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Las preferencias de alimentacin de Morpho peleides en el Jardn de Mariposas Monteverde
Feeding preferences of captive Morpho peleides at the Monteverde Butterfly Garden
Morpho peleides is one of many tropical butterfly species that are fruit-feeding butterflies. In this paper, I will (1) explain the importance of biochemical components in banana pulp including amino acids, sugar, and alcohol in the diet of M. peleides and how it affects the butterflies food choices, and (2) suggest how fermentation might encourage or discourage food preferences banana pulp with high or low concentrations of alcohol. Six combinations were tested in this study: mature pulp with amino acids, mature pulp with water, mature pulp with rum, overripe pulp with water, overripe pulp with amino acids, and overripe pulp with rum. There was no significant difference among average feeding durations (ANOVA: F ratio = 1.60; p value = 0.16; df = 5) or among frequencies of visits between each food type (Chi-square Goodness of Fit: 2 = 8.77, p > 0.05). The similarity between visit times and frequencies for banana pulp samples with higher concentrations of amino acids and rum can be explained by (1) previous research suggesting that it is more common for nectar-feeding butterflies to have preferences for amino acids than it is for fruit-feeding butterflies like M. peleides, and (2) nutritional compensation of overripe bananas via (a) more amino acids released from the peel into the pulp or (b) higher sugar concentration peaks later on in the maturation process combined with stronger odor cues that attract the butterflies.
Morpho peleides es una de las muchas especies de mariposas tropicales que comen frutas. En este reporte explicar (1) la importancia de los compuestos bioqumicos en la pulpa de banano incluyendo el amino cido, la azcar, y el alcohol en la dieta de M. peleides y como le afecta su eleccin de diferentes tipos de comida, y (2) sugiere en como el proceso de fermentacin puede animar o disuadir las preferencias de pulpa de banano con concentraciones altas o bajas en alcohol.
Text in English.
Tropical Ecology 2007
Ecologa Tropical 2007
Mariposas que se alimentan de frutas
Mariposas que se alimentan de nctar
t Monteverde Institute : Tropical Ecology
Feeding preferences of captive Morpho peleides at the Monteverde Butterfly Garden Beth Wright . Biology Department, University of Puget Sound ABSTRACT Morpho peleides is one of many tropical butterfly species that are fruit feeding butterflies. In this p aper, I will 1 explain the importance of biochemical components in banana pulp including amino acids, sugar, and alcohol in the diet of M. peleides and how it affects the butterflies food choices, and 2 suggest how fermentation might encourage or disc ourage food preferences banana pulp with high or low concentrations of alcohol . Six combinations were tested in this study: mature pulp with amino acids, mature pulp with water, mature pulp with rum, overripe pulp with water, overripe pulp with amino acids , and overripe pulp with rum. There was no significant difference among average feeding durations ANOVA: F ratio = 1.60; p value = 0.16; df = 5 or among frequencies of visits between each food type Chi square Goodness of Fit: Ã°C 2 = 8.77, p > 0.05. The similarity between visit times and frequencies for banana pulp samples with higher concentrations of amino acids and rum can be explained by 1 previous research suggesting that it is more common for nectar feeding butterflies to have preferences for amin o acids than it is for fruit feeding butterflies like M. peleides , and 2 nutritional compensation of overripe bananas via a more amino acids released from the peel into the pulp or b higher sugar concentration peaks later on in the maturation process combined with stronger odor cues that attract the butterflies. RESUMEN Morpho peleides es una de muchas especies de mariposas tropicales que comen frutas. En este reporte explicarÃ© 1 la importancia de los compuestos bioquÃmicos en la pulpa de banano c omo amino Ã¡cidos, azÃºcar, y alcohol en el dieta de M. peleides y como lo afecta su elecciÃ³n de diferentes tipos de comida, y 2 explorar como el proceso de fermentaciÃ³n puede animar o disuadir la elecciÃ³n por comida con mÃ¡s o menos alcohol. Se probaron se is combinaciones en este estudio: pulpa de banano s m aduros con agua, pulpa de banano s maduro s con proteÃna, pulpa de banano s maduro s con alcohol, pulpa de banano s mÃ¡s maduro s con agua, pulpa de banano s mÃ¡s maduros con proteÃna, y pulpa de banano s mÃ¡s madu ros con alcohol. No hubo una diferencia considerable entre el tiempo de alimentaciÃ³n entre los tipos de comida o entre la frecuencia de visitas a los diferente s tipos de comida. Esta homogeneidad entre los tiempos en los diferentes tipos de comida y frecuen cias de visitas a pulpa con mÃ¡s alcohol y mÃ¡s proteÃna se puede explicar de dos maneras. 1 Proyectos anteriores sugieren que es mÃ¡s comÃºn para mariposas que comen nÃ©ctar a preferir comida con proteÃna que para mariposas que comen frutas y 2 hay una com pensaciÃ³n de nutricional en la pulpa de bananos mÃ¡s maduros en dos maneras: a mÃ¡s proteÃna fue transportado a la pulpa de la cÃ¡scara y b mÃ¡s azÃºcar en la fruta durante fases mÃ¡s tarde en el proceso de maduraciÃ³n en combinaciÃ³n con olores mÃ¡s fuerte s que atraen las mariposas. INTRODUCTION Tropical butterflies can feed on a number of sources such as nectar from flowers, and other sugar and mineral rich sources like different types of fruit, sap from trees, mud, carrion, dung, and pollen Molleman et al. 2 005. One of the largest feeding guilds in butterfly species is the fruit feeding guild, which includes Morpho peleides , commonly known as the Blue Morpho butterfly. In this paper, I will 1 explain the importance of different biochemical components in ba nana pulp including amino acids, sugar, and alcohol in the diet of M. peleides and how it affects the butterfliesÂ€ food choices, and 2 suggest how fermentation might encourage or discourage food preferences between banana pulps with high or low concentrat ions of alcohol .
In contrast to nectar, fruit is known to contain higher amounts of necessary proteins, creating less dependence on the larval food reserves by fruit feeding butterflies than the nectar feeding butterflies Molleman et al. 2005. It was su ggested that this difference could affect longevity, but there has been little supportive evidence of such a relationship in fruit feeding butterflies. In contrast, there has been some evidence to the contrary demonstrating that neither longevity nor egg p roduction was influenced by proteins in the diets of Bicyclus anynana , a fruit feeding butterfly species Molleman et al. 2005. Tropical fruit feeding butterflies typically feed on overripe or decaying fruit that are found on the ground in forests Molle man et al. 2005. This could be due to the increase in sugar concentrations followed by a rapid decrease as the ripening process occurs. As the amount of starch declines in bananas during ripening, the concentrations of sugar increase due to chemical conve rsions Emaga et al. 2007. Varying sugar concentrations are confirmed by Zhang et al. 2005, who determined that pulp from a green banana contains 1.23% sucrose, which increases to 53.2% sucrose in a fully ripe banana. This conversion from starch to high er amounts of sugar could enable a female butterfly to acquire lacking amino acids, and the differentiation between food choices regarding sugar concentrations may be based on a fitness decision in an effort to gain more proteins. Alternatively, the nitrog en content of the same fruit, required for the production of proteins and nucleic acids, was higher than known nitrogen amounts in nectar from flowers Molleman et al. 2005. Fruits, like bananas, also contain secondary compounds, alcohols Molleman et al. 2005, sugar, starch, fiber, minerals such as potassium Emaga et al. 2007, and amino acids such as leucine, valine, phenylalanine and threonine in consistent concentrations throughout fruit maturation Emaga et al. 2007, which are often used by female butterflies for egg production O'Brien et al. 2003 . As bananas mature, the amount of ethylene alcohol increases as well Taiz & Zeigler 1991 via fermentation, a result of anaerobic processes Biale 1964 and degradation processes. Fermentation should l ogically create shorter feeding spans, as it has been suggested that butterflies can become inebriated from high amounts of alcohol Molleman et al. 2005, and might avoid food sources with that component. In this study, I examined the food preferences o f M. peleides based on two sources of banana pulp mature bananas and overripe bananas, as they have vastly different concentrations of alcohol due to natural fermentation. I also added rum to increase the alcohol concentration in the samples and amino a cids to supplement the pulp with more nutrients to samples from each source to determine if there was a preference between different combinations. Since my hypothesis was that M. peleides could recognize variances between the nutritional values of each f ood combination to choose pulp samples accordingly, I expected to see differences in both number of visits and time spent feeding at each sample. I predicted that the visits to the pulp samples with rum and the pulp samples from the overripe bananas would have shorter and less frequent visits than the other samples, as Molleman et al. 2005 noted that butterflies generally do not like high amounts of alcohol. Additionally, I predicted that since amino acids provide necessary proteins to some species of but terflies, there would be longer and more frequent visits to the samples with amino acids. In decreasing order, I expected to see
longer visits to mature pulp with amino acids, mature pulp, mature pulp with rum, overripe pulp with amino acids, overripe pulp , and overripe pulp with rum. MATERIALS AND METHODS Study Site The study was conducted at the Monteverde Butterfly Garden, in Monteverde, Costa Rica. There were approximately 70 adult individuals of M. peleides within the net enclosed area that were reare d in captivity. The butterflies were fed banana pulp prior to each of the six observation days. Food Preparation The peels of two mature bananas and three overripe bananas were removed and the fruit was mashed with to a pulp consistency and stored in tw o separate, labeled jars. All sample mixtures of mature and then overripe bananas were mashed thoroughly to ensure homogeneity. Five ml of water, rum, or amino acids were added to 45 ml of both a mature banana pulp sample and 45 ml of an overripe banana pu lp sample, making six combinations: mature pulp with water 10% by volume or 3.6 g of water and 90% by volume or 43.31 g of banana, mature pulp with amino acids 10% by volume or 3.08 g amino acids and 90% by volume or 44.13 g of banana, mature pulp with rum 10% by volume or 4.61 g of rum and 90% by volume or 45.9 g of banana, overripe pulp with water, overripe pulp with amino acids, and overripe pulp with rum. Observations One set of three mature banana samples mature pulp with water, mature pulp wi th amino acids, and mature pulp with rum and one set of three overripe samples overripe pulp with water, overripe pulp with amino acids, and overripe pulp with rum were set on plastic lids, with the mature samples consistently on the left and overripe s amples consistently on the right Figure 1. Additionally, a replica of each set as described above was located on two nearby leaves. Prior to collecting data, all other bananas in the butterfly garden, typically left out for butterfly feeding, were remo ved for the duration of the observations to ensure that butterflies would have no other feeding options apart from the pulp. Since the juice from the banana pulp samples did travel from the point of origin, visits were recorded if the butterflies were feed ing on the juices from identifiable specific pulp samples. Data collection occurred for five hours in mornings from 9:30 AM to 1:30 PM due to peak butterfly activity during that time period. Two types of data were collected. First, for each feeding visit b y a single individual, the start time and stop time were recorded as accurately as possible to calculate the amount of time per visit. On some occasions, estimates were made based on local position on the leaves or plates. Sample sizes for the time based data on the six food types ranged from 15 visits for mature pulp with water to 46 visits for overripe pulp with rum. Alternatively, the frequency of feeding visits were recorded for each pulp sample by utilizing scan sampling every five minutes to record t he amount of butterflies present on each sample. Sample sizes for the frequency based data on six food types ranged from 28 visits for mature pulp with water to 46 visits to mature pulp with rum Table 1. Three days of data collection were completed for e ach of the two types of data, and after over 100 visits observed for frequency data, observations were switched to timed visits. Statistics An ANOVA was run to compare the average lengths of time M. peleides spent feeding on each of the six different foo d types. Additionally, a Chi Square Goodness of
Fit analysis was conducted on the frequency of visits to each of the six pulp types to determine if some pulp samples were visited more frequently than others. RESULTS There was no significant difference in t he average feeding times between the six locations Figure 2, though the range of time spent feeding at the banana pulp ranged from less than one minute to 47 minutes at mature pulp with water. Additionally, though there was no significant difference in f requencies of visits between the banana pulp samples, the number of visits received for each food type ranged from 28 to 43 Table 1. 0 2 4 6 8 10 12 14 Mature Banana Pulp with Water Mature Banana Pulp with Amino Acids Mature Banana Pulp with Rum Overripe Banana Pulp with Water Overripe Banana Pulp with Amino Acids Overripe Banana Pulp with Rum Type of Food Average Feeding Time min n = 15 n = 29 n = 33 n =33 n = 25 n = 46 Figure 2. Average time spent feeding on six different combinations of banana pulp: mature bana na pulp with water, mature banana pulp with amino acids, mature banana pulp with rum, overripe banana pulp with water, overripe banana pulp with amino acids, and overripe banana pulp with rum. Timed observations were run at the Monteverde Butterfly Garden. There was no significant difference in the average feeding times between the six food types ANOVA: F ratio = 1.60; p value = 0.16; df = 5. Therefore, the butterflies were not spending more or less time on a given type of food. Sample sizes visits for each food type were as follows: n = 15 Mature Banana Pulp with Water, n = 29 Mature Banana Pulp with Amino Acids, n = 33 Mature Banana Pulp with Rum, n = 33 Overripe Banana Pulp with Water, n = 25 Overripe Banana Pulp with Amino Acids, n = 46 Ov erripe Banana Pulp with Rum. Standard error bars are shown. Table 1. Amount of feeding visits received by each food type. There were no significant differences between the frequencies of visits to each of the food types Chi square Goodness of Fit: Ã°C 2 = 8.77, p > 0.05.
Food Type Frequency of Visits Mature banana pulp with water 28 Mature banana pulp with amino acids 29 Mature banana pulp with rum 46 Overripe banana pulp with water 34 Overripe banana pulp with amino acids 29 Overripe banana pulp with rum 43 DISCUSSION My hypothesis regarding M. peleides detecting varying nutritional values and cho osing a food source accordingly was not supported by my results. Morpho peleides butterflies did not spend a longer or shorter length of time on a given type of food as was predicted Figure 2. Additionally, they did not visit any specific pulp sample wit h higher or lower frequencies Table 1. The results indicate that amino acids were not favored nor were the overripe samples with rum avoided Figure 2. There are multiple possibilities for this outcome. First, amino acids may not have been preferred be cause female M. peleides individuals either had received enough nutrition as larvae or because amino acids do not affect fecundity in this particular species. Referring to the former reason, the M. peleides individuals at the Monteverde Butterfly Garden li kely received sufficient amounts of food and high quality nutrients during their larval stage. In regards to the latter reason, Mevi Shutz & Erdhart 2003 noted that there was a female preference for amino acids in some species for nectar feeding butterfl ies and that food choice in these species was dependent on larval food quality caterpillars, like other herbivores gain nutrients by consuming various plant parts Mattson 1980 of their host plant. Additionally, some female butterflies oviposit their eg gs on the same plant that they feed on Janz et al. 2005 indicating that perhaps there is a dietary quality in the plant like amino acids that the butterflies can detect and actively decide to lay eggs on that plant. Since M. peleides are fruit feeding butterflies, it is less likely that female individuals would exhibit a preference for amino acids due to the observation that preferences for amino acids seems to be found only in nectar feeding butterflies. This is supported by the fact that relationships between amino acids in adult butterfly diets, longevity and egg production have not been found for fruit feeding butterflies Molleman et al. 2005. Second, banana peels contain many nutrients such as amino acids that may seep into the banana pulp as th e outer peel erodes away Emaga et al. 2007. Since the overripe bananas had the peels on longer than the mature bananas, the overripe bananas may have gained more amino acids from the peel than the mature bananas. This increase in amino acids in overripe bananas could balance out the decreased appeal of the higher concentrations of alcohol, by adding enough nutritional value to the overripe bananas so that visitation lengths of times and frequencies between the banana pulp samples were more equivalent. H owever, it appears that amino acids are not attractive by themselves as there was no preference for this in overripe samples or amino acid samples, refuting the banana peel theory. Third, as noted by Molleman et al. 2005, butterflies use cues such as foo d odors to find food. Stronger odors of alcohol via fermentation present stronger signals than food sources with less alcohol Molleman et al. 2005. The previous suggestion by Molleman
et al. 2005 that fermented fruits causes butterflies to become inebr iated possibly making fermenting fruits less desirable as food is contrary to the results presented in this paper. Molleman et al. 2005 also found that mature bananas did receive a large amount of feeding visits, but not as many visits as the overripe, fermenting banana reportedly due to volatiles likely ethylene in the fruits. Molleman et al. 2005 s results support the notion that there is some biochemical compound in overripe bananas that make it attractive, even with the less desirable higher co ncentration of alcohol. This additional important compound could be sugar, and the combination of higher amounts of sugar and alcohol could explain the compensation demonstrated in longer and more feeding times than expected for the less desirable possib ility of becoming inebriated. Biologically, then, the butterflies are attracted to the strong odors and feed on the relatively higher amounts of sugar. As a side note, there is a weaker odor in mature pulp with rum than in the overripe pulp due to the earl y stage of maturation and a relatively smaller amount of sugar. Therefore, the hypothesis that butterflies would make decisions between food choices based on detectable nutritional values was not supported in this study, but it is logical to think that cho osing food sources with higher sugar and protein concentrations would prove beneficial for egg production O'Brien et al. 2003 , longevity as a result of adult dietary intakes Mevi Schutz & Erdhardt 2005, body size, and development, especially if the but terfly did not receive enough nutrition as a larva Mevi Schutz & Erdhardt 2005; Mevi Shutz & Erdhart 2003. In the future, it would prove beneficial to address questions to increase the understanding of how M. peleides decides which food sources to consum e, such as 1 how amino acid supplements affect fecundity or longevity in M. peleides to determine if they are foraging adaptively as well as 2 examining how different odors affect the ability of M. peleides to detect food. There is a sufficient amoun t of information still unknown about butterfly feeding choices. The answers to these questions could assist researchers in determining what exactly is the deciding factor in food choices, which could prove to be important for butterfly conservation. Ackno wledgements Thanks to Karen and Alan Masters for the inspiration for this project. Thanks to Mark Wainwright who led a butterfly diversity field problem in Santa Rosa, which encouraged me to conduct research with butterflies. Thanks to Jim Wolfe and the Mo nteverde Butterfly Garden for letting me conduct my project there. Thank you to the staff at the Monteverde Butterfly Garden for being so friendly. Thanks to Karen Masters, Camryn Pennington and Kalle Larson for reading drafts and offering suggestions for improvement. Thanks to Pablo Allen for helping me get the Power Point presentation ready. Thanks to the EstaciÃ³n BiolÃ³gi c a de Monteverde for access to computers and the hospitality. Literature Cited: BIALE, J. 1964. Growth, maturation, and senescence in fr uits. Science : New Series. 146: 3646. 880 888. EMAGA, T., R. ANDRIANAIVO, B. WATHELET, J. TCHANGO, AND M. PAQUOT. 2007. Effects of the stage of maturation and varieties on the chemical composition of banana and plantain peels. Food Chemistry. 103: 590 600 . JANZ, N., A. BERGSTROM, AND A. SJOGREN . 2005. The role of nectar sources for oviposition decisions of the common blue butterfly Polyommatus icarus. Oikos. 109 : 3: 535 538. MATTSON, W. 1980. Herbivory in Relation to Plant Nitrogen Content. Annual Revie w of Ecology and Systematics . 11. pp. 119 161 .
MEVI SCHUTZ, J., AND A. ERDHARDT. 2003. Larval nutrition affects female nectar amino acid preference in the Map Butterfly Arashchnia levana . Ecology. 84 10:2788 2794. MEVI SCHUTZ, J.AND A.ERDHART. 2005. Amino acids in nectar enhance butterfly fecundity: A long awaited link. The American Naturalist. 165: 4. MOLLEMAN, F. M. VAN ALPHEN, P. BREAKEFORD, AND B. ZWAAN. 2005. Preferences and food quality of fruit feeding butterflies in Kibale Forest, Uganda. Biotropica 37: 4. 657 663 O'BRIEN, D., C. BOGGS, AND M. FOGEL. 2003. Pollen feeding in the butterfly Heliconius charitonia : isotopic evidence for essential amino acid transfer from pollen to eggs. Proceedings: Biological Sciences . 270: 2631 2636. TAIZ, L. AND E. ZEIGLER. 1991. Plant Physiology . Benjamin/ Cummings Publishing Company Inc. USA. ZHANG, P., R. WHISTLER, J. BEMILLER, AND B. HAMAKER. 2005. Banana Starch: production, physiochemical properties and digestibility a review. Carbohydrates Polymers. 59 : 443 458. Appendix Figure 1. Diagram of banana pulp samples. The left plate is from mature bananas while the right plate is from overripe bananas. M = Mush with water, O = Mush with water, MAA = Mush with amino acids, OAA = Mush with amino acids, MR = Mush with rum, OR = Mush with rum. M MAA O OAA OR MR