An eyespot (sometimes ocellus) is an eye-like marking. They are found in butterflies, reptiles, cats, birds and fish.
Eyespots could be explained in at least three different ways. They may be a form of mimicry in which a spot on the body of an animal resembles an eye of a different animal, to deceive potential predator or prey species. They may be a form of self-mimicry, to draw a predator's attention away from the prey's most vulnerable body parts. Or they may serve to make the prey appear inedible or dangerous. Eyespot markings may play a role in intraspecies communication or courtship; the best-known example is probably the eyespots on a peacock's display feathers.
The pattern-forming biological process (morphogenesis) of eyespots in a wide variety of animals is controlled by a small number of genes active in embryonic development, including the genes called Engrailed, Distal-less, Hedgehog, Antennapedia, and the Notch signaling pathway.
Artificial eyespots have been shown to reduce predation of cattle by lions.
The eye-like markings in some butterflies and moths and certain other insects, as well as birds like the sunbittern, serve functions in addition to mimicry;  indeed, it is unclear whether they actually mimic eyes.  There is evidence that eyespots in butterflies are antipredator adaptations, either in deimatic displays to intimidate predators, or to deflect attacks away from vital body parts.   In species such as Hipparchia semele , the conspicuous eyespots are hidden at rest to decrease detectability, and only exposed when they believe potential predators are nearby.  Butterfly eyespots can mimic dead leaves for camouflage from predators, as seen in Bicyclus anynana ; this is a response to a seasonal fall in temperature, causing a shift in selection towards smaller, less conspicuous eyespots among those individuals developing at that time.  Butterfly eyespots may play a role in mate recognition and sexual selection.  Sexual selection drives the diversification of eyespots in different species of butterflies, as mates select for characteristics like size and brightness.  
Some species of caterpillar, such as those of hawkmoths (Sphingidae), have eyespots on their anterior abdominal segments. When alarmed, they retract the head and the thoracic segments into the body, leaving the apparently threatening large eyes at the front of the visible part of the body. 
Butterflies such as the blues (Lycaenidae) have filamentous "tails" at the ends of their wings and nearby patterns of markings, which combine to create a "false head". This automimicry misdirects predators such as birds and jumping spiders (Salticidae). Spectacular examples occur in the hairstreak butterflies; they commonly perch upside down with the false head raised and shift their rear wings repeatedly, causing antenna-like movements of the "tails" on their wings. Studies of rear-wing damage support the hypothesis that this deflects attacks from the insect's head.  
Some reptiles, such as the sand lizard of Europe, have eyespots; in the sand lizard's case, there is a row of spots along the back, and a row on each side. 
Many species of cat, including Geoffroy's cats, jungle cats, pampas cats, and servals, have white markings, whether spots or bars, on the backs of their ears; it is possible that these signal "follow me" to the young of the species. There may be an evolutionary trade-off in this case between nocturnal camouflage and intraspecific signalling.  
Male birds of some species, such as the peacock, have conspicuous eyespots in their plumage, used to signal their quality to sexually selecting females. The number of eyespots in a peacock's train predicts his mating success; when a peacock's train is experimentally pruned, females lose interest.   Several species of pygmy owl bear false eyes on the back of the head, misleading predators into reacting as though they were the subject of an aggressive stare. 
Some fish have eyespots. The foureye butterflyfish gets its name from a large and conspicuous eyespot on each side of the body near the tail. A black vertical bar on the head runs through the true eye, making it hard to see.  This may deceive predators in two ways: into attacking the tail rather than the more vulnerable head, and about the fish's likely direction of travel. The foureye butterflyfish eyespot is thus an example of self-mimicry.  For the same reason, many juvenile fish display eyespots that disappear during their adult phase.  Some species of fish, like the spotted mandarin fish and spotted ray, maintain their eyespots throughout their adult lives. These eyespots can take a form very similar to those seen in most butterflies, with a focus surrounded by concentric rings of other pigmentation. 
Butterfly eyespots are formed during embryogenesis as a result of a morphogenetic signalling centre or organizer, called the focus. This induces neighbouring cells to produce specific pigments which pattern the eyespot.   
Early experiments on eyespot morphogenesis used cautery on the butterfly wing eyespot foci to demonstrate that a long range signaling mechanism or morphogen gradient controlled the formation of eyespots in both space and time.  The findings cannot be explained by a simple source/diffusion model,  but could be explained by either a source/threshold model, in which the focus creates the morphogen, or by the sink model, in which the focus generates a gradient by removing a morphogen which was created elsewhere.  Several genes involved in eyespot formation have been identified that can fit into these models, but only two of them have been functionally tested. These genes are the transcription factor Distalless (Dll) and the ligand (a signalling substance that binds a cell surface receptor) Hedgehog (Hh). 
Butterfly eyespot morphology appears to be the result of the evolution of an altered version of the regulatory circuit which patterns the wings of other insects. This rogue regulatory circuit is able to pattern both the anterior and posterior eyespots independent of the usual anterior/posterior wing compartmentalization restrictions seen in the fruit fly Drosophila .   The altered regulatory circuit redeploys early developmental signaling sources, like the canonical hedgehog (Hh) pathway, Distal-less (Dll), and engrailed (En), breaking the anterior/posterior compartmentalization restrictions through increased localized levels of Hh signaling.    In turn, this raises expression of its receptor Patched (Ptc) and transcription factor.  Normally, in Drosophila , engrailed acts in the posterior compartment to restrict Ptc and Cubitus interruptus (Ci) expression to the anterior compartment by repressing transcription of Ci, thereby preventing Ptc expression.  From the perspective of evolutionary developmental biology, understanding the redeployment and plasticity of existing regulatory mechanisms in butterfly eyespot locus development has given more insight into a fundamental mechanism for the evolution of novel structures.  
The Distal-less gene is present in almost all eyespot organizers, making it an ideal candidate to carry out major functions of eyespot formation. During the wing imaginal disc development Dll, has two expression domains separated by a temporal component. First Dll is expressed in a group of cells in the center of what will become the focus and eventually the eyespot. This expression starts during the middle of the fifth instar larva and lasts until the pupal stage. The second domain starts around 20 hours after pupation around the original central cluster of cells, in an area in which a black ring of the eyespot will be formed. Functional experiments using transgenic Bicyclus anynana (the squinting bush brown butterfly) have shown that overexpression or down-regulation of Dll in the first expression domain correlates with bigger and smaller eyespots respectively. However, if this is done on the second domain then the overall size of the eyespots remains the same, but the width of the black ring raises with a higher amount of Dll. This suggests that Dll might be responsible for the differentiation of the focus in the first expression domain and might be involved in establishing the ring colour patterns in the second domain. These experiments together with the wide distribution of Dll across eyespot forming butterflies suggest that this transcription factor is a central regulator for the correct patterning of the eyespots. 
The Hedgehog (Hh) gene is the other element that has been functionally tested in the formation of eyespots. Investigating genes involved in wing development and morphogenetic activity has led to the discovery that Hh has a primary role in the morphogenetic signaling center of the foci.  In a manner that is similar to the development of Drosophila fruit flies, Hh is expressed in all cells in the posterior compartment of the developing butterfly wing during the mid fifth instar of butterfly wing development. However, in butterflies, Hh expression is significantly higher in those cells that flank the potential foci.  Higher transcription levels of Hh, along with other known associates of the Hh pathway, namely patched (Ptc) the Hh receptor, and cubitus interruptus (Ci), the Hh transcription factor is seen throughout the mid to late fifth instar as well, which further implies a role for Hh signaling in eyespot development and patterning. 
Furthermore, cells that are flanked by the cells expressing the highest level of Hh signaling are fated to become the foci, indicating that focus cell fate determination relies on high concentrations of Hh in surrounding cells.  However, this observation has not been totally confirmed as a rule for multiple butterfly species.  Studies tried to extrapolate the result of Hh pathway involvement by looking for the expression of Ci in Bicyclus anynana.  Here they observed that both seem to be expressed in eyespots, suggesting a relation with the Hh signaling pathway. However, other studies did not find evidence of Hh expression in B. anynana. 
The Notch (N) gene expression precedes an upregulation of Dll in the cells that will become the center of the focus. This makes N the earliest developmental signal, so far studied, that is related with the establishment of the eyespots. Loss of N completely disrupts Dll expression, and eventually eyespot formation, in several butterfly species. A variety of other wing patterns are determined by N and Dll patterns of expression in early development of the wing imaginal disc, suggesting that a single mechanism patterns multiple coloration structures of the wing. 
Butterfly eyespots are formed by an interplay of at least 3 genes, namely Distal-less (Dll), spalt (sal), and Antennapedia (Antp), hence their evolution has been shaped by differential expression of these genes in different butterfly taxa, as shown in Bicyclus anynana . 
Eyespots painted on the rumps of cows have been shown to reduce cattle predation in Africa. The study authors, Cameron Radford and colleagues, note that in the Sundarbans, forest users wear face masks with eye markings on the backs of their heads in the hope of reducing tiger attacks. In the study on 2061 cattle in 14 herds over 4 years, 683 were given eye markings, 543 were painted with crosses, and 835 were unpainted. None of the eyed cattle were predated, but 4 cross-marked and 15 unmarked cattle were killed, one by a leopard, the rest by lions. Both the eyespots and the cross markings provided statistically significant protection. The cattle were always in mixed groups of marked and unmarked animals; it is not known whether marking all animals in a herd would provide effective protection. 
Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises the large superfamily Papilionoidea, which contains at least one former group, the skippers, and the most recent analyses suggest it also contains the moth-butterflies. Butterfly fossils date to the Paleocene, about 56 million years ago.
Evolutionary developmental biology is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved.
Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most animals and some plants. Differences may include secondary sex characteristics, size, weight, color, markings, or behavioral or cognitive traits. Male–male reproductive competition has evolved a diverse array of sexually dimorphic traits. Aggressive utility traits such as “battle” teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to sexual selection and natural selection. The opposite of dimorphism is monomorphism, when both biological sexes are phenotypically indistinguishable from each other.
In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry functions to protect a species from predators, making it an anti-predator adaptation. Mimicry evolves if a receiver perceives the similarity between a mimic and a model and as a result changes its behaviour in a way that provides a selective advantage to the mimic. The resemblances that evolve in mimicry can be visual, acoustic, chemical, tactile, or electric, or combinations of these sensory modalities. Mimicry may be to the advantage of both organisms that share a resemblance, in which case it is a form of mutualism; or mimicry can be to the detriment of one, making it parasitic or competitive. The evolutionary convergence between groups is driven by the selective action of a signal-receiver or dupe. Birds, for example, use sight to identify palatable insects and butterflies, whilst avoiding the noxious ones. Over time, palatable insects may evolve to resemble noxious ones, making them mimics and the noxious ones models. In the case of mutualism, sometimes both groups are referred to as "co-mimics". It is often thought that models must be more abundant than mimics, but this is not so. Mimicry may involve numerous species; many harmless species such as hoverflies are Batesian mimics of strongly defended species such as wasps, while many such well-defended species form Müllerian mimicry rings, all resembling each other. Mimicry between prey species and their predators often involves three or more species.
Aglais io, the European peacock, more commonly known simply as the peacock butterfly, is a colourful butterfly, found in Europe and temperate Asia as far east as Japan. It was formerly classified as the only member of the genus Inachis. It should not be confused or classified with the "American peacocks" in the genus Anartia; while belonging to the same family as the European peacock, Nymphalidae, the American peacocks are not close relatives of the Eurasian species. The peacock butterfly is resident in much of its range, often wintering in buildings or trees. It therefore often appears quite early in spring. The peacock butterfly has figured in research in which the role of eyespots as an anti-predator mechanism has been investigated. The peacock is expanding its range and is not known to be threatened.
Stabilizing selection is a type of natural selection in which the population mean stabilizes on a particular non-extreme trait value. This is thought to be the most common mechanism of action for natural selection because most traits do not appear to change drastically over time. Stabilizing selection commonly uses negative selection to select against extreme values of the character. Stabilizing selection is the opposite of disruptive selection. Instead of favoring individuals with extreme phenotypes, it favors the intermediate variants. Stabilizing selection tends to remove the more severe phenotypes, resulting in the reproductive success of the norm or average phenotypes. This means that most common phenotype in the population is selected for and continues to dominate in future generations.
Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avoiding detection, warding off attack, fighting back, or escaping when caught.
The viceroy is a North American butterfly. It was long thought to be a Batesian mimic of the monarch butterfly, but since the viceroy is also distasteful to predators, it is now considered a Müllerian mimic instead.
A morphogen is a substance whose non-uniform distribution governs the pattern of tissue development in the process of morphogenesis or pattern formation, one of the core processes of developmental biology, establishing positions of the various specialized cell types within a tissue. More specifically, a morphogen is a signaling molecule that acts directly on cells to produce specific cellular responses depending on its local concentration.
Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.
Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.
Limenitis arthemis, the red-spotted purple or white admiral, is a North American butterfly species in the cosmopolitan genus Limenitis. It has been studied for its evolution of mimicry, and for the several stable hybrid wing patterns within this nominal species; it is one of the most dramatic examples of hybridization between non-mimetic and mimetic populations.
Compartments can be simply defined as separate, different, adjacent cell populations, which upon juxtaposition, create a lineage boundary. This boundary prevents cell movement from cells from different lineages across this barrier, restricting them to their compartment. Subdivisions are established by morphogen gradients and maintained by local cell-cell interactions, providing functional units with domains of different regulatory genes, which give rise to distinct fates. Compartment boundaries are found across species. In the hindbrain of vertebrate embryos, rhobomeres are compartments of common lineage outlined by expression of Hox genes. In invertebrates, the wing imaginal disc of Drosophila provides an excellent model for the study of compartments. Although other tissues, such as the abdomen, and even other imaginal discs are compartmentalized, much of our understanding of key concepts and molecular mechanisms involved in compartment boundaries has been derived from experimentation in the wing disc of the fruit fly.
In zoology, automimicry, Browerian mimicry, or intraspecific mimicry, is a form of mimicry in which the same species of animal is imitated. There are two different forms.
Decapentaplegic (Dpp) is a key morphogen involved in the development of the fruit fly Drosophila melanogaster and is the first validated secreted morphogen. It is known to be necessary for the correct patterning and development of the early Drosophila embryo and the fifteen imaginal discs, which are tissues that will become limbs and other organs and structures in the adult fly. It has also been suggested that Dpp plays a role in regulating the growth and size of tissues. Flies with mutations in decapentaplegic fail to form these structures correctly, hence the name. Dpp is the Drosophila homolog of the vertebrate bone morphogenetic proteins (BMPs), which are members of the TGF-β superfamily, a class of proteins that are often associated with their own specific signaling pathway. Studies of Dpp in Drosophila have led to greater understanding of the function and importance of their homologs in vertebrates like humans.
Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male has strong patterns, conspicuous colours and is iridescent, while the female is far less visible.
Bicyclus anynana is a small brown butterfly in the family Nymphalidae, the most globally diverse family of butterflies. It is primarily found in eastern Africa from southern Sudan to Eswatini. It is found mostly in woodland areas and flies close to the ground. Male wingspans are 35–40 mm and female wingspans are 45–49 mm.
Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey animal an opportunity to escape. The term deimatic or dymantic originates from the Greek δειματόω (deimatóo), meaning "to frighten".
The evo-devo gene toolkit is the small subset of genes in an organism's genome whose products control the organism's embryonic development. Toolkit genes are central to the synthesis of molecular genetics, palaeontology, evolution and developmental biology in the science of evolutionary developmental biology (evo-devo). Many of them are ancient and highly conserved among animal phyla.
In evolutionary biology, mimicry in vertebrates is mimicry by a vertebrate of some model, deceiving some other animal, the dupe. Mimicry differs from camouflage as it is meant to be seen, while animals use camouflage to remain hidden. Visual, olfactory, auditory, biochemical, and behavioral modalities of mimicry have been documented in vertebrates.