Masters of Camouflage and Mimicry: Unusual World of Lepidoptera

Abstract

Camouflage is derived from the French word “camoufler”, which originally means disguise. Camoufler derives from camouflet, means a puff of smoke that obscures visibility. As a kind of defense, Lepidoptera uses camouflage, also known as cryptic coloration means to conceal its appearance and blend it with its surroundings. This adaptation prevents Lepidoptera from being detected or recognized by other animals. Lepidoptera, in particular, uses a variety of strategies, including disruptive coloration, active camouflage, concealing coloration, disguise, mimicry, counter-shading, aposematic coloration, color matching, self-decoration, and blending into their surroundings. Camouflage involves animals resembling inanimate objects, while mimicry refers to the similarities between different Lepidoptera species. Since the earliest evolutionists, these ideas have been used as examples of natural selection and adaptation. For surviving in their environments, Lepidoptera employs physical, behavioral, and structural adaptations. Such adaptations include hibernation, migration, instinct, and learning. A Lepidoptera’s body has features called structural adaptations that help it for surviving, such as water retention capacity and defensive coloration. In camouflage, there are eight factors of recognition: position, shape, shadow, texture, color, tone, movement, and shine, those must be considered to ensure that the animal’s location remains concealed. Nature has provided its creatures with great weapons for survival through these amazing techniques.

1. Introduction

During roughly 400 million years on this planet, one million species of insects have developed with a great diversity of shape and color to protect them from predators [1]. Animals have evolved several strategies in prey-predator interactions due to selective pressures, such as mimicry and camouflage [2]. However, several fossil insects have been reported showing plant-like mimesis and debris-carrying camouflage from the Mesozoic [3]. It was reported that the earliest mimetic and defensive strategies of a stick insect from the Middle Jurassic of China, Klimpel’s Hedgehog Cactus, Aclistophasma echinulatum (Weidlich and Werderm) exquisitely preserved abdominal extensions and femoral spines. The distribution of these characteristics mapped onto the phylogeny of (F: Phasmatodea) reveals that abdominal extensions and femoral spines developed multiple times during the evolution of stick insects, Spanish walking stick, Leptynia hispanica Nicolás Vega, indicates that the origin of abdominal extensions predates other modifications, while tergal extensions predate other expansions of the body, such as those of the sterna and pleura, as well as defensive femoral spines [4].

Antipredator defenses among insects commonly involve the interplay of two functional categories. The primary defense, also called passive defense, is the prey’s avoidance of detection by the predator, usually by means of hiding or shifting periods of activity, crypsis, aposematism, or pseudaposematism. The prey’s secondary defense is evading capture after the initiation of a predator’s attack. Secondary defenses involve active escape, antipredator displays, flash coloration, defensive chemical secretion, and feigning death [5, 6]. The active fighting of the prey against a predator, when seized, is sometimes referred to as a separate third category [7, 8]. But, together with the previous series of behaviors, they are referred to as active defenses. Naturally, the ideal situation for any prey is to invest sufficiently in passive mechanisms of defense to avoid the chances of requiring active defense and the increased probability of death that comes when an attack has been initiated [9].

1.1 History

For well over a century, the zoologists have interested in and researching the topic of camouflage. Natural selection, a hypothesis put out by Charles Darwin in 1859, contends those traits, like camouflage, arose as a result of given specific animals a reproductive advantage that allow them to produce, on average, more offspring than other members of the same species. His research demonstrated that the pupae of the swallow-tailed moth, Ourapteryx sambucaria Linnaeus, 1758 [Family (F): Geometridae] were camouflaged to resemble the backgrounds on which they were raised as larvae (Figure 1) [11].

Figure 1.

(i): Swallow-tailed moth, Ourapteryx sambucaria Linnaeus, 1758 [Family (F): Geometridae]: a; pupa showing color assumed when larva has been placed on white paper before pupation: b; pupa with usual dark color assumed in cocoon with natural size: c [2]; (ii): Camouflage, the alluring coloration in the orange tip butterfly, Anthocharis cardamines (Linnaeus, 1758) (F: Pieridae); A. cardamines ♂: d; A. cardamines ♂: e; A. cardamines ♂: f; A. cardamines ♂: g [3]; (iii): Peppered moth, Biston betularia Linnaeus, 1758 (F: Geometridae): h; its camouflage with substrate of natural background: i; a B. betularia caterpillar mimics a twig: j [10].

The avoidance of detection by predators and parasites is critical to survival. Two complex mechanisms for such avoidance are mimicry and camouflage, with fossils providing valuable insight into the evolution of these strategies. It was reported that a diverse insect assemblage exhibited these adaptations from mid-Cretaceous Kachin amber (99 million years ago), including plant mimesis in Tridactylidae (pygmy mole crickets) and debris-carrying camouflage in Gelastocoridae (toad bugs) and Psocodea (bark lice) [12].

Poulton reported that the color of animals, particularly camouflage. Different types of camouflage, such as specific defensive resemblance, where an animal mimics another object, or general aggressive resemblance, where a predator blends in with the background to approach prey, were classified by him in his book. For colors of animals, his research demonstrated that O. sambucaria pupae were camouflaged to blend with the environments in which they were raised as larvae. The major means of camouflage at the time was thought to be Poulton’s overall protective resemblance (Figure 1,i:a-c) [13].

The orange-tipped butterfly, Anthocharis cardamines (Linnaeus, 1758) (F: Pieridae), has attractive colorations, according to Beddard (1940) that the sporadic green patches on the underside of the wings may have been intended to represent a crude drawing of the plant’s miniature flowerets (an umbellifer), which would explain how they are similar to one another (Figure 1,ii:d-g) [14].

The peppered moth, Biston betularia Linnaeus, 1758 (F: Geometridae), is a temperate species of night-flying moth. Natural selection and population genetics are examples of evolution in B. betularia. Prey animals frequently replicate their surroundings, as seen in the B. betularia caterpillar’s imitation of a branch in Figure 1(iii:h-j) [10].

1.2 Camouflage and disguise

It is difficult to distinguish between disguise and camouflage, but generally speaking, camouflage is used to describe something that can blend in nicely with a natural background or substrate, thanks to its color, pattern, and texture. Various types of butterflies and moths have colors and patterns that blend in with each of these backdrops when they rest on a variety of substrates, such as leaf, soil, rocks, and tree trunks. On the other side, disguise refers to a butterfly or moth that resembles another natural object, like a leaf or flower [15, 16].

Numerous caterpillars are skilled twig impersonators. The geometrids (F: Geometridae) is a special huge group that goes by this name (Figure 2a). When disturbed, they will freeze and give the impact of heighten. They have the appearance and feel of twigs. They lack middle-body legs to make them appear like twigs, and as a result, they walk in a loop, which gives the group its name [15, 16].

Figure 2.

Camouflage and disguise in Lepidoptera: a: caterpillar (F: Geometrids); b: green hairstreak, Callophrys rubi (Linnaeus, 1758) (F: Lycaenidae); c: lysippus metalmark, Riodina lysippus (Linnaeus, 1758) (F: Riodinidae); d: brimstone, Gonepteryx rhamni (Linnaeus, 1758) (F: Pieridae); e: orange daggerwing, Marpesia berania (Hewitson 1852) (F: Nymphalidae); f: dead leaf butterfly, Kallima inachus (Doyère, 1840) i: dorsal surface; ii: ventral surface (F: Nymphalidae); g: Ithomiine glasswings, Greta oto (Hewitson, 1854) (F: Nymphalidae); h: blushing phantom, Cithaerias pireta (Stoll, 1780) (F: Nymphalidae); i: uncolored clearwing-satyr, Dulcedo polita (Hewitson, 1869) (F: Nymphalidae); j: octauius swordtail, Chorinea octauius (Fabricius, 1787) (F: Riodinidae); k: white dragontail, Lamproptera curius (Fabricius, 1787) (F: Papilionidae); l: grayling butterfly, Hipparchia semele (Linnaeus, 1758) (F: Nymphalidae); m: peruana skipper, Hylephila peruana Draudt, 1923 (Family: Hesperiidae); n: dingy skipper, Erynnis tages (Linnaeus, 1758) (F: Hesperiidae); o: merveille du jour, Griposia aprilina (Linnaeus, 1758) (F: Noctuidae); p: buff-tip, Phalera bucephala (Linnaeus, 1758) (F: Noctuidae); q: Chinese character, Cilix glaucata (Scopoli, 1763) (F: Drepanidae); r: angle shades, Phlogophora meticulosa (Linnaeus, 1758) (F: Noctuidae); s: eyed hawk-moth, Smerinthus ocellatus (Linnaeus, 1758) (F: Sphingidae) [16].

To evade predators, butterflies employ a variety of strategies. Sometimes, like in the case of the green hairstreak, Callophrys rubi (Linnaeus, 1758) (F: Lycaenidae) resting on green foliage and the Neotropical Lysippus Metalmark, Riodina lysippus (Linnaeus, 1758) (F: Riodinidae) hides under leaves to avoid being seen, respectively (Figure 2b and Figure 2c). A nice example of a British butterfly that is challenging to see when resting on white garlic mustard blooms is the orange-tipped, Anthocharis cardamines (F: Pieridae) (Figure 1,ii:d,e,f,g) [16].

The brimstone, Gonepteryx rhamni (Linnaeus, 1758) (F: Pieridae) (Figure 2d), is a great example of deception since it has the coloration, shape, and elevated veins of a living leaf. The orange dagger wing, Marpesia berania (Hewitson, 1852) (F: Nymphalidae) (Figure 2e), and the pale-spotted leafwing, Memphis pithyusa (Felder, 1869) (F: Nymphalidae) are two examples of the many species that may be found in the tropics those mimic decaying brown leaves [16].

In Malaysia, the dead leaf butterfly, Kallima inachus (Doyère, 1840) (F: Nymphalidae), is the ideal illustration of camouflage. It has wings (dorsal surface) (Figure 2:fi) that can tolerate rain and wetness in deep forests, allowing it to fly high and quickly. When threatened or pursued by birds, it flies erratically and swiftly, falls to the ground, and buries itself in the undergrowth by folding its wings to blend in. Its wings resemble a dry leaf with dark veins, when they are closed (ventral surface (Figure 2:fii). The K. inachus prefers to dwell on tree trunks or the ground with fallen leaves, but it occasionally visits flowers. Its natural predators include birds, ants, and spiders, and it feeds on rotting fruits and animal excrement [17].

Butterflies that use transparency to obscure themselves comprise the Ithomiine (Tribe: Ithomiini) glasswings, Greta oto (Hewitson, 1854) (F: Nymphalidae) (Figure 2g), the Neotropical satyrines, such as the blushing phantom, Cithaerias pireta (Stoll, 1780) (F: Nymphalidae) (Figure 2h) and the uncolored clearwing-satyr, Dulcedo polita (Hewitson, 1869) (F: Nymphalidae) (Figure 2i). The Octauius swordtail, Chorinea octauius (Fabricius, 1787) (F: Riodinidae) (Figure 2j), and the white dragon tail, Lamproptera curius (Fabricius, 1787) (F: Papilionidae) (Figure 2k) are additional specimens from other families [18].

The grayling butterfly, Hipparchia semele (Linnaeus, 1758) (F: Nymphalidae), which is expertly disguised when resting on decaying wood, is depicted in Figure 2. The skipper from Peru, Hylephila peruana Draudt, 1923 (F: Hesperiidae) (Figure 2m) is exceedingly challenging to spot among the dry grasses of its Andean environment due to the disruptive patterning. In leaf litter, the dingy skipper, Erynnis tages (Linnaeus, 1758) (F: Hesperiidae) (Figure 2n) blends well [19].

Because moths are frequently eaten, they have developed a range of camouflage techniques, including delicate colors and patterns that fit in with their environment. The outcomes are frequently rather beautiful, in addition to being incredibly brilliant. They have colors and patterns to match the locations, where they rest because they are particularly vulnerable to being seen in daylight. Because they blend into the bark of branches and are difficult to spot in glooms, many moths have patterns of gray and brown on their wings.

The merveille du jour, Griposia aprilina (Linnaeus, 1758) (F: Noctuidae) (Figure 2o), is the ideal complement to bark that has been covered with lichen. The buff-tip, Phalera bucephala (Linnaeus, 1758) (F: Noctuidae) (Figure 2p), has taken things a step further by resembling branches in both color and shape. It also looks like a particular kind of birch tree twig with its distinctive silvery-colored bark. In case that was not enough, it even resembles a broken birch twig exactly [15].

The Chinese character moth, Cilix glaucata (Scopoli, 1763) (F: Drepanidae) (Figure 2q), is quite noticeable, but it looks exactly like a bird dropping, both in shape and color; consequently, it is definitely not a tempting morsel. A few moths practice camouflage not to be firm to see but to disguise themselves as something unpleasant; subsequently, those predators will not even deliberate on eating them [15].

Numerous moths have markings that obstruct their contour, making it difficult to identify their shape. The common garden moth known as the angle shades, Phlogophora meticulosa (Linnaeus, 1758) (F: Noctuidae) (Figure 2r), employs a variety of tactics. Its form is divided into less moth-like pieces by the triangle patterns. The hues of this material are also suitable for hiding among fallen leaves or on bark. Additionally, when at rest, the wings’ crumpled shape resembles a dry leaf or the folds and fissures in bark [15].

A cunning combination of two techniques (camouflage and mimicry) is used by the eyed hawk-moth, Smerinthus ocellatus (Linnaeus, 1758) (F: Sphingidae) (Figure 2s). It is difficult to see against bark because, while it is resting, its forewings, which are camouflaged, conceal its hind wings. However, if it is startled, it abruptly unfolds its hind wings to reveal a flash of bright eyes sufficient to surprise and scare away a predator [15].

1.3 Mimicry

An animal or plant’s defense against predators is its close external similarity to another animal, plant, or inanimate item. Unappealing butterfly patterns are frequently noticeable. According to experiments, some bird predators may memorize these patterns and learn to steer clear of preying on species with similar patterns in the future. Naturalists claim that leaf mimicry offers some of the most striking illustrations of disguise-based on camouflage. Plentiful species of Lepidoptera that bear a resemblance to leaves also have wing patterns that closely look like the irregularly shaped holes left by insect or decay damage. Notwithstanding claims that these patterns can either make them look more like damaged leaves or interfere with surface presence by giving off deceptive profundity indications. The buddha moth, Siculodes aurorula (Guenée, 1858) (F: Thyrididae) was used in two field trials employing artificial butterfly-like targets. It is shown in Figure 3i that untruthful pit characters proposition extensive subsistence profits against avian predation. Additionally, it was shown in supercomputer-built visual research experimentation that humans were similarly hindered from detecting such targets. Similar to how dark marks do not have the same effect as light ones, in fact, they increase detection. It was determined that the mechanism is the disruption of a homogeneous wing surface (surface disruptive camouflage), and because the holes resemble those occasionally present in actual leaves, the disruptive benefits are not negated by the costs associated with being noticeable [19].

Figure 3.

Mimicry in Lepidoptera: i: buddha moth, Siculodes aurorula Guenée, 1858 (F: Thyrididae); ii: North American viceroy, Limenitis archippus (Cramer, 1776) (F: Nymphalidae); iii: Monarch, Danaus plexippus (Linnaeus, 1758) (F: Nymphalidae); iv: tiger mimic white, Dismorphia amphione (Cramer, 1779) (F: Pieridae); v: postman butterfly, Heliconius melpomene (Linnaeus, 1758) (F: Nymphalidae); vi: milkweed butterfly, Ithomiines mechanitis polymnia Godman and Salvin, 1879 (F: Nymphalidae); vii: spicebush swallowtail, Papilio troilus (Linnaeus, 1758) (F: Papilionidae); viii: pipevine swallowtail, Battus philenor (Linnaeus, 1771) (F: Papilionidae); xiii: Omphale’s king shoemaker, Lycorea pasinuntia (Stoll, 1780) (F: Nymphalidae); xiv: Tropical milkweed butterflies, Lycorea ilione (Cramer, 1775 ) (F: Nymphalidae); xv: Dismorphiine asionym, Patia orise (Boisduval, 1836) (F: Pieridae); xvi: tiger moths, Cosmosomaspp. Hübner, [1823] (F: Erebidae) [20].

1.3.1 Batesian mimicry

Batesian mimicry is that in which an edible species is protected by its resemblance to one avoided by predators. The mimic gains protection and the predator does not eat it because the predator made a mistake. For example, the palatable North American species, viceroy, Limenitis archippus (Cramer, 1776) (F: Nymphalidae) (Figure 3ii), which bears a quite remarkable resemblance to the highly toxic Monarch, Danaus plexippus (Linnaeus, 1758) (F: Nymphalidae) (Figure 3iii). However, recent studies have revealed that both of these species are unpalatable, making them Mullerian mimics. There are, however, many other instances of genuine Batesian mimicry, including the palatable, the tiger mimic white, Dismorphia amphione (Cramer, 1779) (F: Pieridae) (Figure 3iv), and the postman butterfly, Heliconius melpomene (Linnaeus, 1758) (F: Nymphalidae) (Figure 3v) species which mimic toxic milkweed butterfly, Ithomiines Mechanitis polymnia Godman and Salvin, 1879 (F: Nymphalidae) (Figure 3vi); and the palatable spicebush Swallowtail, Papilio troilus (Linnaeus, 1758) (F: Papilionidae) (Figure 3vii), which mimics the toxic pipevine swallowtail, Battus philenor (Linnaeus, 1771) (F: Papilionidae) (Figure 3viii). Bates proposed the hypothesis that occasionally, edible species created mutant forms with visual traits resembling poisonous species. Therefore, he reasoned, they would be less likely to be eaten by birds and would pass those traits on to their progeny. Bates hypothesized that the palatable species had developed to nearly resemble the dangerous species as a result of subsequent mutations over millennia [20].

1.3.2 Mullerian mimicry

Mullerian mimicry is a form of mimicry in which two or more harmful or unpalatable Lepidoptera develop similar appearances as a shared protective device. In 1879, Mullerian realized that there were also many cases, where both the mimic and the model were unpalatable. When a bird catches any one of these butterflies, either model or mimic, and realizes it is unpalatable or toxic, it quickly learns to keep away from all similarly patterned species. This type of evolutionary cooperation is referred to as Mullerian mimicry and is a very common phenomenon among the families Ithomiinae, Danainae, and Pieridae. Muller demonstrated mathematically that this form of mimicry is biased in favor of the scarcer species by a factor of the square of the ratio of species abundance. It is advantageous for there to be a large number of species involved in a Mullerian mimicry complex as it increases the power of the warning signal [21].

1.3.3 Tiger complex

The tiger complex, a collection of roughly 200 Neotropical species that all have a common pattern of orange and yellow stripes on a black background, is a well-known instance of butterfly mimicry. The complex includes many unpalatable Ithomiines such as the harmonia tiger-wing, Tithorea harmonia (Cramer, 1777) (F: Nymphalidae) (Figure 3ix); the tarricina longwing, Tithorea tarricina Hewitson, 1858 (F: Nymphalidae) (Figure 3x); the clearwing (Ithomiine) butterflies, Melinaea marsaeus (Hewitson, 1860) (F: Nymphalidae) (Figure 3xi) and the forbestra, Forbestra equicola (Cramer, 1780) (F: Nymphalidae) (Figure 3xii), the unpalatable Danaines such as Omphale’s king Shoemaker, Lycorea pasinuntia (Stoll, 1780) (F: Nymphalidae) (Figure 3xiii), and several highly toxic day-flying moths from the Arctiid subfamily Pericopinae. At the end of the dry season, members of the tiger complex typically congregate in huge groups in damp gullies in the forest. Mimicry is most effective as a defense during this period, when they are very submissive and easy prey for birds. Any bird that has the bad experience of tasting one of the tiger-complex members soon learns to stay away from any species that has a similar appearance, and they might even be able to communicate their disagreeable character to other birds [22].

1.3.4 Mimicry rings

There are several more mimicry rings. The first example of mimicry ring, the glass-wing ring is a group of fairly large species with transparent wings. The butterflies that contain toxic Mullerian mimicry, mostly belong to the genera of the tribe, Ithomiine such as Methona and Thyridia, and the tribe, toxic Danaini with genera Monarch. The tropical milkweed butterfly, Lycorea ilione (Cramer, 1775) (F: Nymphalidae) (Figure 3xiv) shows Mullerian mimicry, and the palatable Dismorphiine basionym, Patia orise (Boisduval, 183) (F: Pieridae) (Figure 3xv) shows Batesian mimicry. The second example of mimicry ring, the orange-ring is comprised of a group of bright orange species, including the Julia butterfly, Dryas iulia (Fabricius, 1775) (F: Nymphalidae), the Juliette, Eueides aliphera (Godart, 1819) (F: Nymphalidae) and the ruddy daggerwing, Marpesia petreus Cramer, 1776 (F: Nymphalidae) [23].

1.3.5 Wasp mimicry

The day-flying moths (F: Sesiidae) include many species with short, translucent wings and black-and-yellow-banded bodies. They remarkably resemble wasps and hornets, and because of their likeness to these stinging insects, they almost definitely avoid predation. Numerous Arctiid moth species that belong to the Cosmosoma spp. Hübner, [1823] (F: Erebidae) (Figure 3xvi) are found in the Neotropics. These moths have brightly patterned red, orange, or yellow bodies and translucent wings. The majority of them are nocturnal in behavior and hide out among vegetation during the day, where their intimidating demeanor may prevent them from being eaten [24].

1.3.6 Transformational mimicry

Mimicry is not just limited to fully-grown butterflies and moths. Many tasty caterpillars imitate unappealing species. After molting, caterpillars frequently alter their look on regular basis. It is possible for a caterpillar to imitate different models at different instars. Additionally, the adult butterfly or moth that develops from that caterpillar could look just like a different species [25].

1.3.7 Aposematic coloration

Aposematic coloration is a term used to describe colors and patterns those 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 color schemes that reflect these facts in order to label themselves as being unpleasant to eat, thereby dissuading birds from attacking them. Equally, it is unsurprising that a significant number of palatable species have evolved to mimic the patterns of toxic species in order to trick birds into avoiding eating. For example, the unpalatability of the traditional Chinese, Altinote dicaeus callianira(Geyer, 1837) (F: Nymphalidae) (Figure 4a). Birds can remember the colors 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 [26].

Figure 4.

Aposematic coloration in Lepidoptera: a: traditional Chinese, Altinote dicaeus callianira (Geyer, 1837) (F: Nymphalidae); b: bullseye moths, Automeris liberia (Cramer, 1780) (F: Saturniidae); c: peacock butterfly, Aglais io (Linnaeus, 1758) (F: Nymphalidae); d: giant atlas moth, Attacus atlas (Linnaeus, 1758) (F: Saturniidae) e: silk moths, Rothschildia spp. Grote, 1896 (F: Saturniidae); f: Neotropical owl butterfly, Caligo teucer (Linnaeus, 1758) (F: Nymphalidae); g: gatekeeper, Pyronia tithonus (Linnaeus, 1771) (F: Nymphalidae) h: old world swallowtail, Papilio machaon Linnaeus, 1758 (F: Papilionidae); i: prola beauty or red flasher, Panacea prola (Doubleday, 1848) (F: Nymphalidae); j: helenor or common blue morpho, Morpho helenor (Cramer, 1775) (F: Nymphalidae) i: dorsal surface; ii: ventral surface [21].

1.3.8 Diematic patterns

The diematic patterns are defensive markings that have the power to terrify or startle prospective predators. The usage of ocelli is the most prevalent type of diematic defense. These typically appear as a pair of false-eye marks that can scare off a predator. These ocelli are extremely noticeable and resemble the eyes of monkeys or raptors in many butterflies and moths. The eyed hawk-moth, Smerinthus ocellatus (Linnaeus, 1758) (Family: Sphingidae) (Figure 2s) is one example. The ocelli of the Neotropical bullseye moths, Automeris liberia (Cramer, 1780) (F: Saturniidae) (Figure 4b) are typically concealed beneath the forewings of dead leaves with cryptic patterns. When startled, they fall to the ground and begin to twitch rhythmically, bringing as much attention to the ocelli as possible. Its frightening appearance would be more than enough to stop a bird from attacking. Figure 4c shows the peacock butterfly, Aglais io (Linnaeus, 1758) (F: Nymphalidae), which is stunning to humans but terrifying to small birds. Figure 4e shows the adult snake-head markings on the apex of the enormous atlas moth, Attacus atlas (Linnaeus, 1758) (F: Saturniidae), while Figure 4d shows the adult snake-head markings on the apex of the silk moth, Rothschildia spp. Grote, 1896 (F: Saturniidae). These species’ forewing apexes are lobed and have markings that resemble a snake’s mouth and eyes. The illusion is strengthened by the wings’ rhythmic movements, which highlight the snake-head markings [27, 28].

1.3.9 Decoys to distract birds

The terrifying effect of diematic marks frequently last a short while. A bird may renew its attack, once it has recovered from its first fright. The Neotropical owl butterfly, Caligo teucer (Linnaeus, 1758) (F: Nymphalidae) (Figure 4f) and the gatekeeper, Pyronia tithonus (Linnaeus, 1771) (F: Nymphalidae) (Figure 4g) are two examples of predators that the ocelli on the butterfly’s wings divert and confuse in such situations. An attacking bird will always try to predict, where its victim will flee; thus, it will aim to attack just in front of the head. The bird is tricked by the phony head into shooting behind the butterfly instead. The butterfly then escapes by darting off in the opposite direction from where the bird anticipates it [29, 30].

1.3.10 Patterns to confuse

There are patterns on many butterflies that, at first glance, appear to have no purpose. What would be the purpose, for instance, of a visually appealing pattern that is simple to recall, like the chequered pattern on the old world swallowtail, Papilio machaon Linnaeus, 1758 (F: Papilionidae) (Figure 4h). When disturbed, it suddenly flicks its closed wings open in the same manner as the peacock and other ocelli-equipped species. It generally rests with its wings closed [31, 32].

1.3.11 Signaling danger to other butterflies

Although, it is well-known that warning coloration is intended to directly deter birds and other predators, the prola beauty or red flasher, Panacea prola (Doubleday, 1848) (F: Nymphalidae) (Figure 4i) from Peru serves an entirely opposite purpose. On riverbanks, a lot of males congregate to consume mineralized moisture. They expose their metallic blue dorsal surface while basking and feeding, which aids other passing males in identifying them and homing in on the feeding grounds. The butterflies must use a defense mechanism since birds may readily attack a gathering of butterflies on the ground. When there is even a slight disturbance, 1 or 2 butterflies become agitated and begin nervously fanning their wings, causing the other butterflies to see their bright red undersides [14, 33].

1.3.12 Flash coloration

Many butterflies have alternate display of a bright dorsal surface and somber underside. This is recognized as flash coloration. An example is the South American Telenor blue morpho or common blue morpho, Morpho helenor (Cramer, 1775) (F: Nymphalidae) (Figure 4j), which has a brilliant iridescent blue dorsal surface (Figure 4ji) and black with eyes on ventral surface (Figure 4jii) that makes it highly visible to predators as well as to potential mates. A colorful dorsal surface and a somber underside are common displays of many butterflies. This is recognized as flash coloration. For instance, the South American the helenor blue morpho or common blue morpho, Morpho helenor (Cramer, 1775) (F: Nymphalidae) (Figure 4j) is highly visible to predators as well as for potential mates due to its blue brilliant iridescent on the dorsal surface (Figure 4ji) and several eyes pattern on black ventral surface (Figure 4jii). When startled, a butterfly will land quickly and close its wings, consequently, that only the dark brown underside is visible. The secondary decoy-ocelli defense may enable the butterfly to flee, if a chasing bird spot it at rest after landing, since it will direct the bird’s beak away from the body of the butterfly and toward the wing margins [34].

2. Conclusion

Lepidoptera have a wonderful defense mechanism called camouflage that enables them to efficiently hide their appearance and blend with their surroundings. This fascinating subject has been investigated by the zoologists for well over a century. It is distinguished from disguise by its ability to harmoniously merge with a natural background or substrate through color, pattern, and texture. Many butterflies and moths have developed colors and patterns that seamlessly blend with their surroundings, such as leaves, soil, rocks, and tree trunks. Disguise, on the other hand, refers to species that resemble other natural objects like leaves or flowers. Another fascinating facet of how Lepidoptera defend themselves is mimicry. Batesian mimicry involves edible species mimicking avoided ones to protect themselves from predators, while Mullerian mimicry involves multiple harmful or unpalatable Lepidoptera developing similar appearances for shared protection. In addition to the well-known tiger complex, the Lepidoptera kingdom contains several other mimicry rings, such as the glass-wing ring, which represents species with transparent wings, and the orange-ring, which is made up of a collection of species with brilliant orange wings. The diversity of mimicry strategies extends to wasp mimicry, transformational mimicry, aposematic coloration, diematic patterns, decoys to distract predators, patterns to confuse them, and even signaling danger to other butterflies through flash coloration. As scientists work to understand the intricate workings of these astonishing adaptive tactics, the world of camouflage and mimicry in Lepidoptera never ceases to astound both scientists and nature lovers.