Nesting behavior in the resplendent quetzal (Pharomachrus mocinno: Trogonidae)


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Nesting behavior in the resplendent quetzal (Pharomachrus mocinno: Trogonidae)
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Carr, Amber
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The Resplendent Quetzal (Trogonidae) lives in the cloud forests of Central America. In Costa Rica, nest boxes have been placed in the Monteverde Cloud Forest Preserve to increase the population sizes of these birds, which are listed as near threatened by C.I.T.E.S. Cameras placed inside the boxes from 2000 to 2003 as web cams and captured stills inside the nests. I used these four years of captured images and made external observations of two nests to study if there were differences in parental investment for incubation and rearing. I found evidence that 1. During incubation the female spent more total time in the nest than the male (Nest 2 tied p-value =0.0001). 2. The male had significantly more visits per day during incubation than the female (Nest 3 tied p-value =0.0360 and Nest 4 tied p-value =0.0391). 3. The female had significantly more visits per day during rearing than the male (Nest 1 tied p-value =0.0209). 4. The female and male had significantly more total time in the nest for incubation than rearing (Nest 1 female tied pvalue = < 0.0001 male tied p-value =0.001 and Nest 8 female tied p-value =0.02853 and male tied p-value = <0.001) and the mean visits were significantly higher for incubation than rearing for both males and females (Nest 1 female tied p-value =<0.0001 and male tied p-value = < 0.0001, Nest 6 female tied p-value = 0.0315, and Nest 8 female tied p-value = 0.0285 and male tied p-value = < 0.0001). 5. The female had a significantly higher number of visits to the nest during rearing than incubation (Nest 1 tied p-value =0.0097 and Nest 6 tied p-value =0.0031). These results suggest that the female invests more in the young than the male. The female might be willing to invest more in her young due to the fact that she initially invests more in her egg than the male does in his sperm. The study of the nesting behavior of these birds is important to their conservation. Fledging success of their young will be what determines whether their populations will be threatened by extinction. ( ,, )
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El Quetzal (Trogonidae) vive en los bosques nubosos de América Central. En Costa Rica se han instalado cajas para nidos en la Reserva Biológica del Bosque Nuboso de Monteverde para aumentar las poblaciones de estas aves que están clasificadas como amenazadas en C.I.T.E.S. Se instalaron cámaras dentro de las cajas del año 2000 al 2003 y se tomaron fotos del interior de los nidos. Yo usé estos cuatro años de fotos y observé dos nidos para determinar si había alguna diferencia en la inversión paternal en la incubación y cría de los polluelos. Se encontró evidencia de que: 1. La hembra paso más tiempo en el nido que el macho durante la incubación (p del Nido 2 =0.0001); 2. El macho tuvo más visitas por día que la hembra durante la incubación (p del Nido 3 =0.0360 y p del Nido 4 =0.0391); 3. La hembra tuvo más visitas por día que el macho durante el período de crianza (p del Nido 1 =0.0209); 4. La hembra y el macho tuvieron más tiempo total dentro del nido durante la incubación que durante la crianza (p del Nido 1 para la hembra =<0.0001, p del Nido 1 para el macho =0.001 y p del Nido 8 para la hembra =0.02853 y para el macho=<0.001) y las visitas promedio fueron significativamente más altas durante la incubación que durante la crianza tanto para los machos como para las hembras (p del Nido 1 para la hembra = < 0.0001 y para el macho = < 0.0001, p del Nido 6 para la hembra =0.0315, y Nido 8 para la hembra =0.0285 y para el macho = < 0.0001). 5. La hembra presentó un número mayor de visitas al nido durante la crianza (p del Nido 1=0.0097 y p del Nido 6 = 0.0031). Estos resultados sugieren que la hembra invierte más tiempo que el macho en los polluelos. Es posible que la hembra invierta más tiempo en sus polluelos porque ha invertido más en sus huevos que lo que el macho ha invertido en su esperma. El estudio del comportamiento de anidación de los quetzales es importante para su conservación. La sobrevivencia de los polluelos será lo que determine si sus poblaciones serán amenazadas por la extinción.
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Nesting Behavior in the Resplendent Quetzal Pharomachrus mocinno : Trogonidae Amber Carr Department of Ecology and Environmental Sciences, University of Maine ABSTRACT: The Resplendent Quetzal Trogonidae lives in the cloud forests of Central Amer ica. In Costa Rica, nest boxes have been placed in the Monteverde Cloud Forest Preserve to increase the population sizes of these birds, which are listed as near threatened by C.I.T.E.S. Cameras placed inside the boxes from 2000 to 2003 as web cams and cap tured stills inside the nests. I used these four years of captured images and made external observations of two nests to study if there were differences in parental investment for incubation and rearing. I found evidence that 1. During incubation the femal e spent more total time in the nest than the male Nest 2 tied p value =0.0001. 2. The male had significantly more visits per day during incubation than the female Nest 3 tied p value =0.0360 and Nest 4 tied p value =0.0391. 3. The female had significan tly more visits per day during rearing than the male Nest 1 tied p value =0.0209. 4. The female and male had significantly more total time in the nest for incubation than rearing Nest 1 female tied p value =<0.0001 male tied p value =0.001 and Nest 8 female tied p value =0.02853 and male tied p value =<0.001 and the mean visits were significantly higher for incubation than rearing for both males and females Nest 1 female tied p value =<0.0001 and male tied p value =<0.0001, Nest 6 female tied p value =0.0315, and Nest 8 female tied p value =0.0285 and male tied p value =<0.0001. 5. The female had a significantly higher number of visits to the nest during rearing than incubation Nest 1 tied p value =0.0097 and Nest 6 tied p value =0.0031. These resu lts suggest that the female invests more in the young than the male. The female might be willing to invest more in her young due to the fact that she initially invests more in her egg than the male does in his sperm. The study of the nesting behavior of th ese birds is important to their conservation. Fledging success of their young will be what determines whether their populations will be threatened by extinction. RESUMEN: El Quetzal Trogonidae vive en los bosques nubosos de América Central. En Costa Ri ca se han instalado cajas para nidos en la Reserva Biológica del Bosque Nuboso de Monteverde para aumentar las poblaciones de estas aves que están clasificadas como amenazadas en C.I.T.E.S. Se instalaron cámaras dentro de las cajas del año 2000 al 2003 y s e tomaron fotos del interior de los nidos. Yo usé estos cuatro años de fotos y observé dos nidos para determinar si había alguna diferencia en la inversión paternal en la incubación y cría de los polluelos. Se encontró evidencia de que: 1. La hembra paso m ás tiempo en el nido que e l macho durante la incubación p del Nido 2 =0.0001; 2. El macho tuvo más visitas por día que la hembra durante la incubación p del Nido 3 =0.0360 y p del Nido 4 =0.0391; 3. La hembra tuvo más visitas por día que el macho dur ante el período de crianza p del Nido 1 =0.0209; 4. La hembra y el macho tuvieron más tiempo total dentro del nido durante la incubación que durante la crianza p del Nido 1 para la hembra =<0.0001, p del Nido 1 para el macho =0.001 y p del Nido 8 para la hembra =0.02853 y para el macho=<0.001 y las visitas promedio fueron significativamente más altas durante la incubación que durante la crianza tanto para los machos como para las hembras p del Nido 1 para la hembra =<0.0001 y para el macho =<0.0001, p del Nido 6 para la hembra =0.0315, y Nido 8 para la hembra =0.0285 y para el macho=<0.0001. 5. La hembra presentó un número mayor de visitas al nido durante la crianza p del Nido 1=0.0097 y p del Nido 6 =0.0031. Estos resultados sugieren que la hemb ra invierte más tiempo que el macho en los polluelos. Es posible que la hembra invierta más tiempo en sus polluelos porque ha invertido más en sus huevos que lo que el macho ha invertido en su esperma. El estudio del comportamiento de anidación de los que tzales es

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importante para su conservación. La sobrevivencia de los polluelos será lo que determine si sus poblaciones serán amenazadas por la extinción . INTRODUCTION: The Resplendent Quetzal Trogonidae is the only of five Quetzal species that lives in Central America. The Resplendent Quetzal is divided into two subspecies: Pharomachrus m ocinno is found in Mexico, Guate mala, Honduras, and Nicaragua, while P. m. costaricensis is found in Costa Rica and western Panama Johnsgard 2000. The northern sub species€ longer upper tail coverts and more prominent crest on the males€ head can tell the two apart Skutch 1944. Both have iridescent plumes like a hummingbird. The Resplendent Quetzal is found in cloud forests between 1,000 to 3,300 m in elevatio n Johnsgard 2000. In Costa Rica they are restricted to the highlands of the Tilarán, Central Volcanic, and Talamanca mountain ranges García Rojas 2006. The diet of the adult quetzal is mostly fruits from the Lauraceae family. After they digest the peri carp they regurgitate the seed and act as important seed dispersers for some plant species Avila et al 1996. Wheelwright 1983 found that peaks in breeding season in Monteverde, Costa Rica for the Resplendent Quetzal correlated with peaks in fruiting o f species from the Lauraceae. Both male and female take part in excavating the nest cavity, incubating the eggs and caring for the young Fogden and Fogden 1996. Hole cavities are rare and both sexes may be needed to establish ownership. The same nest is often used for a number of years, or a new hole may be made in the same tree but farther down on the tree because of rot Fogden and Fogden 1996. There is often competition for nests Fogden and Fogden 1996. To help protect these birds that are classif ied as near threatened by CITIES Johnsgard 2000. The Monteverde Cloud Forest Preserve has taken measures to help conserve the Resplendent Quetzal by putting up nest boxes Thaden 2002. The MVCFP contains 10,500 hectares of forest lying mostly in the l ower montane wet forest life zone Haber et al 2000. The preserve is located between the elevations of 1550 1850 m Haber 2000. In the year 2000 the MVCFP placed cameras in some of the quetzal nesting boxes that were known to be active nesting sites. The se cameras took pictures and documented incubation and rearing behavior of quetzals every 20 seconds for the years of 2000, 2001, 2002, and 2003. There were some instances where the cameras took pictures every hour or two hours. Since accurate observations could not be made these data were not used. Some of the pictures can be viewed on the webpage http://www.cloudforestalive.org . There have been two previous studies done by C.I.E.E. students using images fro m the cameras for nest building, egg incubation, and chick rearing. Heller 2001 observed nesting behavior between the two sexes for one nest in 2000, which included incubation of eggs and raising the chicks. Her results showed that the female spent signi ficantly more time than the male for both incubation and rearing. Thaden 2002 observed nest building. Her results showed that for the pair that was not monitored by the camera in the nest box quetzalcam the female spent significantly more time building the nest than the male, but for the quetzalcam pair there was no significant difference between the amount of time spent on nest building. Even though Thaden 2002 showed that the female invested more time in nested building and Heller 2001 showed that the female invested more time than the male in egg incubation and chick rearing both studies had small sample sizes, for just one Heller 2001 or two Thaden 2002 nests.

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Fortunately, data for eight nests are now available from camera equipped nest boxe s. In addition, I took external observations from two nests, which were located in or near the MVCFP. I used these 10 nests to see if there are general patterns of differential investment for Resplendent Quetzal males and females throughout incubation and chick rearing. I expected to see that the female spent more time incubating the eggs and rearing the chicks than the male. Trivers 1972 argues, since the female invests more in their eggs initially then the males do in their sperm, they will also invest more in the clutch itself. I also expected to see that the total time spent in the nests and the mean visit duration would be greater for incubation than rearing, while the number of visits would be greater during rearing than incubation. During rearing t he parents would be expected to only bring food to their young and then leave, while during incubation the parents would be required to stay in the nest for extended periods of time to keep the eggs at a constant temperature. MATERIALS AND METHODS: Quetz alcam Data: I processed images from camera nest boxes for four years, between 2000 and 2003. I looked for differences between male and female investment as 1. Total time spent per day for incubation and rearing, 2. Total number of visits per day for incub ation and rearing, and 3. Mean visit duration per day for incubation and rearing. The quetzalcam took a picture of the nest every 20 seconds. Each picture was labeled with the date and time the picture was taken. The data for the four years were placed o n 64 compact disks. The disks were labeled with the year and date ranges for the pictures they contained. I looked at the disks recording down the year, date, time duration of visits, and the sex of the parent in the nest. Only the male or the female could be in the nest at a certain time so there were no overlapping visits. I also noted the period of the chicks€ development. I divided it into three periods incubation, transition, and rearing. The incubation period was when all eggs in the nest were un hatc hed. The transition period was when there was at least one un hatched egg and at least one hatched chick in the nest at the same time. Rearing was defined as when all eggs in the nest were hatched. The times for the male and female quetzal were recorded by looking at the time they entered the nest and the time they exited. The beginning time was then subtracted from the exiting time to calculate the total number of minutes spent inside the nest. If a visit was less than a minute it was disregarded to simpl ify calculations. Since the female is the only one that sleeps in the nest with the chicks those times were ignored. I classified the female as sleeping between the times when the female entered the nest and did not leave until morning, even if she was mov ing in the nest during that time. The period of time when the female was sleeping usually occurred between 18:00  06:00. In the 64 compact disks that were given to me from the reserve there were only data for nest building on April 18 th 2002, so this was discarded since it was only a period of one day. The data were also insufficient for the transition periods between incubation and rearing since they were only available for Nest 1 and Nest 8 see Table 1. These periods had no statistical analysis done, but were included in graphs for each nest where data were available.

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Since the locations of the cameras were not recorded and whether it was the same or a different pair of quetzals from year to year were not recorded on the compact disks, I treated each c amera as a different nest even though it could be the same pair for different years or the same pair during the same year, but with their second clutch. There were a total of two nests for 2000; three nests for 2001, two nests for 2002, and one nest for 20 03 for a total of eight nests during the four years see Table 1. Direct Observations: External observations were made at two different nests during 2006. One of these nests Nest 9 was located within the MVCFP on Sendero Bosque Nuboso. I observed the nest from a distance of 9m and 11cm with 8x42 binoculars. The nest was located in one of the artificial nest boxes. My observations of this nest occurred on four days: 4/22/06, 4/25/06, 4/29/06, and 5/1/06. During this time only observations of incubation were observed see Table 1. Nest 10 was located outside of the reserve. It was along the road leading to the reserve just above the Trapp Family Lodge. This nest was observed at a distance of 16m with 8x42 binoculars. This nest was located in an old sna g within view from the road. My four observations of this nest took place on 4/23/06, 4/27/06, 4/30/06, and 5/2/06. Only observations of incubation behavior were recorded during this time see Table 1. Table 1: The number of days for each period in c hick development is shown below for nests monitored in Monteverde, Costa Rica. The year for each nest is shown as well as comments on chicks€ survival. Nests 1 8 were monitored by a quetzalcam while Nests 9 and 10 were externally observed. Period in chic k development Number of days Nest number Year Incubation Transition Rearing Comments 1 2000 16 9 15 Fledged 2 2000 16 0 0 Egg broke 3 2001 7 0 0 4 2001 6 0 0 5 2001 0 0 16 Fledged 6 2002 14 0 4 Chicks eaten 7 2002 1 0 9 Fledged 8 2003 17 8 12 Fledged 9 2006 4 0 0 10 2006 4 0 0 RESULTS: Quetzalcam data: 2000: For Nest 1 there were sixteen days of incubation, nine days of transition and fifteen days of rearing. At the end of the rearing period one of the chicks fledged because the othe r never hatched and was rolled out by the male see Table 1. For Nest 1 there was no significant difference between the sexes during incubation for the total time spent in the nest per day, the total number of visits to the nest per day, or the mean visit duration per

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day. For rearing the female made significantly more visits per day than the male tied p value = .0209; see Figure 1 A. Females spent significantly longer amount of time in the nest during incubation than rearing tied p value = <.0001; see Figure 1 B. Females for this nest also made significantly more visits to the nest during rearing than incubation tied p value = <.0001; see Figure 1 A.. The mean of each visit to Nest 1 was significantly more for females during incubation than rearing tied p value =<.0001; see Figure 1 C. The male also spent significantly longer amount of time in the nest during incubation than rearing tied p value = <.0001; see Figure 1 B.. The means of visits during incubation were significantly higher than rear ing for males tied p value =<.0001; see Figure 1 C. For Nest 2 only data for incubation of the eggs were available for both sexes. The incubation period was sixteen days and was shortened due to the male breaking the only egg during incubation on 6/17/0 0 see Table 1. The female spent significantly more time in the nest see Figure 2 B and made significantly more visits per day than the male See Figure 2 A for this nest during incubation tied p value = .0001 and tied p value = .0474 respectively. 2001: For Nest 3 only incubation data were available for both sexes. The incubation period was seven days long see Table 1. The male of this nest made significantly more visits per day during incubation than the female tied p value = .0360 see Figu re 3 A. For Nest 4 only incubation data were available for both sexes. The incubation period was six days long see Table 1. The male for this nest also made significantly more visits per day than the female for incubation tied p value = .0391 see Fi gure 4 A. For Nest 5 only rearing data were available for both sexes. The rearing period was sixteen days long when both chicks fledged see Table 1. There were no significant differences between total time spent in the nest per day see Figure 5 B, vi sits per day see Figure 5 A, or the mean duration of visits see Figure 5 C during rearing for Nest 5. 2002: For Nest 6 there were data for both incubation of eggs and rearing of the chicks. The incubation period was fourteen days long and the rearing period was four days long. The rearing period was ended prematurely due to a weasel entering the nest and eating both of the chicks on 5/16/02 see Table 1. At this nest however only the female made visits to the nest so there are no data for differences between sexes. The female made significantly more visits per day during rearing than incubation tied p value = .0031; see Figure 6 A. The mean duration of visits was significantly higher for incubation than rearing for the female tied p value =.0315; se e Figure 6 C. For Nest 7 there were data for both sexes during incubation and rearing. The incubation period was one day long while the rearing period was nine days long. After the rearing period the chick successfully fledged see Table 1. There were no significant differences between the amount of time each sex invested in the eggs or young, nor were there significant differences for each sex between the incubation and rearing periods see Figure 7 A, B, and C. 2003: For Nest 8 there were data for i ncubation and rearing for both sexes. The incubation period was seventeen days long, the transition period was eight days long, and the rearing

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period was twelve days long. Only one chick fledged successfully after the rearing period because an un hatched egg was rolled out of the nest see Table 1. There were no differences between the sexes for the amount of time devoted to caring for young during incubation and rearing. The female spent significantly more time in the nest during incubation than rearing tied p value = .02853; see Figure 8 B. The female also had a significantly higher mean visit duration for incubation than rearing tied p value = .0285; see Figure 8 C. The male also spent significantly more time in the nest during incubation than reari ng see Figure 8 B and had a significantly higher mean duration of visits for incubation than rearing see Figure 8 C; tied p value =<.0001 and tied p value =<.0001 respectively. Personal Observations: 2006: For Nest 9 only partial incubation data were taken from direct observations outside the nest. The incubation period that was documented was four days long see Table 1. The female spent more total time than the male in the nest during observation periods see Figure 9 B. The male had a greater number of visits per day than the female for incubation see Figure 9 A. Both of these observations correspond with the general trends found with the quetzalcam nests see Table 2. On the last day the behavior of the quetzals changed and I think that it is possible that at least one of the eggs hatched. I cannot prove that rearing had begun though because no food items were observed being brought to the nest. For Nest 10 only partial incubation data were taken from direct observations outside the nest. The incubation period that was documented was four days long see Table 1. The male spent more total time in the nest than the female during incubation see Figure 10 B. This does not correspond with the general trend that the female spends significantly more time in the nest than the male during incubation see Table 2. The male had a greater number of visits per day than the female for incubation see Figure 10 A. This does agree with the general trend found with the quetzalcam nests see Table 2. T able 2: The different investment in young between sexes for periods of incubation and rearing for nests monitored in Monteverde, Costa Rica. Nests 1 8 were monitored by quetzalcams while Nests 9 and 10 were externally monitored. The female had a greater nu mber of visits than the male during rearing for Nest 1. For Nest 2 the female had a greater number of visits and a greater amount of total time than the male during incubation. The male had a greater number of visits than the female during incubation for N est 3 and Nest 4. Incubation Rearing Nest Number of visits Total time Mean visit duration Number of visits Total time Mean visit duration 1 Female 2 Female Female 3 Male 4 Male 5 6 7 8

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9 10 Table 3: The different investment in young for males and females during incubation and rearing in nests monitored in Monteverde, Costa Rica. . Nests 1 8 were monitored by quetzalcams while Nests 9 and 10 were externally monitored. The female in Nest 1 had a greater number of visits during rearing than incubation. Both the male and the female for nest 1 had a higher total time and mean visit duration during incubation than rearing. The female in nest 6 had a higher number of visits in rearing than incubation and a higher mean visit duration in incubation than rearing. Both the male and the female for Nest 8 had higher total time and mean visit duration for incubation than rearing. Female Male Nest Number of visits Total time Mean visit duration N umber of visits Total time Mean visit duration 1 Rearing Incubation Incubation Incubation Incubation 2 3 4 5 6 Rearing Incubation 7 8 Incubation Incubation Incubation Incubation 9 10 DISCUSSION: The general trends found that were significant were 1. That during incubation the female spent more total time in the nest than the male see Table 2. 2. The male had significantly more visits per day during incubation than the female see Table 2. 3. T he female had significantly more visits per day during rearing than the male see Table 2. 4. The female and male had significantly more total time in the nest and the mean visit durations were significantly higher for incubation than rearing see Table 3 . 5. The female had a significantly higher number of visits to the nest during rearing than incubation see Table 3. Nesting behavior for quetzals can be divided into three main categories, they are nest building, egg incubation, and chick rearing. Thade n 2002 found that for one of her pairs, a pair not monitored by the quetzalcam, the female spent a significantly greater amount of time than the male excavating the nest. For her other pair, the one monitored by the quetzalcam, there was no significant d ifference between sexes for the time spent building the nest. I found that during incubation one of my six tested nests with incubation data Nest 2 had a female who spent significantly more time in the nest than the male. These results are then same tha t Heller 2001 found in her study with the quetzalcam data for 2000. These results are the same because my Nest 2 was the same nest as Heller€s. I also found that during incubation the male had significantly higher visits per day than the female for two o ut of my six tested nests with incubation data Nest 3 and Nest 4. This implies that these visits were of shorter time and a lower investment to incubating the

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eggs, since these did not add up to a greater total amount of time spent in the nest than the f emale. In one of my nests Nest 2 the female not only spent a significantly more time incubating the eggs but also had more visits per day to the nest. This shows that she invested more than the male in incubating the eggs. Only Nest 1 showed that the fem ale had significantly more visits per day than the male for rearing. This may be because this female might have brought fruits to the young. According to Wheelwright 1983, females generally tend to bring more fruit to the nest during rearing while males bring small insects. Young that have a diet of fruits are slower growing than when they have a diet of insects and fruits which may be the reason why the female comes back more often than the male Wheelwright 1983. These results contradict what Wheelwri ght 1983 found. During his study, the male made more visits than the female at the first nest for rearing while at the second nest, or second clutch, he found no significant difference between the sexes and the number of visits per day during rearing. I also found there to be no difference between the sexes and the number of visits per day during rearing for Nests 5, 7, and 8. My results suggest that the female spends either equal or a greater time on nesting than the male. These trends would leave the fe male at a higher risk to predation as mentioned by Heller 2001. Since the female is in the nest she is not able to see potential predators and is more vulnerable than the male. The female may take this risk because according to Trivers 1972 the female invests more in her eggs than the male invests in his sperm, which would lead to her to invest more into the chicks wellbeing as they develop. In my study I also found at Nest 1 and 8 both the female and male spent more time in the nest with longer visits during incubation than rearing. at Nest 1 and Nest 8. At Nest 1 and Nest 6 the female made significantly more visits per day during rearing than incubation. This makes sense because the visits from the males and females would need to be longer during incu bation to keep the eggs at a constant temperature to develop, as opposed to during rearing when the male and female would be quickly carrying food to the chicks. According to Wheelwright 1983 nesting mortality was high during his study. My observations agree with these findings. Out of my eight nests from the quetzalcam, I observed both chicks in Nest 6 eaten by a weasel, an un hatched egg in Nest 1 and Nest 8 rolled out by male, and the only egg in Nest 2 being broken by the male. The Resplendent Que tzal is unique in that it nests in hole cavities, is sexually dimorphic and both sexes take active parts in raising the young. Most birds that nest in hole cavities need both sexes to protect the nest from competitors, but these birds such as kingfishers, motmots, and toucan are not sexually dimorphic. Birds that are sexually dimorphic such as the peacock, birds of paradise, or hummingbirds do not share responsibilities and the female is the one who is left to raise the young Fogden and Fogden 1996. Since there is a decrease in available habitat for the quetzal nesting availability may be a factor in the Resplendent Quetzal€s survival Johnsgard 2000. Programs to increase the quetzal€s populations, such as the nesting box program at the MVCFP, are importa nt to help keep numbers high. Protecting the range of migration is also very important since quetzals migrate to lower elevations outside of their breeding season. These habitats tend to have a greater percentage of habitat destruction Johnsgard 2000. R esearch on the bird€s nesting behaviors are important to understand because they are

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what are responsible for producing a viable population size so that the species doesn€t become more in danger of extinction. ACKNOWLEDGEMENTS: I would like to thank the Monteverde Cloud Forest Preserve for lending me the quetzalcam compact disks and letting me do observations there. Javier Méndez that you for requesting permission to borrow the compact disks I would like to thank Esteban showing me the locations of my tw o active nests. Alan Masters thank you very much for advising me throughout the project and making sure I was headed in the right direction and helping me get permission to borrow the compact disks from the Preserve. Ollie Hyman and Maria Jost thank you fo r your eternal support, patience and help with my statistical analysis and graphs. Finally, thanks to the whole group at the Estación Biológica de Monteverde for their wonderful atmosphere and positive attitudes. LITERATURE CITED: Avila H., M. L., V. H . Hernández O., and E. Velarde. 1996. The diet of Resplendent Quetzal in a Mexican cloud forest. Biotropica 28 4b: 720 727. Fogden, M., and P. Fogden 1996.The Resplendent Quetzal. Green Mountain Publishing, Monteverde, Costa Rica. García Rojas, M. 2006. Diet and Habitat Preference of the Resplendent Quetzal Pharomachrus mocinno costaricensis in Costa Rican Montane Oak Forest. Ecological Studies 185 : 325 336. Haber, W. A. 2000. Plants and vegetation. In Monteverde ecology and conservation of a tropical cloud forest, Nalini M. Nadkarni and Nathaniel T. Wheelwright, eds. Oxford University Press, New York, New York. Haber, W. A., W. Zuchowski, and E. Bello. 2000. An introduction to cloud forest trees: Monteverde Costa Rica. Mountain Gem Publications, Mo nteverde, Costa Rica. Heller, J. 2001. Nesting behavior of the female and male Resplendent Quetzal Pharomachrus mocinno . In C.I.E.E. Spring 2001 Tropical Ecology and Conservation. Johnsgard, P.A. 2000. Trogons and quetzals of the world. Smithsonian Ins titute Press, Washington. Skutch, A.F. 1944. Life History of the Quetzal. Condor 46 : 213 235. Thaden, E. S. 2002. Nest building behavior of the Resplendent Quetzal Pharomachrus mocinno : Trogonidae. Pages 157 165 in C.I.E.E. Spring 2002 Tropical Ecology and Conservation. Trivers, R.L. 1972. Parental Investment and sexual selection. In Sexual selection and the decent of man. B. Campbell ed.. Aldine, Chicago. In Animal Behavior and Evolutionary Approach. J. Alcock. Sunderland , Massachusetts. As found in Heller, J. 2001. Nesting behavior of the female and male Resplendent Quetzal Pharomachrus mocinno . In C.I.E.E. Spring 2001 Tropical Ecology and Conservation. Wheelwright, N.T. 1983. Fruits and the ecology of Resplendent Quetzals. Auk 100 :286 301.

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FIG URES: A. Number of visits to Nest 1 0 5 10 15 20 25 4/5/00 4/12/00 4/19/00 4/26/00 5/3/00 5/10/00 Date Number of Visits female male incubation transition rearing B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 4/5/00 4/12/00 4/19/00 4/26/00 5/3/00 5/10/00 Date Number of Minutes female male incubation transition rearing C. Mean minutes inside nest per visit 0 50 100 150 200 250 300 350 4/5/00 4/12/00 4/19/00 4/26/00 5/3/00 5/10/00 Date Mean Minutes female male incubation transition rearing Figure 1: The incubation period was between 4/5/00 4/20/00, there was a transition period between 4/21/00 4/29/00, and a rearing period between 4/30/00 5/14/00. The female made significantly more visits per day than the male during th e rearing stage and made significantly more visits to the nest during rearing than incubation A. The female and male spent a significantly longer time in the nest during incubation than rearing B. The mean of visit duration was significantly higher for females and males during incubation than rearing C.

PAGE 11

A. Number of Visits to Nest 0 1 2 3 4 5 6 7 8 9 10 6/2/00 6/3/00 6/4/00 6/5/00 6/6/00 6/7/00 6/8/00 6/9/00 6/10/00 6/11/00 6/12/00 6/13/00 6/14/00 6/15/00 6/16/00 6/17/00 Date Number of Visits female male B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 6/2/00 6/3/00 6/4/00 6/5/00 6/6/00 6/7/00 6/8/00 6/9/00 6/10/00 6/11/00 6/12/00 6/13/00 6/14/00 6/15/00 6/16/00 6/17/00 Date Number of Minutes female male C. Mean minutes inside nest per visit for Nest 2 0 50 100 150 200 250 300 6/2/00 6/3/00 6/4/00 6/5/00 6/6/00 6/7/00 6/8/00 6/9/00 6/10/00 6/11/00 6/12/00 6/13/00 6/14/00 6/15/00 6/16/00 6/17/00 Date Mean Minutes female male Figure 2: There was an incubation period between 6/2/00 6/17/00. The incubation period was prematurely shortened due to the male breaking the only egg on 6/17/00. The female spent signi ficantly more time in the nest B and made significantly more visits per day than the male A. There were no significant differences for mean visit duration between males and females C.

PAGE 12

A. Number of visits to Nest 3 0 1 2 3 4 5 6 7 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 4/21/01 Date Number of Visits female male B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 4/21/01 Date Number of Minutes female male C. Mean minutes inside nest per visit for Nest 3 0 50 100 150 200 250 300 350 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 4/21/01 Date Mean Minutes female male Figure 3: There was an incubation period betwee n 4/14/01 4/21/01. There were no data available for 4/15/01. The male made significantly more visits per day than the female A. There were no significant differences in the total number of minutes spent in the nest B between sexes. There also were not significant differences for mean visit duration between sexes C.

PAGE 13

A. Number of visits to Nest 4 0 1 2 3 4 5 6 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 Date Number of Visits female male B. Total number of minutes in the nest 0 100 200 300 400 500 600 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 Date Number of Minutes female male C. Mean minutes inside nest 0 50 100 150 200 250 300 350 400 4/14/01 4/15/01 4/16/01 4/17/01 4/18/01 4/19/01 4/20/01 Date Mean Minutes female male Figure 4: There was an incubation period between 4/14/01 4/20/01. There were no available data for 4/15/01. The male made significantly more visits per day than the femal e A. There were no significant differences in total time spent in the nest and sex of the parent B. There were also no differences between sexes for the mean visitation duration C.

PAGE 14

A. Number of visits to Nest 5 0 2 4 6 8 10 12 14 5/1/01 5/2/01 5/3/01 5/4/01 5/5/01 5/6/01 5/7/01 5/8/01 5/9/01 5/10/01 5/11/01 5/12/01 5/13/01 5/14/01 5/15/01 5/16/01 Date Number of Visits female male B. Total number of minutes in the nest 0 20 40 60 80 100 120 140 160 180 200 5/1/01 5/2/01 5/3/01 5/4/01 5/5/01 5/6/01 5/7/01 5/8/01 5/9/01 5/10/01 5/11/01 5/12/01 5/13/01 5/14/01 5/15/01 5/16/01 Date Number of Minutes female male C. Mean minutes inside nest per visit for Nest 5 0 5 10 15 20 25 30 35 5/1/01 5/2/01 5/3/01 5/4/01 5/5/01 5/6/01 5/7/01 5/8/01 5/9/01 5/10/01 5/11/01 5/12/01 5/13/01 5/14/01 5/15/01 5/16/01 Date Mean Minutes female male Figure 5: There was a rearing period between 5/1/ 01 5/16/01. There were no data available for 5/15/01. There were no significant differences between the number of visits per day A, the total number of minutes per day B, or the mean visit duration between the sexes C.

PAGE 15

A. Number of visits to Nest 6 0 2 4 6 8 10 12 14 16 18 20 4/22/02 4/24/02 4/26/02 4/28/02 4/30/02 5/2/02 5/4/02 5/6/02 5/8/02 5/10/02 5/12/02 5/14/02 5/16/02 Date Number of Visits female male incubation rearing B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 800 4/22/02 4/24/02 4/26/02 4/28/02 4/30/02 5/2/02 5/4/02 5/6/02 5/8/02 5/10/02 5/12/02 5/14/02 5/16/02 Date Number of Minutes female male incubation rearing C. Mean minutes inside nest 0 20 40 60 80 100 120 140 4/22/02 4/24/02 4/26/02 4/28/02 4/30/02 5/2/02 5/4/02 5/6/02 5/8/02 5/10/02 5/12/02 5/14/02 5/16/02 Date Mean Minutes female male incubation rearing Figure 6: T here was an incubation period between 4/22/02 5/10/02. There were no available data for the periods between 5/1/02 5/5/02 and 5/11/02 5/12/02. There was a rearing period between 5/13/02 5/16/02. This nest only had visits from the female so no male data are available. The female made significantly more visits per day during rearing than incubation A. Then mean duration of visits was significantly higher for incubation than rearing for the female C. There was no difference for the total time spent in the nest during incubation and rearing for the female B.

PAGE 16

A. Number of visits for Nest 7 0 5 10 15 20 25 5/17/02 5/18/02 5/19/02 5/20/02 5/21/02 5/22/02 5/23/02 5/24/02 5/25/02 5/26/02 5/27/02 Date Number of Visits female male incubation rearing B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 5/17/02 5/18/02 5/19/02 5/20/02 5/21/02 5/22/02 5/23/02 5/24/02 5/25/02 5/26/02 5/27/02 Date Number of Minutes female male incubation rearing C. mean minutes inside nest per visit for Nest 7 0 20 40 60 80 100 5/17/02 5/18/02 5/19/02 5/20/02 5/21/02 5/22/02 5/23/02 5/24/02 5/25/02 5/26/02 5/27/02 Date Mean Minutes female male incubation rearing Figure 7: There was a recorded incubation period for one day on 5/17/02. There is a rearing period between 5/18/02 5/27/02. There were no available data for 5/26/02. There were no signi ficant differences between the number of visits per day A, the total number of minutes per day B, or the mean visit duration between the sexes throughout the chicks€ development C.

PAGE 17

A. Number of visits to Nest 8 0 2 4 6 8 10 12 14 16 18 20 4/11/03 4/18/03 4/25/03 5/2/03 5/9/03 5/16/03 Date Number of Visits female male rearing incubation transition B. Total number of minutes in the nest 0 100 200 300 400 500 600 700 800 4/11/03 4/18/03 4/25/03 5/2/03 5/9/03 5/16/03 Date Number of Minutes female male incubation transition rearing C. Mean minutes inside nest 0 50 100 150 200 250 4/11/03 4/18/03 4/25/03 5/2/03 5/9/03 5/16/03 Date Mean Minutes female male incubation transition rearing Figure 8: There was an incubation period between 4/1 1/03 4/27/03, a transition period between 4/28/03 5/5/03, and a rearing period between 5/9/03 5/20/03. There were no available data for the period between 5/6/03 5/8/03. The female and male spent significantly more time in the nest during incubation that r earing B and they also both had a significantly higher mean for incubation than rearing C. There was no difference between sexes for the number of visits to the nest during the chicks development A.

PAGE 18

A. Number of visits to Nest 9 0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 4/22/06 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 Date Number of Visits female male B. Total number of minutes in the nest 0 50 100 150 200 250 300 4/22/06 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 Date Number of Minutes female male C Mean minutes inside nest per visit for Nest 9 0 50 100 150 200 250 300 4/22/06 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 Date Mean Minutes female male Graph 9: There were four days of obse rvation of incubation that took place on 4/22/06,4/25/06, 4/29/06, and 5/1/06. These observations were made at an artificial nesting box along the Sendero Bosque Nuboso in the MVCFP in Monteverde, Costa Rica. The female spent more time than the male in the nest B. The male had a greater number of visits than the female A. Both of these observations correspond with those that were found as significant trends. There were no differences between the sexes for the mean visit duration to Nest 9 C.

PAGE 19

A. Number of visits to nest 0 0.5 1 1.5 2 2.5 3 3.5 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 5/2/06 Date Number of Visits female male B. Total number of minutes in the nest 0 20 40 60 80 100 120 140 160 180 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 5/2/06 Date Number of Minutes female male C. Mean minutes inside nest 0 20 40 60 80 100 120 4/23/06 4/24/06 4/25/06 4/26/06 4/27/06 4/28/06 4/29/06 4/30/06 5/1/06 5/2/06 Date Mean Minutes female male Graph 10: Four days of observation for incubation were taken on 4/23/06, 4/27/06, 4/30/06, and 5/2/06. These observations were taken at a snag off the road close to the MVCFP near the Trapp Family Lodge in Monteverde, Costa Rica. The male spent more t ime in the nest than the female B. This does not correspond with the general trends found. The male also had a greater number of visits per day than the female A. This does correspond with the general trends. There were no differences between the mean visit duration and sex for Nest 10 C.


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