Sugar preference across butterfly families


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Sugar preference across butterfly families

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Title:
Sugar preference across butterfly families
Translated Title:
Preferencia del azúcar a través de las familias de mariposas
Creator:
Olsson, Juliana
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Text in English

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Butterflies ( lcsh )
Mariposas ( lcsh )
Costa Rica--Puntarenas--Monteverde Zone
Costa Rica--Puntarenas--Zona de Monteverde
CIEE Spring 2008
CIEE Primavera 2008
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Reports

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Abstract:
Adult butterflies have a carbohydrate-based diet, acquiring their sugars mostly from nectar or from rotting fruits. Fruit-eating butterflies most often encounter fructose and glucose (Omura and Honda, 2003), while nectarivorous butterflies may encounter hexose-rich nectars in short-corolla flowers or sucrose-rich nectars in long-corolla flowers (Baker and Baker, 1983). This study explores the possibility that butterfly species exhibit certain sugar preferences that reflect not only their diet but their phylogeny as well, since feeding habits are often the result of co-evolution with pollination syndromes in the case of nectar-feeding butterflies. Butterflies from as many species as possible were given a solution of either 20% sucrose by weight or a 20% solution of combined glucose and fructose. The amount imbibed was recorded for the 26 species caught. No obvious preference for either sucrose or hexose nectars were uncovered. Butterflies would drink from either solution, indicating that fruits and flowers offering either sugar would be acceptable to most butterflies and that sugar preference is not as rigidly an evolved trait as was thought. However, minor trends towards sucrose preference are compelling grounds for further study. ( ,,, )
Abstract:
Las mariposas adultas tienen una dieta basada en carbohidratos, consiguen los azucares principalmente del néctar o de los frutos en descomposición. Las mariposas nectarívoras encuentran néctar rico en hexosa en flores con corolas cortas o néctar rico en sucosa en flores con corolas largas. Este estudio explora la posibilidad de que algunas mariposas exhiban alguna preferencia por azucares que refleje no solo su dieta pero también su filogenia, ya que los hábitos alimenticios a veces son el resultado de co-evolución con los síndromes de polinización.
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Student affiliation : Department of integrative biology, University of California at Berkeley
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Born Digital

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Monteverde Institute
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M39-00455 ( USFLDC DOI )
m39.455 ( USFLDC Handle )

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Adult butterflies have a carbohydrate-based diet, acquiring their sugars mostly from nectar or from rotting fruits. Fruit-eating butterflies most often encounter fructose and glucose (Omura and Honda, 2003), while nectarivorous butterflies may encounter hexose-rich nectars in short-corolla flowers or sucrose-rich nectars in long-corolla flowers (Baker and Baker, 1983). This study explores the possibility that butterfly species exhibit certain sugar preferences that reflect not only their diet but their phylogeny as well, since feeding habits are often the result of co-evolution with pollination syndromes in the case of nectar-feeding butterflies. Butterflies from as many species as possible were given a solution of either 20% sucrose by weight or a 20% solution of combined glucose and fructose. The amount imbibed was recorded for the 26 species caught. No obvious preference for either sucrose or hexose nectars were uncovered. Butterflies would drink from either solution, indicating that fruits and flowers offering either sugar would be acceptable to most butterflies and that sugar preference is not as rigidly an evolved trait as was thought. However, minor trends towards sucrose preference are compelling grounds for further study.
Las mariposas adultas tienen una dieta basada en carbohidratos, consiguen los azucares principalmente del nctar o de los frutos en descomposicin. Las mariposas nectarvoras encuentran nctar rico en hexosa en flores con corolas cortas o nctar rico en sucosa en flores con corolas largas. Este estudio explora la posibilidad de que algunas mariposas exhiban alguna preferencia por azucares que refleje no solo su dieta pero tambin su filogenia, ya que los hbitos alimenticios a veces son el resultado de co-evolucin con los sndromes de polinizacin.
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Sugar Preference across Butterfly Families Juliana Olsson Department of Integrative Biology, University of California at Berkeley ABSTRACT Adult butterflies have a carbohydrate based diet, acquiring their sugars mostly from nectar or from rotting fruit s. Fruit eating butterflies most often encounter fructose and glucose Omura and Honda, 2003, while nectarivorous butterflies may encounter hexose rich nectars in short corolla flowers or sucrose rich nectars in long corolla flowers Baker and Baker, 1983 . This study explores the possibility that butterfly species exhibit certain sugar preferences that reflect not only their diet but their phylogeny as well, since feeding habits are often the result of co evolution with pollination syndromes in the case o f nectar feeding butterflies. Butterflies from as many species as possible were given a solution of either 20% sucrose by weight or a 20% solution of combined glucose and fructose. The amount imbibed was recorded for the 26 species caught. No obvious prefe rence for either sucrose or hexose nectars were uncovered. Butterflies would drink from either solution, indicating that fruits and flowers offering either sugar would be acceptable to most butterflies and that sugar preference is not as rigidly an evolved trait as was thought. However, minor trends towards sucrose preference are compelling grounds for further study. RESUMEN Las mariposas tienen una dieta basada en carbohidratos, consiguen los azucares principalmente en la forma fructosa y glucosa, de ne ctar o de frutos en decomposici ón. Las mariposas nectarívoras encuentran néctar rico en hexosa en flores con corolas cortas o néctar rico en sucrosa en flores con corolas largas. Este estudio explora la posibilidad de que algunas mariposas exhiban alguna p referencia por azucares que refleje no solo su dieta pero también su filogenia, ya que los habitos alimenticios a veces son el resultado de co evolución con los sindromes de polinización. Mariposas de varias especies fueron alimentadas con sucrosa al 20% o una solución al 20% de glucosa/fruxtosa. La cantidad consumida fue medida para 26 especies. No se detecto ninguna preferencia. Esto indica que las flores y frutos ofreciendo cualquiera de los azucares serán aceptados por la mayoría de las mariposas, y que la preferencia de azúcar no es un carácter evolutivo tan rigido con se pensaba. INTRODUCTION Most adult butterflies depend o n sugar solutions for their diet, which they get either from nectars Romeis and Wackers, 2002 or juices of rotting fruit Omur a and Honda, 2003. The nectar feeding butterflies are important pollinators of the plants they depend on, and certain pollination syndromes have evolved to ensure a tighter relationship between flower and butterfly Baker and Baker, 1983. While flower sh ape and color are agreed to be important components of pollination syndromes, the importance of the content of the nectar is less certain. Baker and Baker 1983 found that plants are fairly constant within a species in terms of sucrose dominance or hexo se dominance of their nectars, regardless of variation

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in the environment. Plants with the same pollinator type exhibit similarities in sugar ratios even without being taxonomically related. This strongly suggests that pollinators, like butterflies, have i mportant sugar preferences that can shape the evolution of nectars. Most nectar contains sucrose, glucose and fructose; only a few have only one detectable sugar, and none of these contain fructose alone, or sucrose and fructose without glucose. In general , Percival 1961 noted that families with deep tubed flowers tend to be sucrose rich while shallow tubed or cupped flowers are most often hexose rich. Baker and Baker 1983 distinguish butterfly visited flowers into two groups: those with deep narrow cor ollas characteristically rich in sucrose and visited primarily by butterflies, and those smaller short tubed hexose rich flowers visited equally by bees and butterflies. Sugar preference studies have been done in birds Martinez del Rio et al., 1992, sh owing that nectar feeding hummingbirds have specialized on sucrose, but that most fruit eating passerines cannot digest sucrose and instead show a preference for fruits containing the monosaccharide sugars fructose and glucose. Plants pollinated and disper sed by different birds thus use different types of sugars in their nectars and fruit pulps as rewards. If birds exhibit such diet related preferences for sugar types, one would expect fruit eating butterflies to possibly specialize on fructose and glucose, while nectar feeding butterflies of at least long corolla flowers may prefer sucrose. Baker and Baker 1983 did a survey of plant families and found that pollinator syndrome is the most important component of which sugar type a flower offers. Hummingbi rd pollinated flowers are sucrose rich and hummingbirds prefer sucrose. The case for butterflies and their flowers is still unknown. Butterflies may exhibit sugar preferences that reflect their diet, on fruit or either type of nectar. These patterns of pr eference may fall along taxonomic lines if mechanisms like the enzyme deficiencies reported in birds Martinez del Rio et al., 1992 exist in butterflies. Another possibility, that most butterflies have the enzymatic ability to use sucrose and hexose sugar s, would explain why butterfly flowers supply either sugar type. Some studies have been done to determine sugar preference in individual butterfly species, but one does not get an idea of big picture trends from them. Watt et al. 1974 found no preferenc e in Colias sp., as it was found that this genus has enzymes to process both hexose and sucrose sugars. Pieris brassicae , a species of Pierid, was found to prefer sucrose to fructose Romeis and Wackers, 2000, as Battus philenor in Erhardt s 1991 feeding experiments. Even though fructose ands glucose were the major sugars found in the fruits eaten by Nymphalis xanthomelas , Kanisca canace , and Vanessa indica , sucrose was the most effective feeding stimulant in these species, with fructose following close be hind Omura and Honda 2003. Ithomiines specialize on members of Asteraceae and Boraginaceae Baker and Baker, 1983, which have hexose rich nectars, so it is hypothesized that they prefer hexose sugars to sucrose. Besides these findings, few studies have been devoted to discovering sugar preferences in butterfly species let alone in multiple butterfly families. This experiment aims to see if a relationship between sugar preference and butterfly family phylogenies exists. Butterflies in this study were fed a solution containing only the sugars of interest, without presenting flower options. Though other monosaccharide and disaccharide sugars may be present in nectars, they occur in minute amounts, which is why this study focuses on sucrose, glucose and fruc tose in terms of sugar preference in butterflies.

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MATERIALS AND METHODS Study Site The experiment was conducted at the Selvatura butterfly garden and the Monteverde Butterfly Garden, both in Monteverde. Butterflies were collected from inside the garde ns, while feedings were conducted in a separate room at the Monteverde Butterfly Garden, and in an eclosion chamber at Selvatura. Preparation of Sugar Solutions The sucrose solution consisted of 20% sucrose by weight. The hexose mix consisted of 10% fr uctose by weight and 10% glucose by weight. Both solutions were stored in capped containers and remade every other day. Catching and Storing Butterflies were caught using a simple butterfly net, and then transferred to a 1 m tall cylindrical holding net , where they were kept for at least two hours before feeding, or overnight in the case of most of the morphos and heliconiines. This delay ensured that the butterflies would be less active and more inclined to eat the sugar solution presented to them witho ut trying to escape. Butterflies were caught in the late morning between 8 and 11 a.m. when they are most active, and were identified using The Butterflies of Costa Rica and their Natural History DeVries, 1987. Feeding Butterflies were transferred ind ividually to the feeding room, where they were placed on wax paper in front of a bead of 100 ðm L of sugar solution. While a nearby light bulb shone on the butterfly in order to heat it and induce it to feed, the butterfly was held lightly by the wings and it s proboscis was unrolled with an unbent paperclip to place the tip of the proboscis in the liquid. At this point, if the butterfly immediately started to feed, I would release its wings and let it feed until it was ready to fly away; otherwise, I would con tinue keeping its proboscis extended in the sugar solution until it kept it extended on its own for at least three seconds. Every butterfly that did not start feeding immediately was given three chances to eat. At the end of a feeding session, when the but terfly was satiated or had used up its three chances, the remaining sugar solution was sucked up by 20 ðm L microcapillary tubes and the volume measured. The wax paper would then be cleaned, a new bead of 100 ðm L would be applied by blowing sugar solution out o f a microcapilllary tube, and a new butterfly would be pulled from the holding net. This same feeding process was used for sucrose feedings and hexose feedings, though they were conducted on separate pieces of wax paper and with separate microcapillary tub es in order to avoid contamination.

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RESULTS Thirty three species from the families Pieridae, Papilionidae and Nymphalidae were tested. Of these, 26 species of three families and five nymphalid subfamilies were surveyed for both sugar solutions. Mean hexo se volumes were subtracted from mean sucrose volumes for all species that had representatives from both feedings, giving positive or negative values. Differences of <5% were listed as zero and discounted. The result of the Sign Test pointed to 14 cases whe re a species preferred sucrose and nine cases where a species preferred the fructose glucose mixture. In three cases there was little or no difference. Overall, butterflies do not show a preference for sucrose or hexose rich sugars Sign test, n=14 plus, 9 negative, p> 0.05. For all three Papilionid species tested, mean sucrose consumption was greater than mean hexose consumption, though in one trial, the difference was just 7 ðm L. Three of four Pierid species seemed to prefer sucrose to hexose. In these cases, there were too few species to test trends statistically. With eight negative and eight positive points for the Sign Test, no trend could be found in sugar preference, either in Nymphalidae as a family, or within the subfamilies Ithomiinae, Morphina e, Heliconiinae and Nymphalinae. Further Observations On the whole, the Pierid butterflies were very finicky eaters, and it was difficult to get them to take up even a little of either sugar solution. The Papilionids and certain members of Heliconiines were hard to feed as well, but to a lesser extent. Ithomiines and Morphos would readily eat, but whereas the smaller glass wing Ithomiines took half an hour to eat about 30 ðm L, the Morphos would eat the entire hundred ðm L to the point where I had to start fe eding them more until they were satiated. Parides individuals and most Pierid individuals were extraordinarily active when caught, but could not be stored overnight since they would die, so they were only stored between two and four hours. TABLE 1. Sug ar consumption in three butterfly families. Members of the family Nymphalidae are shown in red, Papilionidae in green, and Pieridae in blue. Butterfly individuals were collected from the Monteverde Butterfly Garden and the butterfly garden at Selvatura. Sp ecies Sucrose Sucrose Hexose Hexose Number fed Mean volume eaten ðm L Number fed Mean volume eaten ðm L Greta oto 5 30.682 5 26.788 Ithomia heraldica 1 36.05 1 36.8 Pteronymia fumida 1 40.26 0 Morpho granadensis 4 58.2875 3 62.544 Morpho peleides 5 83.364 5 78.474 Caligo eurilochus sulanus 1 33.4 1 400

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Caligo memnon memnon 1 346.3 2 344.2 Danaus plexippus 1 89.7 1 76.6 Dione moneta poeyii 3 55.089 2 44.737 Dryadula phaetusa 6 30.2183 1 49.5 Dryas iulia 2 40.791 3 58.3 Eueides isabella 4 31.9 1 5 34.788 Heliconius charitonius 3 29.123 3 31.929 Heliconius hecale zuleika 2 70.13 3 65.878 Heliconius sapho leuces 4 23.51 4 26.117 Heliconius sara fulgidus 2 23.29 2 24.08 Heliconius erato 2 35.79 2 26.449 Catonephele numilia esite 3 56.228 3 31 .053 Myscelia cyaniris cyaniris 3 30.263 3 36.404 Siproeta steleres 2 103.553 2 59.6 Battus polydamas polydamas 4 49.212 4 24.5 Parides lycimenes lycimenes 2 77.434 1 59.145 Parides arcas mylotes 0 1 12.1 Parides iphidamas iphidamas 0 1 68.4 Par ides erithalion 0 1 11.3 Papilio astyalus pallas 1 89.47 1 81.3 Papilio polyxenes 1 36.3 0 Papilio thoas nealces 0 1 177.6 Ascia limona 1 43.16 1 59.2 Phoebis sennae 2 46.316 3 23.86 Phoebis philea philea 1 59.5 1 55.5 Anteos clorine 0 1 92. 6 Appias drusilla 1 42.1 2 18.421

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FIGURE 1. Preference of sugar types by Papilionidae. Bars show results of a Sign Test where the mean volume of hexose solution eaten was subtracted from the mean volume of sucrose eaten. Bars pointing toward sucrose preference represent positive results. Negative results indicate hexose preference. FIGURE 2. Preference of sugar types by Pieridae. Positive results of a sucrose minus hexose Sign Test are shown as bars pointing to sucrose preference, while negati ve results point to hexose preference.

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FIGURE 3. Preference of sugar types by Nymphalidae. Subfamily Ithomiinae shown in red, Morphinae in blue, Danainae in black, Heliconiinae in green and Nymphalinae in purple. Positive results of a Sign Test indic ate sucrose preference; negative results indicate hexose preference. Neutral preference indicates <5% difference in mean consumed volumes. DISCUSSION Even though the Sign Test showed no overall sugar preference across butterfly species, this is to be e xpected. The butterflies examined include three famil i es and several subfamilies within Nymphalidae, some of which feed on fruits and some of whom have longer and shorter proboscides. What is perhaps most surprising is that, despite possible preferences, al l species readily took both sugar types, suggesting that both would serve as an acceptable reward for any of the species in the survey. Within the broader pattern, there were some trends worth investigating further. For example, all three Papilionidae sp ecies fed exhibited a preference for sucrose. Indeed, this finding supports Erhardt s 1991 study of Battus philenor , where individuals preferred both sucrose and fructose to glucose, but preferred sucrose to fructose. Confirmation of sucrose preference w ould also tie in to DeVries observations 1987 that

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the genus Parides feeds on flowers of the families Balsamaceae and Rubiaceae, which have fairly long tubes and therefore are probably sucrose rich. Three of the four genera of Pierid butterflies fed in this study also preferred sucrose to hexose, so it would be interesting to see if further studies corroborate a preference for sucrose over hexose, since there currently are conflicting lines of evidence for Pierid sugar preferences. Romeis and Wackers 2 000 study of Pieris brassicae revealed a preference for sucrose over fructose, but interestingly enough, Romeis and Wackers 2002 also found that Pieris brassicae fecundity is compromised by sucrose, and glucose is the only sugar with a positive effect on both longevity and fecundity. Furthermore, according to DeVries 1987, Pierids feed on red flowers in general, and as many red flowers are hummingbird pollinated, one would expect them to be sucrose rich. Pierids also feed on shallow tubed Lantana camara , however, which according to Percival 1961 points toward a hexose rich nectar. If hexoses are good for Pierids, we might expect to see a hexose preference, but there is a possibility that the Pieris brassicae preference for sucrose can be expanded to th e whole family. Again, further study is necessary to determine if there is a family trend for sucrose preference. The Nymphalidae did not show a strong preference for either sucrose or hexose, but even if future studies corroborate this lack of feeding pr eference pattern, it would not be entirely unexpected, since certain subfamilies of Nymphalids have become specialized on different diets. For example, adults of the subfamily Morphinae do not visit flowers, and instead feed exclusively on plant sap and th e juices of rotted fruits and fungi DeVries, 1987; meanwhile, Heliconiines and Ithomiines eat nectar, often from Lantana camara . There might then be patterns within smaller clades, but the family as a whole would not have a single evolved preference. It homiines have specialized on members of the families Boraginaceae and Asteraceae, whose nectars are predominantly hexose rich Baker and Baker, 1983. This corresponds with Percival s 1961 observations that shallow flowers, usually found in inflorescence s, tend to be hexose dominant. However, those observations do not correspond with this study s findings, wherein Greta nero ate more sucrose and Ithomia heraldica had no discernable preference. Clearly more feeding experiments need to be carried out. Morp hinae species are specialized fruit eaters, so one would expect them to prefer fructose and glucose, as these sugars are the most common sugars in fruit Omura and Honda, 2003; however, this is not what was found. Two of the four Morphinae species studied M. peliedes and C. eurilochus preferred the hexose solution, while M. granadensis on average ate more sucrose solution and C. memnon showed no discrimination between sugar solutions at all. Looking again at Omura and Honda s 2003 study, where they found that sucrose was the preferred sugar but that fructose was also a very good feeding stimulant and occurred in higher quantities, perhaps concentration plays a role in sugar feeding preferences. Part of the problem could also be that Morphos have a tendenc y to eat the entirety of the volume of solution they are presented with, which made it hard to find a pattern in terms of preference. One way to possibly overcome this would be to feed them until they are fully satiated, which would require one to put down far more than 100 ðm L at a time. Future experiments on Morphinae sugar preference might consider feeding them solutions of fructose, a hexose mixture, and sucrose, each with the necessary ethanol and acetic acid compounds to entice feeding.

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Two of the thr ee Nymphalinae species preferred sucrose, and five Heliconiinae species preferred sucrose as opposed to three heliconiines that preferred hexose. The genus Heliconius eat pollen from Gurania costaricense Gilbert, 1983 , but nothing is mentioned about the sugars they get from nectar, and any conjecture is made more complicated by the fact that hummingbirds may also visit G. costaricense . I was unable to find any literature on the feeding habits of adult Nymphalinae species that would point to either sucrose or hexose preference Again, more studies should be carried out to find out if Heliconiines and other Nymphalid butterflies exhibit preferences for specific sugar types. Overall, data presented here cannot conclusively say that butterflies as a whole a ppear to favor one sugar type over the other even though there were more positive i.e., sucrose preference results from the Sign Test than negative results. A Omura and Honda 2003 point out, much of the existing literature concerning butterfly feeding habits points to a trend of sucrose preference over fructose, and fructose preference over the remaining sugars, but there are not enough studies of multiple butterfly families for one to really get a sense of a pattern. Successful attraction of pollina tors is complex, relying on many factors other than nectar sugar type. Pollinators will ignore one flower if other flowers with more desirable nectar are present Vansell et al., 1942, and will feed on less desirable flowers of attractive competitors are not present von Frisch, 1950. It could very well be that butterflies will eat sugars from whatever flowers are available to them, and their sugar preference will change as the composition of flower species in their environment changes. If butterflies a lso have the ability to digest multiple types of sugars as Watt et al. found 1974, sugar preferences would be further confounded. In this case, future studies would end up with similarly inconclusive results in terms of sucrose vs. hexose preference, and no pattern would be found relating butterfly phylogeny to adult dietary preference. In any event, butters flies do not seem constrained to one or few sugar types as was found for some frugivorous and nectarivorous birds Martinez del Rio et al., 1992. T he ecological implications of these findings for nectar flowers are rather curious. Despite the strong patterns shown by Baker and Baker 1983 with regards to nectar composition, there seems to be no cause for specified sugar compositions in butterfly pol linated flowers. Sugar preference works really well for hummingbirds, as they really strongly prefer sucrose and the hummingbird flowers are all sucrose, but if butterflies don t really have a sugar preference, why would butterfly pollinated flowers be con strained to sugar types within families and pollination syndromes? Why should long corolla flowers have sucrose rich nectar and short corolla flowers have hexose rich nectar? One explanation could be that the long corolla flowers being pollinated by butter flies are also pollinated by hummingbirds, necessitating the production of a sucrose rich nectar. Another similar explanation has to do with bee sugar preference. Apparently long tongued bees tend to be rewarded with sucrose rich nectar, while short tongue d bees rarely are rewarded with sucrose Baker and Baker, 1983. It could well be that certain flowers contain either sucrose or hexose in order to reward bee pollinators, and butterflies have jumped on the existing pollination syndrome wagon. However, thi s

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hypothesis is not without problems: many of the long tongued bee flowers come from a few set families characterized by sucrose rich nectar while the short tongued bee flowers are from hexose rich families like Asteraceae, so it is hard to say if the flow ers are producing specific nectars to conform to bee preference, or the other way around. Finally, one might want to explore the costs of producing sucrose as opposed to hexose nectars, as this might give insight into which pollinators are dictating the co mposition of sugar nectars. One major limitation of the study was that all of the butterflies had to be caught individually, which was time consuming and gave an uneven distribution of representatives from certain families the Heliconiines are much more numerous than the Papilionids. The caught individuals were also much more energetic and resistant to being force fed, so I had to institute a waiting period before feeding in order to let them calm down and get hungry. The Parides and Pierid individuals in particular were difficult to feed since they could only be contained for two to four hours without food, which was not enough time for them to become hungry and less agitated. A way to improve this study would be to acquire at least twenty individual s of each species as pupae, feed them one of the solutions after they hatched, contain them for a day or two, and then feed them the other solution. This would greatly increase the evenness of representation between species. Such experiments would be incre dibly valuable, since to my knowledge, no other studies have been done looking at adult feeding preferences across butterfly families, relating dietary preference to phylogeny or ecology. Further studies could be undertaken to look at the biology of butter fly digestion that may also shed light on evolved dietary preferences. ACKNOWLEDGEMENTS I would like to thank the staff at the Monteverde Butterfly Garden and the garden at Selvatura for allowing me to do my experiment in their gardens, use their butterf lies, and for hel ping me identify individuals. A very special thanks goes to Alan Masters for guiding me towards the idea of this project after all the others failed. I can t thank you enough. Thanks also to Taegan and Pablo for running to the supply close t for me time after time, and Pablo especially for help translating the abstract. Literature Cited B AKER , H. G. AND I. B AKER . 1983. Floral nectar sugar constituents in relation to pollinator type. In C. Eugene Jones and R. John Little Eds.. Handbook of Experimental Pollination Biology . pp. 118 139. Van Nostrand Reinhold Publishing, Ontario, Canada. D E V RIES , P. J. 1987. The Butterflies of Costa Rica and their Natural History . Princeton University Press, Princeton, New Jersey. E RHARDT , A. 1991. Nectar sugar and amino acid preferences of Battus philenor Lepidoptera, Papilionidae. Ecological Entomology. 164: 425 434. VON F RISCH , K. 1950. Bees Their Vision, Chemical Senses and Language . Cornell University Press, Ithaca, New York. In Baker, H.G. and I. Baker. 1983 . Floral nectar sugar constituents in relation to pollinator type. In C. Eugene Jones and R.

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John Little Eds.. Handbook of Experimental Pollination Biology . pp. 118 139. Van Nostrand Reinhold Publishing, Ontario, Canada. J ANZ , N. AND S. N YLIN . 1998. Butterflies and plants: a phylogenetic study. Evolution 522: 486 502. M ARTINEZ DEL R IO , C., H. G. B AKER AND I. B AKER . 1992. Ecological and evolutionary implications of digestive processes: Bird preferences and the sugar constituents of floral nectar and fruit pulp. Cellular and Molecular Life Sciences. 486: 544 551. P ERCIVAL , M. S. 1961. Types of nectar in angiosperms. New Phytology. 60: 235 281. In Baker, H.G. and I. Baker. 1983 . Floral nectar sugar constituents in relation to polli nator type. In C. Eugene Jones and R. John Little Eds.. Handbook of Experimental Pollination Biology . pp. 118 139. Van Nostrand Reinhold Publishing, Ontario, Canada. O MURA , H. AND K. H ONDA . 2003. Feeding responses of adult butter ûies, Nymphalis xantho melas , Kaniska canace and Vanessa indica , to components in tree sap and rotting fruits: synergistic effects of ethanol and acetic acid on sugar responsiveness. Journal of Insect Physiology. 49: 1031 1038. R OMEIS , J. AND F. L. W ACKERS . 2000. Feeding respo nses by female Pieris brassicae butterflies to carbohydrates and amino acids. Physiological Entomology. 253: 247 253. ----, AND ---. 2002. Nurtitional suitability of individual carbohydrates and amino acids for adult Pieris brassicae . Physiological Entomology. 272: 148 156. V ANCELL , G. H., W. G. W ATKINS and R. K. B ISHOP . 1942. Orange nectar and pollen in relation to bee activity. Journal of Econ. Entomology. 35: 321 323. In Baker, H.G. and I. Baker. 1983 . Floral nectar sugar constituents in relat ion to pollinator type. In C. Eugene Jones and R. John Little Eds.. Handbook of Experimental Pollination Biology . pp. 118 139. Van Nostrand Reinhold Publishing, Ontario, Canada. A MBROSE III, H. W., P. A MBROSE , D. J. E MLEN AND K. L. B RIGHT Eds.. 200 2. A Handbook of Biological Investigation 6 th Ed., pp. 78 80. Hunter Textbooks, Inc. Winston Salem, North Carolina. G ILBERT , L. E. Anguria and Gurania Rain forest cucumber. 1983. In Janzen, D. Ed.. Costa Rican Natural History, pp. 190 191. W ATT , W. B., P. C. H OTCH , AND S. G. M ILLS . 1974. Nectar resource use by Colias butterflies. Oecologia. 14: 353 374. In Baker, H.G. and I. Baker. 1983 . Floral nectar sugar constituents in relation to pollinator type. In C. Eugene Jones and R. John Little Eds .. Handbook of Experimental Pollination Biology . pp. 118 139. Van Nostrand Reinhold Publishing, Ontario, Canada.

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APPENDIX APPENDIX 1. Phylogeny of Lepidoptera from Janz and Nylin 1998.

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APPENDIX 2. Sign Test Results Subfamily Species Sucrose mean volume ðm L Hexose mean volume ðm L Sign of S H Ithomiinae Greta oto 30.68 26.79 + Ithomia heraldica 36.05 36.8 0 Morphinae Morpho granadensis 58.29 62.54 Morpho peleides 83.36 78.47 + Caligo eurilochus sulanus 33.4 400 Caligo me mnon memnon 346.3 344.2 0 Danainae Danaus plexippus 89.7 76.6 + Heliconiinae Dione moneta poeyii 55.09 44.74 + Dryadula phasetusa 30.22 49.5 Dryas iulia 40.79 58.3 Eueides isabella 31.91 34.79 Heliconius charitonius 29.12 31.93 Heliconi us hecale zuleika 70.13 65.88 + Heliconius sapho leuces 23.51 26.12 Heliconius sara fulgidus 23.29 24.08 0 Heliconius erato 35.79 26.45 + Nymphalinae Catonephele numilia esite 56.23 31.05 + Myscelia cyaniris cyaniris 30.26 36.4 Siproeta stel eres 103.55 59.6 + Papilioninae Battus polydamas polydamas 49.21 24.5 + Parides lycimenes lycimenes 77.43 59.15 + Papilio astylaus pallas 89.47 81.3 + Pierinae Ascia limona 43.19 59.2 Appias drusilla 42.1 18.42 + Coliadinae Phoebis sennae 46.32 23.86 + Phoebis philea philea 59.5 55.5 + Total positive: 14 Total negative: 9 Family Pieridae Positive: 3 Negative: 1 Family Papilionidae Positive: 3 Negative: 0 Family Nymphalidae Positive: 8 Negative: 8

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APPENDIX 3. Sucrose Feeding Data SPECIES Vo l. Sucrose Eaten €L Pteronymia fumida 40.26 Dryadula phaetusa 68.16 Morpho peliedes 55.26 Morpho peliedes 99.73 Heliconius charitonius 20 Greta oto 25.79 Ascia limona 43.16 Heliconius hecale zuleika 72.89 Heliconius sapho leuces 23.68 Eueides isabella 58.16 Heliconius charitonius 40.79 Greta oto 21.84 Heliconius sara fulgidus 26.84 Heliconius erato 50.53 Greta oto 34.47 Heliconius hecale zuleika 67.37 Heliconius sapho leuces 44.47 Heliconius charitonius 26.58 Helic onius sara fulgidus 19.74 Heliconius erato 21.05 Eueides isabella 24.74 Eueides isabella 13.42 Eueides isabella 31.32 Heliconius sapho leuces 10.26 Heliconius sapho leuces 15.63 Ithomia heraldica 36.05 Morpho peliedes 62.1 Greta oto 44.2 1 Morpho peliedes 100 Morpho peliedes 99.73 Morpho granadensis 99.73 Dryadula phaetusa 32.1 Morpho granadensis 74.47 Papilio astyalus pallas 89.47 Phoebis argante 0 Dione moneta poeyii 55.53 Parides lycimenes 68.95 Battus polydamas 99.7 4 Battus polydamas 66.05 Dione moneta poeyii 50.79 Morpho granadensis 43.95 Dryadula phaetusa 8.95 Dryadula phaetusa 21.05 Morpho granadensis 15 Dryadula phaetusa 42.63 Dryadula phaetusa 8.42 Appias drusilla 42.1

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Greta oto 27.1 Dryas iulia 48.95 Parides lycimenes 29.47368421 Battus polydamas 74.73684211 Danaus plexippus 89.73684211 Parides lycimenes 27.89473684 Catonephele numilia esite 90.26315789 Caligo eurilochus sulanus 33.42105263 Dione moneta poeyii 58.94736842 Papilio polyxenes 36.31578947 Myscelia cyaniris cyaniris 41.31578947 Myscelia cyaniris cyaniris 32.10526316 Phoebis sennae 80.26315789 Catonephele numilia esite 41.05263158 Caligo memnon memnon 346.3157895 Dryas iulia 32.63157895 Myscelia cy aniris cyaniris 17.36842105 Phoebis sennae 12.36842105 Siproeta steleres 131.5789474 Catonephele numilia esite 37.36842105 Siproeta steleres 75.52631579 Battus polydamas 69.21052632 Phoebis philea philea 59.47368421 APPENDIX 4. Glucose Fruct ose Feeding Data SPECIES Vol. Hexose Eaten €L Heliconius charitonius 26.84 Eueides isabella 22.63 Heliconius charitonius 37.63 Eueides isabella 32.89 Heliconius erato 35.53 Heliconius hecale zuleika 60 Greta oto 17.1 Heliconius sapho leuces 16.84 Dryadula phaet usa 36.32 Greta oto 41.05 Greta oto 43.16 Heliconius hecale zuleika 44.74 Heliconius sapho leuces 33.68 Greta oto 20.53 Greta oto 12.1 Heliconius sara fulgidus 28.95 Heliconius sapho leuces 15.78947368 Papilio astyalus pallas 81.31578 947 Morpho granadensis 73.94736842 Phoebis sennae 26.84210526 Parides lycimenes 40.52631579 Heliconius charitonius 31.31578947 Morpho granadensis 64.73684211

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Dryas iulia 61.31578947 Dryas iulia 53.94736842 Heliconius sara fulgidus 19.2105 2632 Heliconius sapho leuces 38.15789474 Eueides isabella 44.73684211 Eueides isabella 43.68421053 Heliconius hecale zuleika 92.89473684 Heliconius erato 17.36842105 Morpho peleides 2.631578947 Morpho peleides 100 Morpho granadensis 48.9473 6842 Dryadula phaetusa 49.47368421 Eueides isabella 30 Morpho peleides 100 Morpho peleides 100 Morpho peleides 89.73684211 Anteos clorine 92.63157895 Battus polydamas 46.31578947 Battus polydamas 98.94736842 Appias drusilla 15.78947368 Appias drusilla 21.05263158 Dione moneta poeyii 33.15789474 Dione moneta poeyii 56.31578947 Ascia limona 59.21052632 Catonephele numilia esite 30.52631579 Parides lycimenes 24.47368421 Caligo eurilochus 400 Papilio thoas nealces 177.6315789 Battus polydamas 87.36842105 Myscelia cyaniris cyaniris 29.47368421 Myscelia cyaniris cyaniris 45.52631579 Catonephele numilia esite 20 Dryas iulia 59.73684211 Caligo memnon memnon 391.5789474 Phoebis sennae 25.26315789 Parides arcas mylot es 12.10526316 Myscelia cyaniris cyaniris 34.21052632 Phoebis sennae 19.47368421 Parides iphidamas iphidamas 68.42105263 Siproeta steleres 87.63157895 Danaus plexippus 76.57894737 Parides erithalion 11.31578947 Catonephele numilia esite 42.63 157895 Caligo memnon memnon 296.8421053 Battus polydamas 3.947368421 Siproeta steleres 31.57894737 Phoebis philea philea 55.52631579 Ithomia heraldica 36.84210526


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