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Mist frequency and butterfly emergence from the chrysalis: Implications for tropical cloud forest climate change

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Title:
Mist frequency and butterfly emergence from the chrysalis: Implications for tropical cloud forest climate change
Translated Title:
La frecuencia de llovizna y aparición de la mariposa de la crisálida: Implicaciones para el cambio del clima tropical del bosque nuboso ( )
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English
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Davies, Lindsay
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Subjects / Keywords:
Butterflies   ( lcsh )
Fog   ( lcsh )
Climatic changes   ( lcsh )
Costa Rica--Puntarenas--Monteverde Zone--San Luis   ( lcsh )
Mariposas
Neblina
Cambios climáticos
Costa Rica--Puntarenas--Zona de Monteverde--San Luis
Tropical Ecology Spring 2011
Mist
Ecología Tropical Primavera 2011
Llovizna
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Reports   ( lcsh )
Reports

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Abstract:
The cloud bank in Monteverde, Costa Rica is rising due to global warming. This is causing fewer misty days and is impacting the habitats of local organisms. Butterflies have been shown to be negatively impacted by moisture in terms of livelihood. Heliconius charithonia, a butterfly species occurring from 0-1200 meters, and Heliconius hecale zuleika, occurring from 0-1700 meters, were exposed to intermittent (five minutes mist, five minutes dry over the course of the day) and intense mist conditions (10 hours of constant mist a day) to measure the effect of mist on survivorship, days to eclosion, and development (in terms of mass, wing length, and wing quality). As the climate of Monteverde becomes dryer, these butterflies may move up in elevation to take advantage of the dry conditions. Intermittent mist had no effect on survival or days to eclosion. Twenty butterflies eclosed from the intermittent mist tank and 27 from the dry. In dry conditions, 88% (n=17) of H. charithonia survived while 71% (n=17) of H. hecale zuleika. Ten of each species survived in the misted environment, therefore 59% (n=17) survival for both. The intermittent mist did significantly decrease wing quality, wing length, and mass for H. charithonia, but not for H. hecale zuleika. For wing length of H. charithonia, the average length was 4.19 +/- 0.16 cm in the dry and 3.53 +/- 0.2 cm in the wet. The masses of H. charithonia were nearly significantly different between the two climates; in the dry tank their mass was 0.19 +/- 0.02 g and in the wet was 0.15 +/- 0.02 g. H. charithonia may not be as well adapted to mist as H. hecale zuleika, since H. charithonia showed decreased development. Constant mist killed most of butterflies and caused the eclosion rates of those that did eclose to increase. A total of seven butterflies eclosed from the constant mist and 29 from the dry. In the dry tank, H. charithonia had 79% (n=17) of the butterflies survive; H. hecale zuleika had 93% (n=17). In the wet tank, 30% (n=17) of the H. charithonia survived and 7% (n=17) of the H. hecale zuleika. In the dry tank, the average days to eclosion was 8.6 +/- 0.5, almost 3 days less than those in the wet tank, 11.8 +/- 0.49; Constant mist was not shown to affect development, but probably because of the low number of butterflies that eclosed. Overall, as the mist recedes and the number of dry days increases, the conditions improve for these butterflies to move up in elevation. H. charithonia may move up to Monteverde for breeding and living, also, H. hecale zuleika may increase their breeding and time in Monteverde.
Abstract:
El banco de nubes en Monteverde, Costa Rica está subiendo debido al calentamiento global. Esto está causando menos días nubosos y está impactando el hábitat de organismos locales. Las mariposas han mostrado un impacto negativo por la humedad en términos de sobrevivencia. Heliconius charithonia es una mariposa que se encuentra entre los 0-1200 metros y Heliconius hecalezuleika se encuentra entre los 0-1700 metros, fueron expuestos a niebla intermitente (5 minutos con niebla, 5 minutos en lo seco) y en condiciones intensas (10 horas constantes de niebla durante el día) para medir el efecto de la sobrevivencia, días para eclosionar y desarrollar (en términos de masa, largo del ala y calidad del ala).A medida que el clima de Monteverde se convierta más seco, estas mariposas pueden ascender a mayores elevaciones para aprovechar las condiciones secas. La neblina intermitente no tiene efecto alguno en la sobrevivencia o días en eclosionar. 20 mariposas eclosionaron de la niebla intermitente y 27 del ambiente seco. En condiciones secas, 88% (n=17) de H. charithonia sobrevivieron mientras que en un 71% (n=17) los de H. hecale zuleika. Diez de cada especie sobrevivió en el ambiente con niebla constante. La niebla intermitente disminuyo significativamente la calidad de las alas, el tamaño del ala y la masa para H. charitona pero no para H. hecalezuleik. La masa de H. charitona fue diferente entre los dos ambientes, en el tanque seco la masa fue de 0.19 +/- 0.02 g y en el húmedo fue de 0.15 +/- 0.02 g. H. charithonia puede estar no tan bien adaptada a la niebla como la otra especie ya que demuestra una disminución en el desarrollo. Un total de siete mariposas eclosionaron de la niebla constante y 29 del seco. En el tanque seco H. charithonia tiene un 79% de sobrevivencia (N=17) y H. hecalezuleika un 93% (N=17). En el tanque húmedo un 30% (N= 17) de H. charithonia sobrevivió y en un 7% (N=17) de H. hecalezuleika. En el tanque seco el promedio de días de eclosión fue de 8.6 +/- 0.5, casi 3 días más que en el tanque húmedo 11.8 +/- 0.49. La niebla constante no muestra un efecto en el desarrollo pero probablemente debido al número bajo de mariposas que eclosionaron. Sobre todo, al aumentar el número de días secos, las condiciones se prestan para que estas mariposas se muevan a elevaciones mayores, H. charithonia puede moverse a Monteverde para reproducirse y vivir, además H. hecalezuleika puede aumentar su reproducción y tiempo en Monteverde.
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Mist Frequency and Butterfly Emergence from the Chrysalis : Implications for Tropical Cloud Forest Climate Change Lindsay Davies Department of Biology, Indiana University Bloomington ABSTRACT The cloud bank in Monteverde, Costa Rica is rising due to global warming. This is causing fewer misty days and is impacting the habitats of local organisms. Butterflies have been shown to be negatively impacted by moisture in terms of livelihood. Heliconius charithonia a butterfly species occurring from 0 1200 meters and Heliconius hecale zuleika occurring from 0 1700 meters, were exposed to intermit tent (five minutes mist, five minutes dry over the course of the day) and intense mist conditions (10 hours of constant mist a day) to measure the effect of mist on survivorship, days to eclosion, and development (in terms of mass, wing length, and wing quality) As the climate of Monteverde becomes dryer, these butterflies may move up in elevation to take advantage of the dry conditions Intermittent mist had no effect on survival or days to eclosion Twenty butterflies eclosed from the intermittent mist tank and 27 from the dry. In dry conditions, 88% (n= 17 ) of H. charithonia survived while 71% (n=17 ) of H. hecale zuleika Ten of each species survived in the misted environment, therefore 59% (n=17) survival for both The intermittent mist did significantly decrease wing quality, wing length, and mass for H. charithoni a, but not for H. hecale zuleika For wing length of H. charithonia the average length was 4.19 +/ 0.16 cm in the dry and 3.53 +/ 0.2 cm in the wet The masses of H. charithonia were nearly significantly different between the two climates ; in the dry tank their mass was 0.19 +/ 0.02 g and in the wet was 0.15 +/ 0.02 g H. charithonia may not be as well adapted to mist as H. hecale zuleika, since H. charithonia showed decreased development. Constant mist killed most of butterflies and caused the eclosion rates of those that did eclose to increase. A total of seven butterflies eclosed from the constant mist and 29 from the dry. In the dry tank, H. charithonia had 79% (n=17) of the butterflies survive; H. hecale zuleika had 93% (n=17). In the wet tank 30% (n=17) of the H. charithonia survived and 7% (n=17) of the H. hecale zuleik a In the dry tank, the average days to eclosion was 8.6 +/ 0.5, almost 3 days less than those in the wet tank, 11.8 +/ 0.49 ; Constant mist was not shown to affect development, but probably because of the low num ber of butterflies that eclosed Overall, as the mist recedes and the number of dry days increases, the conditions improve for these butterflies to move up in elevation. H. charithonia may move up to Monteverde for breeding and living, also, H. hecale zuleika may increase their breeding and time in Monteverde RESUMEN El banco de nubes en Monteverde, Costa Rica esta subiendo debido al calentamiento global. Esto est causando menos das nubosos y est impactando el hbitat de organismos locales. Las ma riposas han mostrado un impacto negativo por la humedad en trminos de sobrevivencia. Heliconius charithonia es una mariposa que se encuentra entre los 0 1200 mts y Heliconius hecalezuleika se encuentra entre los 0 1700 mts, fueron expuestos a niebla inte rmitente (5 minutos niebla, 5 minutos seco) y condiciones intensas (10 horas constantes de niebla durante el da) para medir el efecto de la sobrevivencia, das para eclosionar y desarrollo (en trminos de masa, largo del ala y calidad del ala). En cuanto el clima de Monteverde se torne ms seco estas mariposas se pueden mover a elevaciones mayores. Neblina intermitente no tiene efecto alguno en sobrevivencia o das en eclosionar. 20 mariposas eclosionaron de la niebla intermitente y 27 del ambiente seco. En condiciones seca, 88% (n=17) de H. charithonia sobreviviero mientras 71% (n=17) de H. hecale zuleika. Diez de cada especie sobrevivi en el ambiente con niebla constante. La niebla intermitente disminuyo significativamente la calidad de las alas, el tamao del ala y la masa para H. charitona pero no para H. hecalezuleik. La masa de H. charitona fue difere nte entre los dos ambientes en el tanque seco la masa fue 0.19 +/ 0.02 g and y en el hmedo fue 0.15 +/ 0.02 g. H. charithonia puede estar no tan bien adaptada a

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la niebla como la otra especie ya que demuestra una disminuacin en el desarrollo. Un total de siete mariposas eclosionaron de la niebla constante y 29 del seco. En el tanque seco H. charithonia tiene un 79% de sobrevivencia (N=17) y H. hecalezuleika un 93% (N=17). En el tanque hmedo 30% (N= 17) de H. charithonia sobrevivi y 7% (N=17) de H. hecalezuleika. En el tanque seco el promedio de das de ecloson fue de 8.6 +/ 0.5, casi 3 das ms que en el tanque humedo 11.8 +/ 0.49. La niebla constante no muestra un efecto en el desarrollo pero probablemente debido al nmero bajo de mariposas que eclosionaron. Sobre todo, al aumentar el nmero de das secos, las condiciones se prestan para que estas mariposas se muevan a elevaciones mayores, H. charithonia puede moverse a Monteverde para reproeducirse y vivir, adems H. hecalezuleika puede aument ar su reproduccin y tiempo en Monteverde. INTRODUCTION It is widely accepted that our planet is warming (Walther et al. 2002) A verage surface temperature has increa sed around 0. 6 C during the 20 th Century and is anticipated to increase between 1.4 and 5.8 C by 2100 ( Houghton et al. 2001 ) This temperature increase is caused by an increase of greenhouse gases in the atmosphere ( s uch as carbon dioxide, methane and nitrous oxide) and is largely the outcome of human activities, such as tropical deforestation and burning of fossil fuels (Houghton et al. 2001 ) Global warming has caused different climatic changes. The poles have seen melting of glaciers and the snow and ocean levels are changing worldwide (Houghton et al. 2001). Some areas of the Northern Hemisphere will see increases in precipitation while others will see a decrease (Walther et al. 2002). o phenomenon is expected to be more frequent (Walther et al. 2002). These abiotic changes have lead to biotic changes as well : plants ar e flowering earlier, birds are changing their migration patterns, and invasive plants and diseases are spreading (Walther et al. 2002). Although all areas of the globe will be affected somehow by global warming, diversity hotspots, such as the Tropical mountains will be greatly affected due to their narrow abiotic variability and high s pecies richness and/or endemism The C loud F orest of Monteverde located between 1500 1800 meters often experiences humid, cloudy or foggy conditions. G lobal warming i s causing the cloud bank to rise in Monteverde (Pounds et al. 1999) The R aising C loud B ank theory predicts misty days will be replaced by more dry days (Pounds et al. 1999). This alteration in climate has caused some biological changes for many species in the area, most notably, the disappearance of the G olden T oad (Bufo periglenes). In addition, the disappearance of other amphibians and upward movement of bird specie s are likely the result of warming and consequent drying (Pounds et at 1999). The impact of lower mist frequency and more consecutive dry days on most taxa is poorly understood however The effects on Cloud Forest butterfly communities, for example, is unknown. The life cycle of butterflies depends a great deal on cli mate (Roy et al. 2001) A fter the butterfly emerges from the chrysalis time is needed for the wings to dry before flight can be attempted W arm, dry seasons can favor successful eclosion and may help explain why dryer conditions generally favor the survival and population sizes of many butterfly species (Pollard 1988). At the chrysalis stage, intermittent and constant pre cipitation have been shown to increase morta lity rates and eclosion time (Clifford 2008). Both quality of wing and wing length were negatively impacted by precipitation (Clifford 2008). Also, too much moisture can provide an environment for pathogens (Mur phy et al. 1992). However, a certain amount of mist may be needed for survival, in order to prevent desiccation of the chrysalis (Smetacek 2009).

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Changes in precipitation and moisture levels als o have an impact on butterfly ranges Two checkerspot butterflies were shown to go extinct as a result of variable precipitation caused by global warming (McLaughlin et al. 2002 ) In the temperate United States, a drought caused the displacement of butterflies into a new habitat (Debinski et a l. 2006). These changes in butterfly populations as a result of a decrease in moisture could be seen in the Tropics as well. With misty conditions in Monteverde decreasing, those butterflies accustomed to mist may not survive and /or may become displaced by more dry adapted butterfly populations. Higher elevations in Monteverde experience a higher frequency of misty conditions This study use d an aquarium fogger to mimic the misty conditions of the Monteverde Cloud Forest I aim to test how butterflies fro m lowland areas respond to intermittent mist, continuous mist, and dry conditions by looking at th eir eclosion from the chrysalis. M isty conditions are expected to limit lowland species suggesting that the lowland species may move into higher elevations of Monteverde as mist frequency declines. METHODS Study organisms Two butterfly species of the Heliconiinae (Nymphalidae) were used for this study : H. charithonia, a black butterfly with cream to yellow stripes and dots (Figure 1a) and H. hecale zuleika black with white spots and orange (Figure 1b). Both are found on both slopes of Costa Rica but differ in their elevational habitats H. charithonia the lowland species, and H. hecale zuleika the highland, can both be found on both sl opes of Costa Rica. Both feed on Passiflora spp found in the Monteverde Cloud Forest (DeVries, 1987, Haber, 2000), but while H. charithonia is reported for the Monteverde community, it does not reach the higher elevations of the Monteverde Cloud Forest. 1a. 1b. Figure 1. Morphological differences between the two study butterfly species; both are widespread species from Heliconiinae (Nymphalidae) and found on both slopes of Costa Rica 1a shows Heliconius charithonia, found from 0 1200 meters. 1b shows Heliconius hecale zuleika found from 0 1700 meters H. charithonia are found from sea level to 1200 meters in Costa Rica, they are restricted to environments that, while rainy, have parts of most days th at are sunny and dry. H. hecale zuleika a butterfly species found from sea level to 1700 meters (DeVries 1987) is found higher and well into Monteverde Cloud Forest (Haber, 1983). Also, it is important to note that ranges are based on adult sightings and may not mean that the butterflies differ in breeding range.

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Experiment 1 Intermittent Mist and Butterfly Development The study took place in the laboratory at the University of Georgia Biological Station in San Luis, Costa Rica. This is surrounded by pas ture and secondary forest that used to be Pacific slope Premontane Moist Forest. It experiences a pronounced dry season from January to mid May. This study was performed at the end of the dry season in April 2011. Sixty eight chrysalises 34 of each species, were purchased from a grower that supplies th e Monteverde Butterfly Garden The chrysalises were tacked to Styrofoam and placed in two terrariums each terrarium containing 17 of each species of equal and known age One terrarium expose d the chrysalis to a low humidity environment which was the ambie nt conditions of the laboratory ( 20 C and 60% humidity ) The other terrarium had intermittent, on and off mist from a fogger (Figure 2 ) The intermittent mist terrarium had a fogger (Zoo Med Reptile Fogger Terrarium Humidifier, www.zoomed.com ) attached to a Humidity Controller (Zoo Meds HygroTherm Humidity and Temperature Controller) This controller was set to its highest setting of 95% relative humidity. As a result, the fogger would fill the terrarium with fog, at which time the controller would turn the fogger off. When relative humidity dropped below 85%, the fogger would turn ba ck on and fill the terrarium with mist once more. This resulted in intermittent mist where there was about five minutes of mist followed by five minutes of gradually dissipating mist (Figure 2a). In the end, visib ility in the terrarium was about 30 minutes of a given hour. The fogger generally delivered a liter of water in a 24 hour period. 2 a 2b 2c

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Figure 2. Different climate environments for H. charithonia and H. hecale zuleika Seventeen chrysalises of each butterfly species were exposed to dry, intermittent mist, and constant mist for 16 days and the survival, eclosion, wing length, wing quality, and mass were noted. Mist was provided by a Zoo Med Reptile Fogger Terrar ium Humidifier and the controller 2a shows the intermittent mist tank mist filled the tank until it reached 95% humidity, then the controller would shut it off. The mist would start again after the humidity reached 85% 2b shows the intensely misted en vi ronment constant mist would fill the tank for 10 hours a day 2c shows the dry mist environment. Both terrariums were placed in front of an air conditioning unit, to help keep the tank cool and the temperature in both tanks constant The average temperature in both tanks was 20 C Every day the terrariums were monitored to se e if any of the butterflies had emerged or had died. When a butterfly emerged from its chrysalis it was allowed to dry for at least an hour, before its forewing length wa s measured The adult butterfly was also weighed and the status of the wing was noted. The wing status was ranked on a scale from 1 to 5. 1 corresponded to wings of excellent quality, with no wrinkles and/or folding of the wing s ; 5 corresponded to wings of very poor quality, with ex treme wrinkles and/or folding (Figure 3) 3a. 3b. Figure 3. Differing statuses of wings of H. hecale zuleika and H. charithonia After the butterflies emerged from the intermittent mist, constant mist, or dry environment, they were allowed to dry for at least one hour; then their wing status was noted. Wing status was assigned based on a scale from 1 5 based on quality. A score of 1, shown in 3a, reflects wing of very h igh quality (no folds, wrinkles). A score of 5, shown in 3b, reflects wings of very low quality (significant folds and wrinkles that would affect the flight). Experiment 2 Intense Mist and Butterfly Development The second experiment involved the same set up as the first, except the fog ged terrarium was disconnected from the humidity controller. It misted with intense fog f or an average of 10 hours, from morning until early evening. Without the controller, the fogger would mist for five hours, so I woul d fill it during the day. The fogger would not mist through the night, so it was hooked up to the controller and the chrysalises were exposed to intermittent mist throughout the night.

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Observations H. charithonia is overall a smaller butterfly species in size than H. hecale zuleika T herefore the two were not compared in terms of mass, wing status, and wing length When recording data, t hose butterflies that did not survive were documented as having 14 days until eclosion Chrysalises were watched for a total of 16 days, but the last adult emerging occurred on day 13. Therefore, counting dead chrysalises as Day 14 was a convenient yet, conservative way to include all individuals even those that never eclosed. RESULTS Intermittent Mist and Butterfly Development In the dry tank, 15 H. charithonia emerged (n = 17) and 2 died while 12 H. hecale zuleika (n = 17) survived and 5 died; This represents 88% survival for H. charithonia and 71% for H. hecale zuleika Ten of each species survived in the misted environment, with both species having 7 deaths therefore 59% surviv al for both (Figure 4 ) 27 total butterflies survived in the dry tank, while only 20 in the wet tank survived Figure 4 Number of survivors of H. charithonia and H. hecale zuleika in each climate treatment. Seventeen chrysalises of each species were exposed to dry and intermittent mist conditions for 16 days and survivorship noted. Dark gray represents the intermittent mist treatment while light grey represents the dry. Intermittent mist was provided by Zoo Med Reptile Fogger Terrarium Humidifier with a controller that would shut the mist off after 95% humidity reached. Overall, the misted environment received 30 minute s of mist an hour. The mist had no significant effect on survival of the two species (chi square = 0.095, df = 1, p > .05). 0 2 4 6 8 10 12 14 16 H. Charithonia H. Hecale zuleika Survivorship Species

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Although more survived in the dry tank, t here was no si gnificant effect of mist on survival of the two species of butterflies (chi square = 0.095, df = 1, p > .05 ). The average mass for H. charithonia i n the dry environment was 0.19 +/ 0 .02 g and in the wet was 0.15 +/ 0 .02 g a difference of 0.4g These weights were not statistically different though were nearly so (t = 2. 00, df = 24, p = 0.06). The average mass for H. hecale zuleika in the dry tank was 0 .27 +/ .03 g and in the wet tank 0 .35 +/ .03 g The wet tank had on average, butterflies that were 0 .12g larger than those in the wet tank, however, there was no significant difference between the two tanks ( t = 1.642 df = 21 p = 0 .11 ) The wing status of the H. charithonia differed significantly between the wet and dry tanks. I n the wet tank the butterflies had an average score of 2.5, while in the dry the averag e was 1.2 S ince a lower score signified a better wing condition, the dry tanks had wings in significantly better conditions (t=2.841, df=24, p=.009 ). H. hecale zuleika also showed a difference between the two environments ( t=2.524, df=21, p=.02 ). In the wet tank, the average score was 2.5, while in the dry, the score was 1.25 Again, the butterflies in the dry tank had wings of significantly higher quality. For wing length of H. charithonia the average length was 4.19 +/ 0.16 cm in the dry and 3.53 +/ 0.2 cm in the wet, this difference is statistically significant (t=2.528, df=24, p=.02). The H. hecale zuleika had an average length of 4.23 +/ 0.12 cm in the dry tank and a similar average of 4.16 +/ 0.14 cm in the wet tank. These differences were not statistically significant (t=.379, df=21, p=.71). For the day s to eclosion, the species and climate were both considered. Climate was not shown to have an effect on eclosion (Two Way ANOVA, F= 1.14, df= 1, p=.29) The average days to eclosion in the intermittent mist climate was 11 .0 +/ 0.68 while in the dry environment the average was 10. 3 +/ 0.56 0 .7 days less than the misted (Figure 5 ). Figure 5 Average days +/ SE to eclosion under two different climates for H. charithonia and H. hecale zuleika Both environments had 17 chrysalises of each species. Intermittent mist involved on and off mist during the treatment by Zoo Med Reptile Fogger Terrari um Humidifier while the control was a dry environment. The butterflies were watched for a total of 16 days. Climate did not have an effect on the rate of eclosion ( Two Way ANOVA, F= 1.14, df= 1, p=.29). 8 8.5 9 9.5 10 10.5 11 11.5 12 Intermittent Mist Dry Days to eclosion Climate

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Species also did not statistically affect rate of eclosion (Two Way ANOVA, F=3.61, df=1, p=.06). H. charithonia eclosed on average after 10.0 +/ 0.64 days, while H. hecale zuleika exlosed after 11.2 +/ 0.60 day s, about a day later on average (Figure 6 ). Figure 6 Average +/ SE days to eclosion for H. charithonia and H. hecale zuleika Seventeen chrysalises of each species were placed in dry and intermittent misted environments and monitored for 16 days. Intermittent mist was from the Zoo Med Reptile Fogger Terrarium Humidifier; intermittent mist delivered 30 minutes of mist per hour. The species did not have an effect on rate of eclosion ( Two Way ANOVA, F=3.61, df=1, p=.06). The interaction between species and climate also showed no effect on eclosion (Two Way ANOVA, F=.02, df=1, p=.89 Figure 7 ). H. hecale zuleika eclosed after a similar amount of time i n both climate conditions; after 11.6 +/ 0.53 days in the mist, in the dry environment t hey eclosed after 10.9 +/ 0.67 days (Tu key s HSD test p> .05). H. charithonia also eclosed after similar amounts of time in both tanks; 10.3 +/ 0.82 days to eclosion in the mist and 9.8 +/ 0.45 days in the dry (Tukey s HSD test p>.05). Although both species showed a slower time to eclosion when exposed to intermittent mist, this differe nce was not significant 8 8.5 9 9.5 10 10.5 11 11.5 12 12.5 H. Charithonia H. Hecale zuleika Days to eclosion Species

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Figure 7 Average +/ SE eclosion rates for H. hecale zuleika and H. charithonia exposed to intermittent mist and dry environments. Light gray represents the dry environment while dark gray shows the misted. Seventeen chrysalises of each species were exposed to intermittent and dry conditions. Intermittent mist was from a Zoo Med Reptile Fogger Terrarium Humidifier and would cause the tank to be filled with fog for 30 minutes of every hour. Species and climate showed no effect on days to eclosion ( Two Way ANOVA, F=.02, df=1, p=.89) Intense Mist and Butterfly Development In the dry tank a total of 15 H. charithonia emer ged (n = 17) while 4 died. Of the H. hecale zuleika 14 emerged (n = 17) and 1 died. H. charithonia h ad 79% of the butterflies survive; H. hecale zuleika had 93%. In the intensely fogged tank, 6 H. charithonia emerged and 14 died. One H. hecale zuleika emerged and 14 died. 30% of the H. charithonia survived in the fogged and 7% of the H. hecale zuleika (Figure 8 ) Overall, 2 9 butterflies emerged from the dry tank and 7 emerged from the fogged tank 0 2 4 6 8 10 12 14 H. hecale zuleika H. charithonia Days to eclosion Species

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Figure 8 Number of survivors of H. charithonia and H. hecale zuleika in each climate treatment. Seventeen chrysalises of each species were exposed to dry and intermittent mist conditions for 16 days and survivorship noted. Dark gray represents the constant mist treatment while light gray represents the dry. Constant mist was provided by Zoo Med Reptile Fogger Terrarium Humidifier for 10 hours a day. The mist had a significant effect on survival of H. charithonia (chi square = 3.86 d f = 1, p < .05). The mist also significantly caused a decrease in survival for H. hecale zuleika (chi square = 11.3, df=1, p < .05). Four times as many emerged from the dry than from the intensely mi sted tank H. charithonia showed a significant difference in survivorship between the two climates (chi square = 3.86, df = 1, p < .05). H. hecale zuleika also showed a difference in survival between dry and fogged environments (chi square = 11.3, df = 1, p < .05). In the dry and fogged tanks, the average masses were very similar for H. charithonia, 0 .17 +/ 0.04 and 0 .18 +/ 0.01 g respectively. These masses were not statistically significant (t = 0 .180, df = 20, p = 0 .8594). H. hecale zuleika also showed no difference between masses in both climates (t = 0 .043, df = 14, p = 0 .9663). The average mass in the dry was 0.29 +/ 0.05 g, only 0 .01g less than the average mass in the wet tank, 0 .30 +/ 0.03 g. The average wing status for H. charithonia in the dry tank was 2.1, which was very similar to the s tatus in the wet tank, 2.2; these data were not statistically significant (t=.046, df = 20, p = .9637). H. hecale zuleika also did not show a significant difference between tank environme nts (t = .709, df = 14, p = .4906). The average status in the dry was 1.9, and the average in the wet was 1.0 Wing length between the two tanks of H. charithonia was not significantly different (t = 0 .666, df = 20, p = 0 .51330 ) Average wing length of the H. charithonia in the dry environment was 3.64 +/ 0.10 cm, only 0 .21 cm less than the wing length of those in the wet tank, 3.85 +/ 0.17 cm. The H. hecale zuleika also did not have differences between wing sizes (t = 0 .873, df = 14, p = 0 .3987). In the dry tank, the wing length was 4.22 +/ 0.18 cm and 0.06 cm less in the misted tank, 4.16 +/ 0.09 cm. Climate was shown to have an effect on rates of eclosion (Two Way ANOVA, F = 19.07, df = 1, p < .0001) In the dry tank, the average days to eclosion was 8.6 +/ 0.5 0 2 4 6 8 10 12 14 16 H. charithonia H. hecale zuleika Survivorship Species

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almost 3 days less th an those in the wet tank, 11.8 +/ 0.49 (Tukey s HSD test p<.05 Figure 9 ) Figure 9 Average days +/ SE to eclosion under two different climates for H. charithonia and H. hecale zuleika Both environments had 17 chrysalises of each species. Intense mist involved 10 hours of mist by Zoo Med Reptile Fogger Terrarium Humidifier while the control was a dry environment. The butterflies were watched for a total of 16 days. Climate was shown to have an effect on the rate of eclosion those in the dry tank eclosed earlier ( Two Way ANOVA, F = 19.07, df = 1, p < .0001 ). Species also significantly affected eclosi on (Two Way ANOVA, F = 9.96, df = 1, p = .0024 Figure 10 ). H. charithonia eclosed overall earlier than H. hecale zuleika ; H. charithonia waited 9.3 +/ 0.46 days before eclosing, while H. hecale zuleika came out after 11.5 +/ 0.53 days (Tukey s HSD test p<.05) 0 2 4 6 8 10 12 14 Intense Mist Dry Days to eclosion Climate 0 2 4 6 8 10 12 14 H. charithonia H. hecale zuleika Days to eclosion Species

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Figure 10 Average +/ SE days to eclosion for H. charithonia and H. hecale zuleika Seventeen chrysalises of each species were placed in dry and intermittent misted environments and monitored for 16 days. Constant mist was from the Zoo Med Reptile Fogger Terrarium Humidifier; mist was delivered constantly for 10 hours a day. The species did have an effect on rate eclosion H. charithonia eclosed earlier on average ( Two Way ANOVA, F = 9.96, df = 1, p = .0024 ). The climate and species interaction did not significantly affect eclosion rates (Two Way ANOVA, F = 3.60, p = .0622 Figure 11 ). H. hecale zuleika while in the fogged tank, took 12.3 +/ 0.50 days to eclose, while in the dry environment took about two days less, 10.6 +/ 0.60 These results were not statistically different (Tukey s HSD test p>.05). H. charithonia did differ significantly between the wet and dry tanks (Tukey s HSD test p<.05). In the fogged tank they took 11.5 +/ 0.20 days and in the dry 7.0 days, about 4.5 days less. Within the dry tank, the two butterfly species differed significantly (Tukey s HSD test p<.05). The H. charithonia took 7.1 days and the H. hecale zuleika took 10.6 +/ 0.40 days. Figure 11 Average +/ SE eclosion rates for H. hecale zuleika and H. charithonia exposed to intermittent mist and dry environments. Light gray represents the dry environment while dark gray shows the misted. Seventeen chrysalises of each species were expo sed to intermittent and dry conditions. Constant mist was from a Zoo Med Reptile Fogger Terrarium Humidifier and would cause the tank to be filled with fog for 10 hours a day Species and climate showed no effect on days to eclosion ( Two Way ANOVA, F = 3.60, p = .0622 ). DISCUSSION Overall, the intermittent mist did not have a significant effect on survival or rate of eclosion; there was ample time for the chrysalises to dry and it did not affect their survival and eclosion T he mist did negatively affect wing status of H. charithonia ; H. charithonia had much better quality of wings in the dry tank Wing length was also significantly different; H. charithonia had significantly smaller wings in the wet tank. Also the mass of H. charithonia was shown to be nearly significant between the two tanks; the misted tank 0 2 4 6 8 10 12 14 H. charithonia H. hecale zuleika Days to eclosion Species

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had smaller H. charithonia on average. This could be due to the fact that H. charithonia is not well evolved to deal with misty conditions since it is found from 0 1200 meters, while H. hecale zuleika may be slightly more adapted since it can be found in the Monteverde Cloud Forest. The intensely mist ed environment played a big role in eclosion and survival The intense mist caused the days to eclosion to increase and it also killed most of the butterflies Constant rain has been shown to affect the survival and development of butterflies in their chrysalis (Clifford 2008). The intense mist did not affect the mass, wing length, and wing status of the butterflies. This could just reflect the small number of surviving individuals in constant mist conditions. O verall both species performed worse in the presence of intense mist. This constant mist is more ch aracteristic of Monteverde weather, t herefore, as the mist recedes and the number of dry days increases the conditions improve for these butterflies to move up in elevation. H. charithonia may move up to Monteverde for breeding and living, also, H. hecale zuleika may increase their breeding and time in Monteverde. The populations of butterflies in Monteverde may change to include those that ar e used to dry, warm conditions. These new species will impact host plants, nectar sources, and highland endemic but terflies. Future studies could involve testing the effect of mist on endemic highland species to see if lowland species could displace the endemics that are more adapted to misty conditions. ACKNOWLEDGEMENTS I would like to thank Alan Masters, for hi s guidance and advice while completing this project Thanks to the University of Georgia for allowing me to use their lab space and Thanks also to Amber Brossard, for assisting me in getting materials to the lab. Lastly, thank you to Katie Bradshaw, Hanna h Crane, Amber Brossard, Nathan Sellers, Brendan Boyer, and Daniel Brunelle for keeping me company in the lab. LITERATURE CITE D Clifford, K. 2008. The Effects of Precipitation on Lepidopteran Chrysalis Mortality, Eclosion time and Wing Size CIEE Spring. Debinski, D.M., R.E. VanNimwegen, M.E. Jakubauskas. 2006. Quantifying relationships between bird and butterfly community shifts and environmental change. Ecological Applications 16: 380 393. DeVries, P.J. 1987. The Butterflies of Costa Rica and their Natural History. Princeton University Press. Princeton. Pp 194, 189. Houghton, J.T., Y. Ding, D.J. Griggs, M. Noguer, P.J. van der Linden, X. Dai, K. Maskell, C.A.

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Johnson. 2001. Climate Change 2001: The Scientific Basis. Intergovernmental Panel on Climate Change. http://www.csun.edu/~hmc60533/CSUN_630E_S2004/climate%20change/climate_ change_2001_tech_summary.pdf 4/4/2011 McLaughlin J.F., J.J. Hellmann, C.L. Boggs, P.R. Ehrlich. 2002. Climate change hastens population extinctions. PNAS 99. 70 74. Murphy, D. D., and S. B. Weiss. 1992 Effects of climate change on biological diversity in Western North America: Species losses and mechanisms. Chapter 26 in Global Warming and biological diversity ed. R. L. Peters and T. E. Lovejoy. Castleton, New York: Hamilton Printing Pollard E. 1988. Temperature, Rainfall and Butterfly Numbers. Journal of Applied Ecology 25: 3. 819 828. Pounds, J.A ., M.P.L Fogden, J.H. Campbell. 1999. Biological response to climate change on a tropical mountain. Nature 398 611 615. Roy, D.B., P. Rothery, D. Moss, E. Pollard, J.A. Thomas. 2001. Butterfly numbers and weather: predicting historical trends in abundance and the future effects of climate change. Journal of Animal Ecology 70: 201 217. Smetacek, P. 2009. Detrimental effec ts of low atmospheric humidity and forest fire on a community of western Himalayan butterflies. Journal of Threatened Taxa 3: 1694 1701. Walther, G., E. Post, P. Convey, A. Menzel, C. Parmesan, T.J.C. Beebee, J. Fromentin, O. Hoegh Guldberg, F. Bairlein. 2002. Ecological responses to recent climate change. Nature 416. 389 395.


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The cloud bank in Monteverde, Costa Rica is rising due to global warming. This is causing fewer misty days and is impacting the habitats of local organisms. Butterflies have been shown to be negatively impacted by moisture in terms of livelihood. Heliconius charithonia, a butterfly species occurring from 0-1200 meters, and Heliconius hecale zuleika, occurring from 0-1700 meters, were exposed to intermittent (five minutes mist,
five minutes dry over the course of the day) and intense mist conditions (10 hours of constant mist a day) to measure the effect of mist on survivorship, days to eclosion, and development (in terms of mass, wing length,
and wing quality). As the climate of Monteverde becomes dryer, these butterflies may move up in elevation to take advantage of the dry conditions. Intermittent mist had no effect on survival or days to eclosion. Twenty butterflies eclosed from the intermittent mist tank and 27 from the dry. In dry conditions, 88% (n=17) of H.
charithonia survived while 71% (n=17) of H. hecale zuleika. Ten of each species survived in the misted environment, therefore 59% (n=17) survival for both. The intermittent mist did significantly decrease wing quality, wing length, and mass for H. charithonia, but not for H. hecale zuleika. For wing length of H. charithonia, the average length was 4.19 +/- 0.16 cm in the dry and 3.53 +/- 0.2 cm in the wet. The masses of H. charithonia were nearly significantly different between the two climates; in the dry tank their mass was
0.19 +/- 0.02 g and in the wet was 0.15 +/- 0.02 g. H. charithonia may not be as well adapted to mist as H. hecale zuleika, since H. charithonia showed decreased development. Constant mist killed most of butterflies and caused the eclosion rates of those that did eclose to increase. A total of seven butterflies eclosed from the constant mist and 29 from the dry. In the dry tank, H. charithonia had 79% (n=17) of the butterflies survive; H. hecale zuleika had 93% (n=17). In the wet tank, 30% (n=17) of the H. charithonia survived and 7% (n=17) of the H. hecale zuleika. In the dry tank, the average days to eclosion was 8.6 +/- 0.5, almost 3 days less than those in the wet tank, 11.8 +/- 0.49; Constant mist was not shown to affect development, but probably because
of the low number of butterflies that eclosed. Overall, as the mist recedes and the number of dry days increases, the conditions improve for these butterflies to move up in elevation. H. charithonia may move up to Monteverde for breeding and living, also, H. hecale zuleika may increase their breeding and time in Monteverde.
El banco de nubes en Monteverde, Costa Rica est subiendo debido al calentamiento global. Esto est causando menos das nubosos y est impactando el hbitat de organismos locales. Las mariposas han mostrado un impacto negativo por la humedad en trminos de sobrevivencia. Heliconius charithonia es una mariposa que se encuentra entre los 0-1200 metros y Heliconius hecalezuleika se encuentra entre los 0-1700 metros, fueron expuestos a niebla intermitente (5 minutos con niebla, 5 minutos en lo seco) y en condiciones intensas (10 horas constantes de niebla durante el da) para medir el efecto de la sobrevivencia, das para eclosionar y desarrollar (en trminos de masa, largo del ala y calidad del ala).A medida que el clima de Monteverde se convierta ms seco, estas mariposas pueden ascender a mayores elevaciones para aprovechar las condiciones secas. La neblina intermitente no tiene efecto alguno en la sobrevivencia o das en eclosionar. 20 mariposas eclosionaron de la niebla intermitente y 27 del ambiente seco. En condiciones secas, 88% (n=17) de H. charithonia sobrevivieron mientras que en un 71% (n=17) los de H. hecale zuleika. Diez de cada especie sobrevivi en el ambiente con niebla constante. La niebla intermitente disminuyo significativamente la calidad de las alas, el tamao del ala y la masa para H. charitona pero no para H. hecalezuleik. La masa de H. charitona fue diferente entre los dos ambientes, en el tanque seco la masa fue de 0.19 +/- 0.02 g y en el hmedo fue de 0.15 +/- 0.02 g. H. charithonia puede estar no tan bien adaptada a la niebla como la otra especie ya que demuestra una disminucin en el desarrollo. Un total de siete mariposas eclosionaron de la niebla constante y 29 del seco. En el tanque seco H. charithonia tiene un 79% de sobrevivencia (N=17) y H. hecalezuleika un 93% (N=17). En el tanque hmedo un 30% (N= 17) de H. charithonia sobrevivi y en un 7% (N=17) de H. hecalezuleika. En el tanque seco el promedio de das de eclosin fue de 8.6 +/- 0.5, casi 3 das ms que en el tanque hmedo 11.8 +/- 0.49. La niebla constante no muestra un efecto en el desarrollo pero probablemente debido al nmero bajo de mariposas que eclosionaron. Sobre todo, al aumentar el nmero de das secos, las condiciones se prestan para que estas mariposas se muevan a elevaciones mayores, H. charithonia puede moverse a Monteverde para reproducirse y vivir, adems H. hecalezuleika puede aumentar su reproduccin y tiempo en Monteverde.
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