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Activity patterns of damselflies and dragonflies (Odonata) in Monteverde, Costa Rica

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Title:
Activity patterns of damselflies and dragonflies (Odonata) in Monteverde, Costa Rica
Translated Title:
Patrones de actividad de las libélulas (Odonata) en Monteverde, Costa Rica ( )
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English
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Gaynor, Jenny
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Subjects / Keywords:
Dragonflies--Animal behavior--Costa Rica--Puntarenas--Monteverde Zone   ( lcsh )
Damselflies--Animal behavior--Costa Rica--Puntarenas--Monteverde Zone   ( lcsh )
Libélulas--Comportamiento animal--Costa Rica--Puntarenas--Zona de Monteverde
Tropical Ecology Summer 2008
Ecología Tropical Verano 2008
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Reports   ( lcsh )
Reports

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Abstract:
Daily periods of activity tend to vary across different species of dragonflies and damselflies of the order Odonata. The purpose of this study was to investigate the activity patterns of five different behaviors for five different species of odonates. Observations were made at the Estación Biológica de Monteverde over a weeklong period in late July. Results showed that there were variations in behavior compared with time both within and across the five species. However, each species did not display every behavior, and behaviors were not evenly distributed across different species. This may be a result of variance in population sizes or that observations were not conducted at peak times of activity for certain species.
Abstract:
Los periodos diarios de actividad tienden a variar a través de las diferentes especies de libélulas del orden Odonata. El propósito de este estudio fue investigar los patrones de actividad de los cinco diferentes comportamientos de las cinco especies diferentes de odonatos. Las observaciones se realizaron en la Estación Biológica de Monteverde por un periodo de una semana a finales de julio.
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Text in English.
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usfldc doi - M39-00003
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Activity patterns of damselflies and dragonflies (Odonata) in Monteverde, Costa Rica Jenny Gaynor Department of Biology, University of Wisconsin – Ma dison ABSTRACT Daily periods of activity tend to vary across diffe rent species of dragonflies and damselflies of the order Odonata. The purpose of this study was to investig ate the activity patterns of five different behavio rs for five different species of odonates. Observations w ere made at the Estacin Biolgica de Monteverde ov er a weeklong period in late July. Results showed that there were variations in behavior compared with tim e both within and across the five species. However, each species did not display every behavior, and behaviors were not evenly distributed across differ ent species. This may be a result of variance in population sizes or that observations were not cond ucted at peak times of activity for certain species RESUMEN Los perodos diarios de actividad tienden a variar en las diferentes especies de liblulas y gallitos del diablo. El objetivo de este estudio fue investigar los patrones de actividad de cinco especies de odon atos en cuanto a cinco diferentes comportamientos. Las obse rvaciones se realizaron en la Estacin Biolgica de Monteverde durante una semana a finales de julio. L os resultados mostraron que existen variaciones en el comportamiento a lo largo del tiempo, tanto dentro como entre las cinco especies. Sin embargo, cada especie no mostr cada conducta, y no se distribuye uniformemente en las diferentes especies. Esto pue de ser el resultado de la varianza en los tamaos de p oblacin o que las observaciones no se llev a cabo en las horas punta de actividad de algunas especies. INTRODUCTION Most dragonflies (Anisoptera) and damselflies (Zygo ptera) of the order Odonata spend the majority of their adult life near some kind of water source (Esquivel 2006). However, daily periods of activity tend to vary across diffe rent species and also greatly depend on the amount of sunlight available. Furthermore, per iods of activity also fluctuate between males and females. Females tend to only make brief visits to aquatic sites in order to mate and lay their eggs. Males, on the other hand, will spend most of their time near a water source, waiting for females to arrive (Esquiv el 2006). In a study by Bick and Bick (1965), it was found that within the species A. apicalis females were at a water source for only 20% of their lives, but mated 89% of their days there. Males, on the other hand, spent 44% of their lives at water and only mated du ring 20% of their days there. Male odonates are known to be territorial at the wa ter’s edge. The defense of areas around oviposition sites is critical for drag onflies and damselflies to have reproductive success (Switzer 2002). It has been f ound that there is a positive correlation between quality of oviposition substrate and both f emale densities and rates of males intruding into territories (Wolf et al. 1997). Mal es guard their territory by making patrol flights or by perching on a prominent place. In a study by Kirkton and Schultz (2001), it was found that during 25% of total observation time males performed intraspecific interactive behaviors such as patrol flights, court ship displays, mate guarding, and male-

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male aerial dogfights. In another study it was fou nd that for the dragonfly Perithemis tenera individual males defended a single territory for about half of the time that the aquatic site was occupied (Switzer 2002). When a f emale enters the territory of a male, the male has unrestricted access to the female for reproduction. However, females only arrive intermittently and male individuals will fig ht each other when one invades another’s territory (Esquivel 2006). Males of different species living in the same habit at tend to avoid competition for space. They can accomplish this by mating at diffe rent times, perching at different heights, using different types of perches, or using different areas of the site (Esquivel 2006). This study aims to explore behavioral activ ity patterns of multiple species in a single area. It was hypothesized that different sp ecies would have distinct and differing times of peak activity in order to avoid competitio n for resources. METHODS Study Site The study was conducted near the Estacin Biolgica de Monteverde, Costa Rica at a waterfall of the Quebrada Mquina about 100 m down from the lower lab. The site is at an elevation of 1,465 m, and is surrounded by steep hills on either side that are inhabited by many tall canopy trees. Because of this, sun ra rely struck the entire area. In the morning, the sun generally only hit the hill on the west bank. Sun exposure over the water was most intense at noon, and in the afternoo n the sun traveled up the opposite bank. Study subjects Four species within the suborder Zygoptera and one within Anisoptera were studied. The identification of these species occurred during pre liminary observations with the use of electronic field guides (Haber 2004). Argia anceps (Coenagrionidae), Argia underwoodi (Coenagrionidae), Hetaerina majuscula (Calopterigidae), and Palaemnema sp. (Platystictidae) are damselfly species and Brechmorhoga rapax (Libelullidae) is a dragonfly species. Argia anceps averages 38 mm in length. The males’ bodies are almost completely blue and females are mostly brown The species stays active for most of the day, but becomes inactive during cloudy peri ods. Argia underwoodi is very similar to A. anceps but males do not have quite as much blue on their bodies. It averages 40 mm in length, has small black spots on the wing tip s, and is very abundant on vegetation near water sources. Hetaerina majuscula is a much larger damselfly and ranges between 56–58 mm long. It is black in color and males have bright red spots on the base of their wings, which gives it its common name, “Rubyspots.” Palaemnema sp. is brownishblack and has yellow bands at each segment of its a bdomen. Males are 56-58 mm long while females are usually 48-50 mm long. Little is known about the habits of Palaemnema sp. and it is rarely seen because it spends the ma jority of its time in dark environments and has little activity. The dragonfl y B. rapax has turquoise green eyes, a dark brown thorax, and a black abdomen with a brigh t yellow ring around segment seven. It ranges from 45-55 mm in length, and the males an d females are very similar in

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appearance. Brechmorhoga rapax is the most common species of Brechmorhoga in Mesoamerica. Behaviors After preliminary observations, five behaviors to i nvestigate of the aforementioned species were determined. The first of which is agg ressive behavior. This was defined as any aggressive interaction or fight between two or more individuals. This included when one individual would chase an intruder out of his t erritory. Also, the dragonfly B. rapax exhibited aggressive behavior when two individuals engaged in energetic circular flight patterns at very high speeds. The second behavior was a patrolling behavior. This occurred when a dragonfly or damselfly would act te rritorially and stay in the same small area when there was no individual in close proximit y. Dragonflies would generally patrol an area by hovering close to the water within an ar ea of about one square meter. Damselflies tended to patrol by leaving their perch ing site to survey the area for only a second before returning to the exact same spot. Th e third behavior observed was denoted a “hopping” behavior. This occurred when odonates would fly in an up-and-down motion about a half a meter above the water’s surfa ce. This flight pattern made it appear that the individuals were repeatedly bouncing or ho pping a few inches up and down. The fourth observed behavior was mating and was counted as any time a male and female were seen in tandem. Finally, the fifth behavior o bserved was perching. A perching behavior was counted when the individual was seen p erforming the act of landing on its perching site. An individual that was already perc hed without seeing it land was not counted. Furthermore, patrolling behavior of damse lflies was not counted as perching. Methodology Data were collected during a weeklong period in lat e July between the hours of 9:30 a.m. and 2 p.m. When any of the aforementioned behavior s were observed, I marked down which behavior it was, as well as the time and whic h species performed it. Weather conditions were also noted. I generally stayed in areas where there was sunlight; therefore, in the morning I would begin my observat ions on the west slope. As the day progressed I moved down the hill to the water, and then slightly up the slope on the other side of the water in the afternoon. The data were analyzed using half hour time slots b etween 9:30 am and 2 p.m. I made analyses of behaviors based on time performed by each individual species, and also looked at behavior times across different species. Statistical analyses were performed using the chi-squared test to determine if the beha viors varied depending on the time of day within and across species. RESULTS I found that A. anceps exhibited only aggressive, mating, and perching be haviors (Fig. 1). Peak aggressive activity for A. anceps was observed between 12:00 and 12:29 p.m., and

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peak activity for both mating and perching was seen between 12:30 and 12:59 p.m. Argia underwoodi exhibited aggressive, patrolling, and perching beh aviors (Fig. 2). All three behaviors showed peak activity between 10:30 and 10:59 a.m. Brechmorhoga rapax showed aggressive, patrolling, mating, and perchin g behaviors (Fig. 3). Patrolling and perching were both most frequent between 11:00 and 11:29 a.m., peak aggressive activity was between 12:30 and 12:59 p.m., and mati ng behavior was only seen between 1:00 and 1:29 p.m. Hetaerina majuscula showed aggressive, patrolling, hopping, and perching behaviors (Fig. 4). Aggressive, patrollin g, and perching behaviors were most frequent between 10:30 and 10:59 a.m., and peak act ivity for hopping behavior was between 11:00 and 11:29 a.m. Palaemnema sp. only showed mating and perching behaviors, and peak activity for both was observed between 1:00 and 1:29 p.m. (Fig. 5). Argia underwoodi showed the highest number of aggressive behaviors for a specific time period (Fig. 6). Across all species, B. rapax displayed the most total patrolling behaviors, but for a single time period A. underwoodi patrolled most frequently (Fig. 7). Hetaerina majuscula was the only species to perform the hopping behavi or. Argia anceps showed the highest frequency of mating behaviors (Fig. 8). Finally, both A. underwoodi and H. majuscula showed very high frequencies of perching behaviors between 10:30 and 10:59 a.m. The chi-squared analyses showed statistical signif icance for the distribution of all behaviors within and across species except for perc hing behavior of B. rapax ( l2 = 11; df = 8; P = 0.2). DISCUSSION The data demonstrate that different species at the same site have different and distinct periods of activity for various behaviors. Within the same species, distribution of behaviors was not constant throughout the entire du ration of observation. There were peak time periods when behaviors were observed, and different behaviors of the same species did not always peak at the same time. Acro ss different species, it was found that the temporal distribution of specific behaviors als o varied. This agrees with the hypothesis that different species would perform dif ferent behaviors at different times of the day. Behaviors were not evenly distributed across all t he species. No species displayed every behavior, and some behaviors were very seldom ly displayed. For instance, the hopping behavior was only observed 14 times, and on ly performed by H. majuscula The unequal distributions of behaviors may indicate tha t there is a lot of variation between population sizes of the different species at the wa terfall site. Another explanation for the unequal distribution o f behaviors could be that observations were not conducted at optimal times fo r certain species. However, due to weather conditions and time constraints it was not possible to make more observations later in the afternoon. Around 2:00 p.m. the sun h ad traveled too far up the bank for it to be possible to see substantial odonate activity. F urthermore, frequent rains in the afternoon worsened the conditions for observation. This may have also caused discrepancies in the results because weather condit ions were much more favorable for observation of odonate behavior in the morning. In a study on the behavior of Argia apicalis it was found that peak activity

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occurred around noon (Bick and Bick 1965). In the results from this study it was found that A. anceps displayed peak activities between 12:00 and 1:00 p .m. while A. underwoodi showed peak activity between 10:30 and 10:59 a.m. In the study by Bick and Bick (1965) it was also found that aggressive a ctivity drove off 87% of all intruders and that the most important kinds of aggressive act ivity were both strong flights towards intruders without physical contact and patrolling f lights of an area. This correlates with the high amounts of aggressive behaviors observed b y A. underwoodi It’s possible that the number of aggressive behavi ors as well as patrolling behaviors could be indicators of a hierarchy of dominance amo ng the five observed species. Argia underwoodi and B. rapax displayed the most combined frequencies of aggress ive and patrolling behaviors; however, they had differing p eak periods of activity. Argia underwoodi did have similar peak periods of activity to H. majuscula Hetaerina majuscula had much less combined aggressive and patrolling b ehaviors, indicating that A. underwoodi may be a more territorial species. It was also ob served that A. underwoodi and H. majuscula often had aggressive interspecies encounters, and also used very similar perches; therefore, it is likely that there exists territoriality between these two species. Similarly, B. rapax and A. anceps had similar periods of peak activity for combined aggressive and patrolling behaviors. They also occ upied similar areas and were observed performing aggressive behaviors, so it is likely th at territoriality exists between these two species as well. Not only did the species A. underwoodi display higher frequencies of aggressive and patrolling behaviors than H. majuscula it is also much smaller in size. In a study by Marden and Waage (1990), it was found that the amou nt of energy reserves that males had at the end of aggressive contests had stronger correlations with winning these contests than size or physical attributes related t o flight ability. However, if A. underwoodi is indeed more territorial than H. majuscula then it is curious that more mating behaviors were not seen of A. underwoodi One might expect that greater reproductive success would usually follow higher te rritoriality. Recommendations for future studies involving behav ior analysis of odonates would be to make observations over a longer time period. Starting earlier in the morning and ending later in the afternoon would be beneficial i f species were more active during those times. Another suggestion would be to choose a stu dy site that receives sun over its entire area for the whole duration of observations. The waterfall site created problems because different areas were not sunny at the same time. It would also be interesting to observe how weather conditions affect activity acro ss different species. ACKNOWLEDGEMENTS I would first like to thank Tania Chavarria for all her help when the weather presented problems for m y initial project. I wouldn’t have had any data with out you. Also, big thanks to Pablo Allen Monge and Luke Hillman for helping me catch and mark dragonflies a nd damselflies…even though I ended up not needing that data in the least when my project changed. Th ank you to Karen Masters for showing me what I want to be like when I “grow up.” Thank you, Izzy Place for your sweatshirt. Also, thank you for going t o the same school as me. Finally, thanks to all my fello w students for keeping me on my toes and making thi s an incredible AND incredibly fun experience. You guys are neat. LITERATURE CITED

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Bick, G. H. and J. C. Bick. 1965. Demography and Behavior of the Damselfly, Argia apicalis (Say), (Odonata: Coenagriidae). Ecology 46(4): 46 1-472. Corbet, P. S. 1980. Biology of Odonata. Annual R eview of Entomology 25: 189-217. Esquivel, C. 2006. Dragonflies and damselflies of Middle America and t he Caribbean Instituto Nacional de Biodiversidad (INBio), Santo Domingo de Heredia, Costa Rica. Haber, W. A. 2004. Electronic Field Guide Project UMass-Boston. Kirkton, S. D. and T. D. Schultz. 2001. Age-Speci fic Behavior and Habitat Selection of Adult Male Damselflies, Calopteryx maculata (Odonata: Calopterygidae). Journal of Insect Behavior 14(4): 545-556. Marden, J. H. and J. K. Waage. 1990. Escalated da mselfly territorial contests are energetic wars of attrition. Animal Behavior 39: 9 54-959. Switzer, P. V. 2002. Individual Variation in the Duration of Territory Occupation by Males of the Dragonfly Perithemis tenera (Odonata: Libellulidae). Annals of the Entomological Society of America 95(5): 628-636. Wolf, L. L., E. C. Waltz, D. Klockowski, and K. Wak eley. 1997. Influences on Variation in Territorial Tenures of Male White-Face d Dragonflies ( Leucorrhinia intacta ) (Odonata: Libellulidae). Journal of Insect Behav ior 10(1): 31-47.

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n n rn n rn n rn nr n rn rn nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n rn rn n nr n n rn rn n nrn n rn rn n n nnnrn nnnr !rn" # nrn n rn rn n n nnnrn nnnr !rn" # nrn n rn rn n n nnnrn nnnr !rn"# $.'.& !$'&()*-rn" # $%& !$'&()*r!+rn" # $ nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '& !$'&()rnnrn r nnnnr" # $& !$'&$ nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) n rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) n rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) nr n rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % .nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nr n n rn /--n n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn" # $,& !$'&()*-rn" # $.%& !$'&()*--n nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n # $%.& !$'&()*r!-n" # $,& !$'&()rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n "#$%.&!$'&()*r!-n"#$,&!$'&()rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n "#$%.&!$'&()*r!-n"#$,&!$'&()rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n "#$%.&!$'&()*r!-n"#$,&!$'&()rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n "#$%.&!$'&()*r!-n"#$,&!$'&()rn nrn n rn rn n n nnnrn nnnr !rn"#$.'.&!$'&()*-rn"#$%&!$'&()*r!+rn"#$ '&!$'&()rnnrn rnnnnr"#$&!$'&$ ) % nrn n rn /--n n n nnnrn nn nnnr rn"#$,&!$'&()*-rn"#$.%&!$'&()*--n "#$%.&!$'&()*r!-n"#$,&!$'&()rn

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' ' % nr rn % nr rn % nr rn % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr rn n % % nr n rn n % % nrn rn n n nnnrn -nnnr !+rn % % nrn rn n n nnnrn -nnnr !+rn # $,,& !$'&()r!-n" # $.& !$'&()rn % % nrn rn n n nnnrn -nnnr !+rn # $,,& !$'&()r!-n" # $.& !$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r% % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r% % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r% % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n 0r-r1 % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnr r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 nnnr" # $& !$&() % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 nnnr"#$&!$&() % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 nnnr"#$&!$&() % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 nnnr"#$&!$&() % % nrn rn n n nnnrn -nnnr !+rn "#$,,&!$'&()r!-n"#$.&!$'&()rn % nnrn r! !n 0r-r1 /+r2r n nnnrnrnr-n 3 nnnr"#$&!$&()

PAGE 10

' ' nnr !n nnr !n % nnr !n 0r-r1 % nnr !n 0r-r1 % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr !n 0r-r1 /+r2r % % nnr n !n 0r-r1 /+r2r % % nnrn !n 0r-r1 /+r2r n nnn-rnrnr-n 3 % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr" # $& !$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr" # $& !$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r% % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r% % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r% % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnr n r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 nnnr" # $%.& !$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 nnnr"#$%.&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 nnnr"#$%.&!$%&() % % nnrn !n 0r-r1 /+r2rn nnn-rnrnr-n 3 nnnr"#$&!$%&() nnrn r0r-r1 rr++r n nnn+rnrnr-n3 nnnr"#$%.&!$%&()

PAGE 11

. . . r. nnr r. nnr r. nnr r! !n nnr r! !n 0r-r1 nnr r! !n 0r-r1 % nnr r! !n 0r-r1 % nnr r! !n 0r-r1 /+r2r % nnr r! !n 0r-r1 /+r2r % nnr r! !n 0r-r1 /+r2r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr r! !n 0r-r1 /+r2r rr++r % nnr n r! !n 0r-r1 /+r2r rr++r % nnr n r! !n 0r-r1 /+r2r rr++r % nnrn r! !n 0r-r1 /+r2r rr++r n! nnn-nrnr-n 3 % nnrn r! !n 0r-r1 /+r2r rr++r n! nnn-nrnr-n 3 nnnr" # $.& !$%&() % nnrn r! !n 0r-r1 /+r2r rr++r n! nnn-nrnr-n 3 nnnr"#$.&!$%&() % nnrn r! !n 0r-r1 /+r2r rr++r n! nnn-nrnr-n 3 nnnr"#$.&!$%&() % nnrn r! !n 0r-r1 /+r2r rr++r n! nnn-nrnr-n 3 nnnr"#$.&!$%&()


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Gaynor, Jenny
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Patrones de actividad de las liblulas (Odonata) en Monteverde, Costa Rica
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Activity patterns of damselflies and dragonflies (Odonata) in Monteverde, Costa Rica
260
c 2008-08
500
Born Digital
3 520
Daily periods of activity tend to vary across different species of dragonflies and damselflies of the order Odonata. The purpose of this study was to investigate the activity patterns of five different behaviors for five different species of odonates. Observations were made at the Estacin Biolgica de Monteverde over a
weeklong period in late July. Results showed that there were variations in behavior compared with time both within and across the five species. However, each species did not display every behavior, and behaviors were not evenly distributed across different species. This may be a result of variance in
population sizes or that observations were not conducted at peak times of activity for certain species.
Los periodos diarios de actividad tienden a variar a travs de las diferentes especies de liblulas del orden Odonata. El propsito de este estudio fue investigar los patrones de actividad de los cinco diferentes comportamientos de las cinco especies diferentes de odonatos. Las observaciones se realizaron en la Estacin Biolgica de Monteverde por un periodo de una semana a finales de julio.
546
Text in English.
650
Dragonflies--Animal behavior--Costa Rica--Puntarenas--Monteverde Zone
Damselflies--Animal behavior--Costa Rica--Puntarenas--Monteverde Zone
4
Liblulas--Comportamiento animal--Costa Rica--Puntarenas--Zona de Monteverde
653
Tropical Ecology Summer 2008
Ecologa Tropical Verano 2008
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Reports
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CIEE
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t Monteverde Institute : Tropical Ecology
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u http://digital.lib.usf.edu/?m39.3