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Heliconius Wikipedia Page Edits
I added my edits to the Heliconius butterfly page https://en.wikipedia.org/wiki/Heliconius

I made my changes under different sections. I made the section titled Cyanic Characteristics and the section titled Sexual Selection and aposematism

Cyanic Characteristics
In order to be unpalatable, the Heliconius butterflies use cyanic characteristics, meaning they produce substances that have cyanide group attached to them, ultimately making them harmful. Research has found that the amino acids needed to make the cyanic compounds comes from feeding on pollen. Although feeding on pollen takes longer than nectar feeding, the aposematic characteristics help to warn predators away and gives them more time to feed. While the Heliconius larva feed on the Passifloraceae plant, which also has cyanic characteristics, the larva have evolved the ability to neutralize the cyanic molecules to protect them from the negative effects.

Sexual selection and aposematism
In order for the aposematism and mimicry to be successful in the butterflies they must continually evolve their colors to warn predators of their unpalatability. Sexual selection is important in maintaining the aposematism as it helps to select for specific shades of colors rather than general colors. It is important to select for specific colors to avoid subtle shades in any of the species involved in the mimicry. If any colors are not successful in their warning it will negatively affect the success of the aposematism because it cannot warn predators as efficiently. In order to select for specific colors, neural receptors in the butterflies’ brains give a disproportionate recognition and selection of those shades. In order to test the importance of these neural and visual cues in the butterflies, researchers conducted an experiment where they eliminated colors from butterflies’ wings. When a color was eliminated, the butterfly was less successful in attracting mates and therefore did not reproduce as much as its counterparts

Mating and Offspring
For this section I changed its title from Pupal mating to mating and offspring. Originally, there was only one sentence and I added more to make it clear that there wasn't just pupal mating in the butterflies.

In this form of mating, the male Heliconius finds a female pupae and waits until a day before she is moulted before he mates. In this form of mating there is no sexual selection present. Heliconius female butterflies also disperse their eggs much slower than other species of butterflies. They obtain their nutrients for egg production through pollen in the adult stage rather than the larval stage. Due to nutrient collection in the adult rather than larval stage, adult females have an extensively longer life compared to others species which allows them to better disperse their eggs for survival and speciation. This form of egg production is helpful because larva are much more vulnerable than adult stages, although they also utilize aposematism. Because many of the nutrients needed to produce eggs are obtained in the adult stage the larval stage is much shorter and less susceptible to predation.

Paper Final Draft
Evolution Leading to Successful Müllerian Mimicry in Heliconius Butterflies In nature, species can use a tactic for survival known as Müllerian Mimicry. This form of mimicry, which was first noted by Fritz Müller in 1878, is used by two or more species that are poisonous or unpalatable and utilize the same warning signal to warn predators. Müllerian mimicry can involve signals that are visual, verbal, scent based or other sensory warnings. Different species adopt different approaches to this form of mimicry, but one that is notable involves the genus of Heliconius butterflies. Understanding the mechanisms involved in the evolution of these butterflies can be used in many applications such as applying this knowledge to general medicine for humans. These butterflies are noted for their diverse range of speciation and success using Müllerian mimicry and their evolutionary steps to get to this state are noteworthy to study.

The Heliconius butterfly is found in many different environments. They are mainly found in tropical and subtropical areas. The different butterflies have their own natural predators, but a main predator is the family of Jacamar bird. In order to evade these predators there are different evolutionary forces at play which include sexual selection, natural selection, mutations, runaway signal selection and others which help to improve their fitness and survival.

One of the primary characteristics involved in this Müllerian mimicry is the use of aposematism, or warning colors, as a visual cue for predators. In order for this form of warning to work the butterflies must have colors that successfully warn predators of their unpalatability. The color of the butterflies is actually very specific in selection. Sexual selection is important in the aposematic color as it selects not only for a basic color, but rather a specific shade that works the best to caution predators. The colors that have survived through time have been studied to show that this evolution occurred partially through sexual selection when mating (Llaurens et al. 2014). It is important to select for specific colors to avoid subtle shades in any of the species involved in the mimicry. If any colors are not successful in their warning it will negatively affect the success of the aposematism because it cannot warn predators as efficiently. Vane-Wright and Ackery (1984) found that receptors in the butterflies’ brains give a disproportionate selection to specific color shades. This neural recognition system that came about through evolution gives the butterflies a selective advantage when mating (Vane-Wright and Ackery 1984). Learning about these neural mechanisms can allow scientists to apply the same concepts to neurology in humans to see if receptions can occur the same way as they do in these butterflies.

In testing the importance of these visual cues an experiment was completed utilizing different species of Heliconius butterflies, all of the butterflies showed preference in finding a mate within their own species. The study used techniques to eliminate specific colors from wing pattern to test their ability to attract mates. When a color was eliminated, the butterfly was less successful in attracting mates and therefore less fit (Mavarez et al. 2006). The researchers found that pheromones did not play a role in selection and that it was mainly the color component of the butterflies (Mavarez et al. 2006).This intersexual selection occurs in both males and females when mating which can make the process more selective in both genders. The mechanism involved in this type of sexual selection of mates can be applied to better understand recognition ability in other organisms.

In order for an aposematism to be successful the predator must learn from the warning signal. When aposematic coloration first appeared for the butterflies it was most likely negative for the species because bright colors can actually attract predators more easily until they learn that the color indicates a negative outcome (Prezeczek et al. 2008). In order for the aposematism to be successful there has to be a stronger degree of unpalatability in the butterflies and in their mimics (Prezeczek et al. 2008). This stands true for a few reasons. First, for the aposematism to work the signal needs to stick in the predators mind. They either need to decrease the fitness of the predators that eat the butterflies, so that those that do not eat them will pass on offspring that also avoid the butterflies or, the outcome needs to be strong enough that the predator learns after one or a few events to avoid that prey. The success of these colors in warning predators was studied using different morphs with the Jacamar bird. To test the success of the mimicry, Langhan (2004) studied the ability of Jacamars to recognize the butterfly’s native to their habitat and mimics from other locations. The local butterflies were similar in phenotype to novel butterflies, those taken from different habitats, so that they represented the mimicry seen in nature. The study measured how many of the local butterflies versus the novel butterflies the Jacamars consumed. It was found that although the Jacamars had never seen the novel morphs they only attacked 37% of the novel forms showing that the mimicry had worked (Langhan 2004). In order for the mimicry to work both of the aposematism’s need to be strong so that the predator does not get a palatable butterfly one time and an unpalatable the next because it will take longer for it to learn or it will not learn at all.

In order for the aposematic coloration to work the extent of unpalatability needs to be memorable. How this occurs is important to study to be able to apply to other species. In order to be unpalatable, the Heliconius butterflies use cyanic characteristics, meaning they produce substances that have cyanide group attached to them, ultimately making them harmful. When studying the production of these cyanic molecules, researchers found that the amino acids needed to make the cyanic compounds came from pollen feeding rather than their host plant, the Passifloracea which is also cyanic (Narstehdt and Davis 1980). Pollen feeding is an adaptation that helps contribute to this mimicry. Feeding on pollen takes much longer than nectar feeding. Because of the needed time to feed they are more susceptible to predators, but because of the success of the mimicry and aposematism they do not need to worry about being exposed. Their aposematic characteristics are what allows them to take more time to feed, because they are less likely to be attacked by predators (Gilbert 1972). The Heliconia are noted to have evolved with their host plants, the Passifloraceae. The plant secretes cyanic chemicals and Heliconius butterflies have the ability to neutralize the plants cyanic characteristics so that they are no longer harmful. This prevents the butterfly from having to compete for a host plant. Helicnoia have utilized this to be able to lay their eggs on the plant so their larvae are safe due to the fact that the plant is unpalatable and is less likely to be consumed. It then improves the possibility of offspring survival (Price et al. 1991). All of these small adaptations applied to feeding and offspring contribute to the success of the mimicry and can give insight into better understanding evolutionary forces. Understanding the mechanisms involved in the butterfly’s ability to neutralize cyanic effects can be utilized by humans in medicine to be able to counteract poisons such as cyanide. The Heliconius butterfly has also adapted to use an alternate form of mating to increase reproductive success. When the butterflies mate they participate in either pupal mating or “normal mating”. Pupal mating involves the male Heliconius finding a female pupae and waiting until a day before she is moulted before he mates. The interesting part about the pupal mating is that the males cannot participate in sexual selection especially for colors because they cannot see the female. However, the butterflies that mate normally versus pupal mating are both very similar phenotypically. When the female lay’s her eggs and the larva are present they also use aposematism which can help to increase their survival. The use of aposematism in the larvae also helps to increase the survival of the butterflies. Heliconius female butterflies also disperse their eggs much slower than other species of butterflies. They obtain their nutrients for egg production through pollen in the adult stage rather than the larval stage. Due to nutrient collection in the adult rather than larval stage, adult females have an extensively longer life compared to others species which allows them to better disperse their eggs for survival and speciation (Gilbert 1972). This form of egg production is helpful because larva are much more vulnerable than adult stages, but because many of the nutrients needed to produce eggs are obtained in the adult stage the larval stage is much shorter and less susceptible to predation(Gilbert 1972). The intake of nutrients and how they are maximally utilized can also be applied to humans and understanding nutrition uptake and energy usage and storage.

Helicnoius butterflies are distinguished from other organisms that use similar mechanisms because of their success. Research focuses on the Heliconius butterflies because of the amount of speciation seen. Understanding the combination and importance of each of these evolutionary forces is a key area of research. The aposematism combined with Müllerian mimicry is so successful that when a new butterfly with similar wing coloration was put in with a new predator it was less likely to get eaten even though the predator had never seen the prey before (Prezecezek 2008). Due to the fact that color changes arise through mutation or selection the new color morphs are rare. It would be thought that these rare morphs would be selected against due to the already wide range of species. Mallet and Singer (1987) point out that selection against these rare forms can occur due to frequency-dependent selection. This selection means that the success, or fitness of a species or phenotype is affected by its frequency in comparison to other species or similar morphs. Mallet and Singer (1987) say that that the more rare form has a stronger negative selection, but they point out that this is not the case since rare morphs are often very successful. The reason that the rare morphs are successful rather than being eliminated due to frequency-dependent selection is because the rare color morphs are less likely to be tested and killed by predators (Sillen-Tullberg and Bryant 1982). However, this is only the case until the rare morph becomes more common in which case it is more likely to be tested (Mallet and Singer 1987). At this point the morph is no longer rare and combines as another common species in the Heliconius genus. When a rare morph comes around it is usually due to a mutation. Mallet and Singer (1987) gave the approach of runaway signal evolution which they compare to R.A. Fisher and runaway sexual selection in mating. In this form of evolution, a certain characteristic, such as aposematic coloration, is strongly selected for. In this example, rare morphs and aposematic color are chosen for since the characteristic helps to avoid predators. Therefore, when a new mimic comes about it can increase in proportion to the other forms and expand via runaway signal evolution (Mallet and Singer 1987). Once it reaches the same proportion it will be sampled as frequently as the other morphs and the runaway signal evolution will slow down. Because of the success of the Müllerian mimicry this can happen more often leading to the increased fitness and speciation seen.

Heliconius butterflies have a wide range of speciation that all play a part in a successful Müllerian mimicry. Many adaptations that make the mimicry successful have come about through evolution. These different evolutionary forces including sexual selection, mating adaptations, aposematism, and runaway signal evolution all contribute to the success of Heliconius butterflies. Understanding how these forces combine to lead to the success of the butterflies can be applied to other organisms or in better understanding molecular processes which can be applied to humans and general medicine.

References

Gilbert L.E. 1972. Feeding and Reproductive Biology of Heliconius Butterflies. Proc. Nat. Acad. Sci. 69(6):1403-1407 Langhan G.M., 2004. Specialized Avian Predators Repeatedly Attack Novel Color Morphs of Heliconius Butterflies. Evolution. 58, 12:2783-2787.

Llaurens V, M Joron, and M. Thery. 2014. Cryptic differences in colour among Mullerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?. J. Evol. Biol. 27:531-540.

Mallet J, M.C. Singer. 1987. Individual selection, kin selection, and the shifting balance in the evolution of warning colours: the evidence from butterflies. Biological Journal of the Linnean Society. 32:337-350.

Mavarez J, C Salazar, E Bermingham, C Salcedo, C Jiggins, M Linares. 2006. Speciation by hybridization in the Heliconius butterflies. Nature 441:868-871

Nahrstedt A, R.H. Davis. 1980. The occurrence of the cyanoglucosides linamarin and lotaustralin, in Acraea and Heliconius butterflies. Comp. Biochem. Physiol. 68B:575-577. Prezeczek K, C. Mueller, and S.M. Vamosi. 2008. The evolution of the aposematism is accompanied by increased diversification. Integrative Zoology. 3:149-156.

Price P.W., T.M. Lewinsohn, G.W. Fernandes, W.W. Benson eds. 1991. Plant- Animal Interactions: Evoltionary Ecology in Tropical and Temperate Regions. John Wiley and Sons, Inc, New. York, United States.

Sillen-Tullberg B, E.H. Bryant. 1982. The evolution of aposematic colorations in distasteful prey: an individual selection model. Evolution. 37(5):993-1000.

Vane-Wright R.I, P.R. Ackery eds. 1984. The Biology of Butterflies. Symposium of the Royal Entomological Society of London. Number 11. Academic Press, London, U.K.

Samantha Devenport

Annotated Bibliography
Tuesday 8am Question: What factors contribute to the success of Mullerian mimicry in Heliconius butterflies? Hines, H.M., B.A. Counterman, P. Albuquerque de Moura, M.Z. Cardoso, M. Linares, J Mallet, R.D. Reed, C.D. Jiggings, M.R. Kronforst, and W.O. McMillan. 2011. Wing patterning gene redefines the mimetic history of Heliconius butterflies. PNAS. 108,49:19666-19671. This article is aiming to help and define the phylogenetic tree of the Heliconius butterfly in order to also help better understand the patterns of mimicry. In order to answer these questions they tested genetics of the butterflies via PCR and used different markers to better understand the genes related to color. Two of the species in the Heliconius genus are, erato and melpomene. The article points out that they have almost identical wing patterns but they are rather far apart phylogenetically. Therefore they are similar due to mimicry. They found that on separate occasions the two butterflies have evolved multiple times within different colors to be as similar in appearance as they are today. The study also found that more recently when changes occurred that they occurred very rapidly. This would be important information for my paper because it supports the idea that the mimicry is successful in the butterflies because phenotypic traits are selected for very quickly. A limitation of this study is that they only looked at two of the species of butterfly so it is somewhat limited but the general concepts can be applied to Heliconius as a whole.

Prezeczek K, C. Mueller, and S.M. Vamosi. 2008. The evolution of the aposematism is accompanied by increased diversification. Integrative Zoology. 3:149-156.

This article aims to better understand the effects of aposematism on evolution over time and more specifically the diversification of species. The way they try to answer, or better understand aposematism, is by comparing groups of organisms that expressed aposematism to those that did not and observed diversification within those groups. They found that those in which aposematism was present there was a higher amount of species diversification than in those that did not use aposematism. They also pointed out that the initial cost of being aposematic is negative because the use of bright coloration increases predation until the prey learns that the organism is not palatable. One limit to this paper is that is does not directly reference or study the Heliconius butterfly but the general concept can be applied. This research is helpful to my own as it points out the benefits and also the negatives of aposematism and its effects in the long term. It shows that although there can be initial costs that the long term allows for more diversification and survival. The paper also points out that the degree of the aposematism is important for the organisms. That it is more successful based on the color and unpalatability of the organism. I will use this idea to look at the specific chemical aspects of the butterflies to their predators to see how severe their effects are which would affect the potential success of the aposematism. Llaurens V, M Joron, and M. Thery. 2014. Cryptic differences in colour among Mullerian mimics: how can the visual capacities of predators and prey shape the evolution of wing colours?. J. Evol. Biol. 27:531-540.

This paper is looking to see if there are selections based on specific colors of wings in mimicry. The group used reflectance spectrophotometry to detect differences in color. Then they set up different models for animal and butterfly vision to see how they perceive the color and used models of photoreceptors in their eyes. The group monitored not only visuals from predators but also mating patterns. One main interesting piece of information they found is that the butterflies are also selective for color when mating. They are also adapting colors to avoid predators because if the color is too subtle in one aposematic species then it will not work as well throughout the mimicry. This paper will be important for mine to show that the Mullerian mimicry along with the aposematism is being very selective as to even choose for different color shades rather than a general color. This is making the butterflies specialized to help them avoid prey as best as possible. Mallet J, and L.E. Gilbert. 1994. Why are there so many mimicry rings? Correlations between habitat behavior and mimicry in Heliconius butterflies. Biological Journal of the Linnean Society. 55:159-180.

This article questions if there have been any evolutional changes in color of wings in Heliconius butterflies that correlates with their habitat or behaviors. In order to answer this question the researchers measured different characteristics such as roosting height, flying height, where they spent time in the say versus the night and more. They did this by counting the number of butterflies and using vegetation markings to measure height. This study is important because it shows the use of mimicry in different habitats. Although the butterflies are essentially in the same environment they have still taken on the mimicry when they inhabit different areas. This shows that the mimicry can be successful in different aspects of living. The research showed that some behaviors were not different between species but that they utilized the mimicry in different ways such as at night versus the day time. I can use this source because I can relate the success of mimicry being used in different habitats which means it can be used in a broader sense rather than only in specific situations.

Langhan G.M., 2004. Specialized Avian Predators Repeatedly Attack Novel Color Morphs of Heliconius Butterflies. Evolution. 58, 12:2783-2787.

This article is aiming to see the effects that the Mullerian mimicry in the butterflies can have on predators. In order to understand if this occurs, Langhan collected jacamar birds along with Heliconius butterflies. Some butterflies were selected from the jacamars natural habitat and others from different areas. The local butterflies were similar in phenotype to novel butterflies, those taken from different habitats. She measured how many of the local butterflies versus the novel butterflies that the jacamars consumed. They did not eat any of the local butterflies and only ate about 37% of the novel ones although they did not look exactly alike. This is an example that the Mullerian mimicry is working for the butterflies to avoid predators. One limit to this study is that the author was also focusing on the ability of the birds to be selective but it shows that they were able to quickly learn to recognize the butterflies even before they tasted them. This would benefit my paper to show the success that the aposematism has for the butterflies on predators and the benefit that the Mullerian mimicry plays. The ability for novel butterflies to avoid prey that have never actually seen them before shows how successful the mimicry is.

Comments and Suggestions Assignment

 * https://en.wikipedia.org/wiki/Heliconius

Aposematism, using warning colors to denote unpalatable prey, has also been noted to help improve species diversification which may also attribute to the wide range of Heliconius species.
 * Under the pupal mating section it is almost implied from what is written that all Heliconius butterflies mate via pupal mating. It might be better to clarify that it is just a large portion, however not a majority. There is about 42% of Heliconius that are involved in pupal mating . Those that don’t use pupal mating are actually selective for color which contributes to their mimicry success.
 * The habitats that Heliconius butterflies are in would be important to add since there is no specific information and only general regions. They are actually located in different areas which effects how the mimicry works. Some can even live in the same area and there habitat only differs by where they nest and sleep. Even though their habitats are very similar and only differ on elevation they can still show different characteristics adapted to those environments.
 * In the model for evolutionary study tab it might also be important to note that they are also studied based on their convergence for mimcry. Some butterflies, such as Heliconius erato and melpomene have almost identical wing patters but they are very different phylogenetically so it is interesting to study them and the evolutionary mechanisms that made them look so similar in the mimicry.
 * Added sentence and citation