher
predation levels. The grayscaled wild type tested if the straight horizontal band was an effective
disruptive pattern as any possible aposematic qualities given by the original light green color would have
been removed. The wild type model with distorted wing patterns tested whether the light green color on
its own, without any possible disruptive qualities given by the straight band, was still a good enough
deterrence tool against predators, thus possessing aposematic qualities. The grayscaled model with
distorted wing patterns acted as the control against which all the other variants were compared, since all
possible aposematic or disruptive coloration functions were removed from it. The combination of
different predation levels on each model would tell us about the effectiveness of the light-green color and
the straight band pattern in deterring predation, and also whether aposematism or disruptive coloration, or
a combination of both are at work to protect the Banded Swallowtail.
Figure 1a. Plot of black reflectance spectra of wildtype and artificial Papilio demolion forewings

Figure 1b. Plot of green reflectance spectra of wildtype and artificial Papilio demolion forewings
 Figure 2. Butterfly models that were used in the experiment.

(c) Model preparation

A mealworm larva, Tenebrio molitor, was attached to each paper model. Mealworms have proven to be
more effective as bait compared to other choices such as pastry, clay or plasticine, showing higher attack
rates than the latter options (Ho et al., 2016). They were placed on the middle, underside area of each
model, firmly stuck between two pieces of Blu-Tack. To prevent the mealworm larva from being attacked
by ants and other crawling insects which were not natural predators of the Banded Swallowtail, the model
was elevated with a piece of Blu-Tack placed on the underside of the left wing. This piece of Blu-Tack
had been sprayed with insecticide (Baygon Multi-Insect Killer) beforehand.
The mealworms used in our study are of the brown and not the red variety, bearing in mind the earlier
mentioned point on how certain predators, especially birds could be aversive to the latter.

(d) Field sites and experimental setup

All predation experiments were carried out at locations where the Banded Swallowtail could be naturally
found, and these included Kent Ridge Park (01°17' N, 103°46' E), Tengah forest (01°21’N, 103°43’ E)
and Macritchie Reservoir Park (01°20 N’, 103°49’ E). These locations contained a myriad of avian and
insect predators such as the Forest Wagtail (Dendronanthus indicus), the Javan Mynah (Acridotheres
javanicus) and the Greater Coucal (Centropus sinensisby).

Models were placed at each site on different dates, between December 2016 to February 2017. 100
models, 25 of each type were placed at the Kent Ridge Park location. 200 models, 50 of each type were
placed at the Tengah forest location. Lastly, 360 models, 90 models of each type were placed at the
Macritchie Reservoir Park location. At each location, the models were placed about half a metre apart
from each other, with no more than 3 models on the same tree or plant. Models were left in the field for
up to 4 days and checked for predation each day. A model was considered to have been preyed on if the
mealworm attached to it was partially or fully consumed.

(e) Statistical Analysis

To test for differences in predation across all the different models placed at the various sites, survival
analysis of the models was performed using the OASIS 2 (Online Application For Survival Analysis 2)
program (Han et al., 2016). Within this program, the Kaplan-Meier estimator, which is a non-parametric
statistic, was used to estimate the survival function (probability that an object of interest survives beyond
a specific time) of the models. The mean and median lifespans of each model, and also their
corresponding survival curves at each location were obtained using OASIS 2 as well. Probability values
were obtained from comparing the predation levels of the models against each other.

A chi-squared test was done on the number of surviving models after 3 days at each site. This was
performed using R statistical software (R Core Team, 2012).

A Generalized Linear Mixed-Effects Model (GLMM) analysis with a binomial distribution, logit link
function, followed by post-hoc pair-wise comparisons (Tukey corrected) were done on the predation
results at all 3 locations. These were also performed using R statistical software (R Core Team, 2012).

Results
Of all the models placed in the field, it can be observed that the grey distorted models fared the worse and
showed the highest levels of predation (Figures 3a to 3c). On the other hand, wild type models fared the
best and survival rates hovered around 80 percent at the end of 3 days for these models at all 3 site
locations (Figures 3a to 3c). Green distorted models fared significantly better than grey banded models at
Kent Ridge Park and Tengah forest, but not at the Macritchie site, where percentage survival of the
former models were either similar or lower than that of the latter models. At Kent Ridge park, 9 green
distorted models and 15 grey banded models were predated on. At Tengah forest, 20 green distorted
models and 22 grey banded models were predated on. However, at Macritchie Reservoir Park, 38 green
distorted models and 36 grey banded models were predated on.

We observe that the difference in predation on the wild type models and grey banded models was
significant (p<0.05), with the latter predated on significantly more than the former, at Kent Ridge Park
but not at Tengah and Macritchie sites (Table 1). Across all 3 sites, wild type models were preyed on
significantly less than grey distorted models (Table 1). Grey banded models were significantly less
predated on than grey-distorted models at Macritichie Reservoir Park (p=0.0448). Also, green-distorted
models were preyed on significantly less than grey-distorted models at Kent Ridge Park (p=0.0114).

P-values are not small enough to conclude whether wild type models were significantly less preyed on
than green-distorted or grey banded models; or whether grey banded models were significantly less
preyed on than green-distorted models.

Results from the Chi-square test on the survival of models at the end of the 3 day experiment at each site
indicate that at all 3 locations, differences in number of surviving models were not significant.

Table 1. X2 and P-values of the models versus each other at each site location
Kent Ridge Park Tengah Macritchie Reservoir
Park

Conditions x2 P-value x2 P-value x2 P-value

WT v.s. greybanded 6.9200 0.0085 2.6700 0.1026 0.3300 0.5659

WT v.s. greendistorted 1.0100 0.3142 3.2900 0.0697 1.5100 0.2197

WT v.s. greydistorted 12.0200 0.0005 10.3800 0.0013 6.7000 0.0096

greybanded v.s. greendistorted 2.6800 0.1014 0.0600 0.8041 0.4200 0.5168

greybanded v.s. greydistorted 0.8000 0.3710 2.7800 0.0955 4.0300 0.0448

greydistorted v.s. greendistorted 6.4100 0.0114 1.8200 0.1776 1.7300 0.1878

Figure 3a. Kaplan-Meier survival curve of models at Kent Ridge Park

Figure 3b. Kaplan-Meier survival curve of models at Tengah site

 Figure 3c. Kaplan-Meier survival curve of models at Macritchie Reservoir Park

(a) Mean lifespans as an estimate of best predator deterrence

Using mean lifespans as an estimate for survivability, we observe that the wild type models had the
longest mean survival across all 3 locations and grey-distorted models have the shortest (Table 2).
However, it is important to note the differences in mean lifespans among models are not statistically
significant. We also observe that grey banded models had a longer mean lifespan at Tengah and
Macritchie sites as compared to green distorted models but not at Kent Ridge park.

Mean lifespans of all the models were also considerably different at each of the 3 different location sites
(Table 2).

Table 2. Mean lifespan at the various site locations for each of the different model types
Mean Lifespan at Kent Ridge Mean Lifespan at Tengah Mean Lifespan at Macritchie
(Days) (Days) (Days)

Wildtype 2.72 2.8 2.88

Greybanded 2.35 2.66 2.81

Green-Distorted 2.59 2.54 2.72

Grey-Distorted 2.19 2.46 2.7

 (b) Generalized Linear Mixed-Effects Model Analysis results
There are statistically significant differences in predation levels of model types. The wild type models had
significantly higher survivability than the grey distorted models, and grey banded and green distorted
models were preyed on significantly less when compared with the grey distorted model (Table 3).

Across all three locations, the wild type models had the lowest probability of predation (p=0.225),
followed by green distorted having the next lowest probability of predation (p=0.348), then grey banded
models (p=0.384), and finally the grey distorted which have the highest probability of predation
(p=0.562).

Table 3. Generalized Linear Mixed-Effects Model analysis results

Discussion
For this study, we tested whether the light green band found on the Banded Swallowtail butterfly
possessed any antipredator qualities, and whether aposematism or disruptive coloration could have been
some of these qualities. Our results clearly show that the straight, horizontal light green band of the wild
type model has been greatly effective in deterring predation. It is also apparent from the results that every
component of the band matters. Both the straight horizontal structure of it, and the light green color of it
are crucial components in terms of predator deterrence. Each of these two components have different
possible antipredator qualities associated with it. The effectiveness of these two components are relatively
similar, as shown by the similar predation levels in both the green distorted and grey banded models.

Regarding the light green color associated with the band, we suggest that there could be aposematic
qualities associated with this color. There are a variety of ways to argue for aposematism, and the first is
to rely on the fact that aposematism depends on the prey in question being either toxic or distasteful
(Santos et al., 2003). There appears to be no scientific literature explicitly stating that the Banded
Swallowtail is either toxic or distasteful. Also, while in the larval stage, the larva feeds on the leaves of
mainly 3 different host plants, and these are the Luvunga crassifolia, Melicope luna-ankenda and Citrus
spp. (Tan, 2012). These plants belong to the Rutaceae family and are all non-toxic, giving further reason
to believe that the adult form of the butterfly is non-toxic as well. However, at this point, we still cannot
be absolutely sure that the adult Banded Swallowtail is neither toxic or distasteful. It is entirely possible
for distasteful and aromatic substances to be sequestered from harmless plants by butterflies in the adult
stage (Smetacek, 2006). The larvae of 3 species from the Papilio family, namely polyctor, polytes, and
protenor all feeding on plants belonging to the Rutaceae family too, just like the larvae of the Banded
Swallowtail butterfly. It has also been proven that the adults from the earlier mentioned 3 species are
significantly distasteful to most birds (Smetacek, 2006). As such, there is reason to believe that the adult
Banded Swallowtail butterfly might be distasteful as well, and the light green color of the band might
have developed over time to signal this distastefulness to predators.

Aposematism is also most relevant when dealing with predators which have the ability to perceive a wide
range of colors. Birds are common predators of butterflies and these animals possess highly advanced
visual systems. Simply put, humans possess trichromatic color vision which is based on 3 different
photoreceptor types:red, green and blue cones, while birds possess tetrachromatic color vision, having an
additional photoreceptor type, meaning that birds might be able to better discriminate between different
colors compared to humans (Hill and McGraw, 2006). As such, there is a good chance that the avian
predators in our experiment were able to discriminate between grey and light green model types. Wild
type and green distorted models in the experiment were preyed on less compared to other grey model
types, indicating that avian predators might have had a preference for grey colored models and tended to
avoid the light green models. There is thus a possibility that the light green color has evolved and came to
be over time as a means of deterring avian predators.

We suggest that another quality the light green band might have, is that of disruptive coloration. In this
case, both the straight horizontal structure and light green color of the band must be taken into account. In
the case of disruptive coloration, contrasting markings tend to cancel each other out by their separate and
conflicting patterns (Thayer, 1909). The outline of the animal is broken up against both light or dark
backgrounds, with light failing to show against light, and dark failing to show against dark (Thayer,
1909). It appears that the light green band of the Banded Swallowtail fits this definition of disruptive
coloration, with the light green color of the band resting on top of a black background. As such, the band
adheres to the ‘color contrast’ aspect of disruptive coloration, but structure is also of paramount
importance. In quite a number of animals such as leopards and giraffes, disruptive coloration has taken
the form of relatively random spots scattered throughout the animal’s bodies while in the case of zebras, it
has taken the form of relatively organized, vertical black and white bands on the animal’s body. There is
currently no literature explaining whether more ‘organized’ or random arrangements of patterns, and also
whether certain shapes are more effective in deterring predation. The band patterns in snakes have been
studied to a great degree, and studies have shown that stripes and bands have been significantly effective
forms of disruptive coloration in these snakes, aiding in concealment while these snakes forage
(Brattstrom, 1955). Also, a single horizontal black band cuts across the body of the Papilio demodocus
caterpillar, breaking apart the overall structure of it and rendering it less visible to predators (Evans and
Schmidt, 1990). Arrangement and structure of patterns could affect their effectiveness as a form of
disruptive coloration. This could give us some insight as to why the wild type models did better than the
green distorted models, and also why the grey banded models did better than the grey distorted models in
our experiment, since both of the former models possessed the straight horizontal band structure.

It is important to note that disruptive coloration is not a mutually exclusive trait, and is often combined
with other forms of crypsis such as background matching (Endler, 2006). The natural behaviour of the
Banded Swallowtail is such that they are often found resting on leaf perches or flying close to vegetation
(Tan, 2012). As such, it is entirely possible that the green color on the Banded Swallowtail is similar to
the color of vegetation and helps the butterfly to blend into the surroundings. However, there appears to
be a lack of literature substantiating the idea that a single colored band alone, is able to effectively help a
butterfly blend into the surroundings. Literature seems to instead suggest that in order to function
effectively as a camouflage, a single green band is insufficient, and the whole butterfly has to be covered
in a shade of green as is the case of the Green Hairstreak butterfly (Stavenga, 2010). As a result, there is
more reason to believe that crypsis in the case of the Banded Swallowtail has been achieved by disruptive
coloration, where the green color matches the surroundings, and the stripe disrupts the outline of the
butterfly.

Different aspects of body coloration might well be linked in different ways and can only be truly be
understood from the predator’s perception (Stevens, 2007). For example, camouflage and warning
coloration need not be mutually exclusive (Stevens, 2007). There has been much argument over the role
of the dorsal zigzag pattern present in European vipers (Valkonen et al., 2011). Early studies tend to
suggest that this pattern is an example of disruptive coloration which reduces the vipers’ detectability,
helping it to more easily hunt for prey (Valkonen et al., 2011). However, in more recent studies, where
plasticine snake models with colored zigzag patterns and those with control markings (striped models)
were placed in the field, it was observed that models with zigzag patterns were less predated on than
striped ones, leading scientists to conclude that the zigzag pattern served more of an aposematic effect as
opposed to a disruptive one (Valkonen et al., 2011). There is a certain zigzag pattern in the distorted
models used in the experiment. The earlier point made by Valkonen might help to shed some light on how
the distorted green model was less preyed on than the grey banded model at certain sites. Perhaps the
zigzag pattern might have added small amounts of aposematic and disruptive coloration qualities to the
distorted green models and increase their survivability at certain sites. This shows that a single pattern
could have varying amounts of different anti-predator qualities. It also presents the possibility of
aposematism and disruptive coloration coexisting in the case of the Banded Swallowtail.

Apart from aposematism, disruptive coloration and crypsis, patterns on other butterflies have been shown
to possess other qualities and purposes, such as sexual selection and mimicry (Mallet and Gilbert, 1995).
This is especially so for butterflies of the Heliconius species, where their patterns are postulated to have
mate selection, mimicry and camouflage properties (Mallet and Gilbert, 1995). There is the question of
whether these qualities could apply to the Banded Swallowtail as well. The band patterns found on the
Banded Swallowtail are relatively uncommon, with few butterflies bearing similar color or structure to it.
As such, the possibility of mimicry is ruled out. Also, the same light green band pattern is present on both
males and females, so it is unlikely that the band is used for sexual selection.

In terms of predator deterrence, it is clear that the light green band on the Banded Swallowtail possesses
antipredator qualities, but there is insufficient evidence to pinpoint the qualities to aposematism or
disruptive coloration. We suggest that there could be a combination of both of these qualities at work,
which has led to the significant success of the wild type model as compared to all the other model types.

References
Bennett, A., & Thery, M. (2007). Avian Color Vision and Coloration: Multidisciplinary Evolutionary Biology. The
American Naturalist. Retrieved March 27, 2017, from http://www.journals.uchicago.edu/doi/abs/10.1086/510163

Beatson, R. (1976). Environmental and Genetical Correlates of Disruptive Coloration in the Water Snake, Natrix S.
sipedon. Evolution. doi:10.2307/2407699

Brattstrom, B. (1955). The Coral Snake 'Mimic' Problem and Protective Coloration. Evolution. doi:10.2307/2405591

Chiao, C., Chubb, C., & Hanlon, R. (2007). Interactive effects of size, contrast, intensity and configuration of
background objects in evoking disruptive camouflage in cuttlefish. Vision research. Retrieved March 27, 2017, from
http://www.sciencedirect.com/science/article/pii/S0042698907002076

Cuthill, I., Stevens, M., Sheppard, J., Maddocks, T., Parraga, A., & Troscianko, T. (2005). Disruptive coloration and
background pattern matching. Letters to Nature. doi:10.1038/nature03312

Cuthill, I., Stevens, M., Windsor, A., & Walker, H. (2006). The effects of pattern symmetry on detection of
disruptive and background-matching coloration. Behavioral Ecology. Retrieved March 27, 2017, from
https://academic.oup.com/beheco/article/17/5/828/207206/The-effects-of-pattern-symmetry-on-detection-of.

Diggins, C., Estrada, C., & Rodrigues, A. (2004). Mimicry and the evolution of premating isolation in Heliconius
melpomene Linnaeus. Journal of Evolutionary Biology. doi:10.1111/j.1420-9101.2004.00675.x

Endler, J. (2006). Disruptive and cryptic coloration. Proceedings of the royal society B. doi:10.1098/rspb.2006.3650

Evans, D., & Schmidt, J. (1990). Insect Defenses: Adaptive Mechanisms and Strategies of Prey and Predators.

Exnerová, A., Štys, P., Fučíková, E., Veselá, S., Svádová, K., Prokopová, M., Vojtěch, J., Fuchs, R., Landová, E.
(2007). Avoidance of aposematic prey in European tits (Paridae): learned or innate?. Behav Ecol; 18 (1): 148-156.
doi: 10.1093/beheco/arl061

Finkbeiner, S., Briscoe, A., & Reed, R. (2014). Warning signals are seductive: Relative contributions of color and
pattern to predator avoidance and mate attraction in Heliconius butterflies. International Journal of Organic
Evolution. doi:10.1111/evo.12524

Han, S. K., Lee, D., Lee, H., Kim, D., Son, H., Yang, J., . . . Kim, S. (2016). OASIS 2: online application for
survival analysis 2 with features for the analysis of maximal lifespan and healthspan in aging research. Oncotarget.
doi:10.18632/oncotarget.11269

Harvey, P., & Paxton, R. (1981). The Evolution of Aposematic Coloration. OIKOS. doi:10.2307/3544135

Hatle, J., & Salazar, B. (2000). Aposematic Coloration of Gregarious Insects Can Delay Predation by an Ambush
Predator. Environmental Entomology. Retrieved March 27, 2017, from
http://www.bioone.org/doi/abs/10.1603/0046-225x-30.1.51

Hill, G., & McGraw, K. (2006). Bird Coloration: Mechanisms and measurements.

Ho S, Schachat SR, Piel WH, Monteiro A. (2016). Attack risk for butterflies changes with eyespot number and size.
Royal Society Open Science. doi: 3(1):150614.
Kunte, K. (2008). RESEARCH: WING COLOR PATTERN EVOLUTION, MIMICRY, AND SPECIATION IN
PAPILIO SWALLOWTAIL BUTTERFLIES. Retrieved March 27, 2017, from http://biodiversitylab.org/papilio-
swallowtails

Lindström, L., Alatalo, R., Mappes, J., Riipi, M., & Vertainen, L. (1999). Can aposematic signals evolve by gradual
change? Letters to Nature. doi:10.1038/16692

Mallet, J., & Gilbert, L. (1995). Why are there so many mimicry rings? Correlations between habitat, behaviour and
mimicry in Heliconius butterflies. Linnean Society. doi:10.1111/j.1095-8312.1995.tb01057.x

McGovern, G., Mitchell, J., & Knisley, B. (1984). Field Experiments on Prey Selection by the Whiptail Lizard,
Cnemidophorus inornatus, in Arizona. Journal of Herpetology. doi:10.2307/1564093

Niskanen, M., & Mappes, J. (2005). Significance of the dorsal zigzag pattern of Vipera latastei gaditana against
avian predators. Journal of Animal Ecology. doi:10.1111/j.1365-2656.2005.01008.x

Prudic, K., Skemp, A., & Papaj, D. (2006). Aposematic coloration, luminance contrast, and the benefits of
conspicuousness. Behavioral Ecology. Retrieved March 27, 2017, from
https://academic.oup.com/beheco/article/18/1/41/207914/Aposematic-coloration-luminance-contrast-and-the.

R Core Team (2012). R: A language and environment for statistical computing. R Foundation for Statistical
Computing, Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org/

Santos, J., Coloma, L., & Cannatella, D. (2003). Multiple, recurring origins of aposematism and diet specialization
in poison frogs. PNAS. doi:10.1073/pnas.2133521100

Schaefer, M., & Stobbe, N. (2006). Disruptive coloration provides camouflage independent of background
matching. Proceedings of the royal society B. doi:10.1098/rspb.2006.3615

Stevens, M., & Merilaita, S. (2011). Animal Camouflage: Mechanisms and Function.

Stevens, M., Winney, I., Cantor, A., & Graham, J. (2009). Outline and surface disruption in animal camouflage.
Proceedings of the royal society B. doi:10.1098/rspb.2008.1450

Stevens, M. (2007). Predator perception and the interrelation between different forms of protective coloration.
Proceedings of the royal society B. doi:10.1098/rspb.2007.0220

Svadova, K., Exnerova, A., Stys, P., Valenta, J., Fucikova, A., & Socha, R. (2008). Role of different colours of
aposematic insects in learning, memory and generalization of naïve bird predators. Elsevier. Retrieved March 27,
2017, from http://www.sciencedirect.com/science/article/pii/S0003347208004958

Tan, H. (2012). Life History of the Banded Swallowtail. Retrieved March 27, 2017, from
http://butterflycircle.blogspot.sg/2012/12/life-history-of-banded-swallowtail.html

Thayer, Abbott H.; Thayer, Gerald H. (1909). Concealing-Coloration in the Animal Kingdom. Macmillan, New
York.
Valkonen, J., Niskanen, M., Bjorklund, M., & Mappes, J. (2011). Disruption or aposematism? Significance of dorsal
zigzag pattern of European vipers. Evolutionary Ecology. doi:10.1007/s10682-011-9463-0

Wiklund, C., & Järvi, T. (1982). Survival of Distasteful Insects After Being Attacked by Naive Birds: A Reappraisal
of the Theory of Aposematic Coloration Evolving Through Individual Selection. JSTOR. doi:10.2307/2408077