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Rhea-Fournier, Dylan J.
Fenologa y caros forticos de murcilagos en las flores Calyptrogyne brachystachys (Araceae) en el Bosque Nuboso de Monteverde
Phenology and phoretic bat mites of Calyptrogyne brachystachys (Arecaceae) flowers in the Monteverde Cloud Forest
Digitized by MVI
In November 2005 Calyptrogyne brachystachys palm inflorescences were examined for the presence of phoretic bat mites (tribe Melichari) in the Cloud Forest of Monteverde. Sixty-seven Calyptrogyne brachystachys plants were studied over a two-week period for flower and inflorescence phenology. Forty-six inflorescences were examined thoroughly for mite presence of which 11 were found to harbor mites. Flower phenology studies suggest a temporary flowering lag. Abundances of mites on both inflorescence and site scales were found at much lower frequencies than the Atlantic slope (Tschapka and Cunningham 2004), The lower abundance in Monteverde is attributed to abiotic differences such as recent weather patterns and biotic influences such as low flower abundance.
Se examinaron inflorescencias de la palma Calyptrogyne brachsrachys en Noviembre del 2005 para determinar la presencia o ausencia de los caros forticos en murcilagos (Tribu Melichari) en el bosque nuboso de Monteverde. Se estudiaron sesenta y siete plantas de Calyptrogyne brachsrachys por dos semanas para averiguar la fenologa de las flores e inflorescencias. Once de las cuarenta y seis inflorescencias examinadas contenan caros. Los estudios de la fenologa de las inflorescencias sugieren que hay un retraso temporal en la floracin. Las abundancias de caros, examinadas por escala del sitio e inflorescencia, ocurrieron con frecuencias ms bajas que en la Costa Atlntica de Costa Rica (Tschapka and Cunningham 2004). La abundancia ms baja tal vez sea debida a las diferencias abiticas tales como patrones climticos recientes e influencias biticas como la baja abundancia de las flores.
Text in English.
Monteverde Cloud Forest Reserve (Costa Rica)
Costa Rica--Puntarenas--Monteverde Zone--Monteverde
Reserva del Bosque Nuboso de Monteverde (Costa Rica)
Costa Rica--Puntarenas--Zona de Monteverde--Monteverde
Tropical Ecology Fall 2005
Monteverde Biological Research Station
Bat mites--Costa Rica
Ecologa Tropical Otoo 2005
Estacin Biolgica Monteverde
Acaros de murcilagos
t Monteverde Institute : Tropical Ecology
Phenology and phoretic bat mites of Calyptrogyne brachystachys Arecaceae flowers in the Monteverde Cloud Forest Dylan J. Rhea Fournier Department of Ecology and Evolutionary Biology, University of Oregon ABSTRACT In November 2005 Calyptrogyne brachy stachys palm inflorescences were examined for the presence of phoretic bat mites tribe Melichari in the Cloud Forest of Monteverde. Sixty seven Calyptrogyne brachystachys plants were studied over a two week period for flower and inflorescence phenology. Forty six inflorescences were examined thoroughly for mite presence of which 11 were found to harbor mites. Flower phenology studies suggest a temporary flowering lag. Abundances of mites on both inflorescence and site scales were found at much lower frequ encies than the Atlantic slope Tschapka and Cunningham 2004, The lower abundance in Monteverde is attributed to abiotic differences such as recent weather patterns and biotic influences such as low flower abundance. RESUMEN Se examinaron inflorescencia s de la palma Calyptrogyne brachsrachys en Noviembre del 2005 para determinar la presencia o ausencia de los Ã¡caros forÃ©ticos en murciÃ©lagos Tribu Melichari en el bosque nuboso de Monteverde. Se estudiaron sesenta y siete plantas de Calyptrogyne brachsra chys por dos semanas para averiguar la fenologÃa de las flores e inflorescencias. Once de las cuarenta y seis inflorescencias examinadas contenÃan Ã¡caros. Los estudios de la fenologÃa de las inflorescencias sugieren que hay un retraso temporal en la florac iÃ³n. Las abundancias de Ã¡caros, examinadas por escala del sitio e inflorescencia, ocurrieron con frecuencias mÃ¡s bajas que en la Costa AtlÃ¡ntica de Costa Rica Tschapka and Cunningham 2004. La abundancia mÃ¡s baja tal vez sea debida a las diferencias abiÃ³t icas tales como patrones climÃ¡ticos recientes e influencias biÃ³ticas como la baja abundancia de las flores. INTRODUCTION Commensalism when individuals of one species benefit from interactions with another species whose fitness does not change. Phoresy is a type of commensalism in which an organism travels on another solely for transport. This provides effective dispersal, particularly when the host animal predictably visits preferred patches frequently such as traplining nectavores or frugivores. Boggs 1 987. A phoretic communalistic relationship occurs between hummingbirds, some hummingbird pollinated flowers and a phoretic mite Gamasidae: Ascidae. Up to 200 species of phoretic hummingbird mites have been proposed to exist in the closely related gener a, Rhinoseius and Proctolaelaps Hyneman at al. 1991. These mites can be seen with the naked eye and do not obtain any nutrients from the bird, relying on their hosts solely for transport/dispersal, apparently disembarking in response to olfactory cues C olwell 1983. The mites feed on nectar and pollen. These mites have been found to be extremely specific to floral hosts Heyneman et al. 1991 found only 1 of 200 mites on a
flower host different than their regular hosts. Few flower hosts exist for a give n mite species but presumably enough to provide year round habitats. Heyneman et al 1991 also found that some hummingbird pollinated flowers that meet all apparent criteria for mites are not colonized at all. This suggests high specificity and recognitio n of flower hosts among hummingbird mites as well as criteria still unknown, such as attractant or repellent properties of nectars Heyneman et al. 1991. Tschapka and Cunningham 2004 recently discovered a complex involving phoretic mites on Calyptrogyn e ghiesbreghtiana Arecacea and bats Mammalia: Chiroptera in La Selva, Costa Rica. Eight different species of bats were collected, including two nectarivorous bats Phyllostomidae: Glossophaginae and two genera of primarily frugivorous bats Phyllosto midae: Stenoderminae; Artibeus and Carollia . Phoretic mites were only found Artibeus bats. Twenty nine of 73 Artibeus bats collected carried mites and as many as 19 mites were found on one individual bat. Mites were found on the wing surfaces of the bats, in the same general region as where pollen is collected. These mites Tribe: Melicharini were not found on any flowers other than Calyptrogyne ghiesbreghtiana . While hummingbird mites that have relationships with host plants that flower year round are mo nophagous, other hummingbird mites are polyphagous, having two or more hosts that they alternate due to seasonal phenology Colwell and Naeem 1993. Tschapka and Cunningham 2001 found one species of flower pollen besides C. ghiesbreghtiana Solanaceae: Markea neurantha on Artibeus spp. harboring phoretic mites. Studies of M. neurantha and other bat pollinated flowers have not yielded the presence of these mites, suggesting that these flower mites are truly monophagous. The flowers that were studied for presence and colonization of mites excluded crawling mites by covering the peduncle of the inflorescence with glue. Seventy six percent of the flowers surveyed were found to have flower mites. Tschapka and Cunningham 2004 concluded that flower mites of C . ghiesbreghtiana used Artibeus bats as phoretic carriers. They went further in suggesting that mites were not found on other bats due to feeding behavior of the bats. Nectarivorous bats spend less than one second at each flower, while hummingbirds can hov er for minutes. Frugivorous bats on the other hand perch to feed on nectar, allowing body contact with the inflorescence and longer visitation than hovering bats Tschapka and Cunningham 2004. Monteverde, Costa Rica is the home for populations of both f rugivorous 18 species, mainly of the genera Carollia and Artibeus and nectarivorous 7 species bats Timm and LaVal 2000 as well as Calyptrogyne brachystachys palms Haber 2000 . Artibeus toltechnus is the most abundant fruit bat in Monteverde, the pri mary pollinator of C. brachystachys species locally and is in the same genus as the bats found carrying mites in La Selva. Calyptrogyne brachystachys and C. ghiesbreghtiana are very similar an d in fact Henderson et al 1995 describe the former as a higher elevation phenotype of the latter, with narrower leaflets and base constriction. La Selva, Costa Rica is the first and only place that phoretic bat mites have been documented Tschapka and Cunningham 2004, thus the geographical distribution of these mite s are widely unknown. Given that Monteverde has similar species to those found in the La Selva, determining if a similar complex is present here in Monteverde is
imperative to broadening knowledge of bat mite biogeography. Furthermore due to abiotic differ ence s between La Selva and Montever de, confirmation of bat mite presence will shed light on how environmental factors affect colonization of these mites and what factors are limiting. Elevation has potential effects on plant flowering phenology patterns a nd therefore colonization success of mites. The diversity of species involved in hummingbird flower mite complexes has found to decrease with elevation Heyneman et al. 1991, implying a decrease in diversity in such relationships, and in parallel, bat flo wer mite complexes, in the higher elevation Cloud Forests in Monteverde. Increased seasonality in Monteverde compared to La Selva may also impose influences onto the species interactions involved in a potentially bat mite complex. Seasonal changes in plan t phonologies are likely to be greater on the Pacific side of the continental divide, possibly increasing seasonality of flowering patterns and associated folivores . In addition local weather patterns in Monteverde are much different than those found in La Selva. While both locations receive a substantial amount of rain, Monteverde also experiences periods of high wind and misty conditions. These types of weather patterns may very likely influence pollination, phoresy and bat activity. The goal of this stud y was to reproduce Tschapka and Cunningham s study at a higher elevation, within Monteverde Cloud Forest Reserve to determine if a phoretic mite bat flower complex exists. METHODS Study Sites Fieldwork was conducted between November 3 rd and 19 th , 200 5. Site 1 was located in a primary forest fragment between the Bullpen Pasture on the Campbell Property and Windy Corner, approximately 0.5 km from La Reserva de Monteverde, Costa Rica. This site experienced high frequencies of wind gusts, providing cons tant disturbance and resulting in low canopy trees. Site 2 w as located along the Quebrada MÃ¡ quina off of the Se ndero Jilguero behind La EstaciÃ³n BiolÃ³ gica de Monteverde, Costa Rica. This site consisted mainly of riparian habitats with a low canopy and thic k understory. Inflorescence Phenology Study Fifty one inflorescences at Site 1 and 11 inflorescences at Site 2 of Calyptrogyne brachystachys were monitored throughout the three week study period at 2 to 4 day intervals. Only inflorescences that were pre f lowering were chosen for this study. Inflorescences were tagged and Vaseline was applied to the peduncle in order to exclude the introduction of any walking or crawling insects and/or colonization by crawling mites. The number of male and female flowers al ong with buds and location on the inflorescence were recorded during each date of collection. Local Site Inflorescence Stage Survey At each site a flowering stage survey was conducted to attain understanding of flowering stages of all inflorescences in th e area. On November 16, 2005, 84 inflorescences were surveyed at Site 2 and on November 19, 2005, 145 inflorescences were surveyed at Site
1. Inflorescences were categorized by the following criteria: Non reproductive state no inflorescence stalk present , unopened spathe fully enclosing inflorescence, budding spathe removed, but no flowers in bloom, flowering any number of flowers present, post flowering, fruiting and post fruiting. Multiple inflorescences belonging to the same plant were counted se parately. Analysis of Inflorescences for Mites Inflorescences monitored in the phenology study were collected when five to ten flowers remained or otherwise on the last day of study. Any additional inflorescences in flower, but not included in the phenolo gy census, were also collected. Inflorescences were examined under an Olympus SZ40 dissecting scope for mite presence. Individual flowers were carefully dissected and floral pits investigated along the entire length of the inflorescence with a dissecting p robe and knife. Bat Phenology Data for bat phenology were taken and graphed from Dien e rstein 1983. RESULTS Mite Occurrence Mites did not colonize the majority of the inflorescences examined. Eleven out of 46 inflorescences 23.9% were observed to have the presence of mites Fig. 1. A total of 36 mites were found on these 11 mite positive inflorescences, with one inflorescence housing over 20 mites. This yielded an average of 3.27 +/ 5.59 mites per mite positive inflorescence and an overall average of 0.78 +/ 2.99 mites Table 1. While most mite positive inflorescences had few mites, occasional inflorescences had many mites Fig. 3 histogram. In respect to seasonality, more mites were found in inflorescences collected later in the study period F ig. 2. Table 1. Inflorescence analysis of C. brachystachys for flower mites. n = 46. Total mite abundance 36 Inflorescences containing mites 11 Percent of inflorescences containing mites 23.9% Mean number of mites per inflorescence 0.78 Â€ 2.99 Mean n umber mites per mite positive inflorescence 3.27 Â€ 5.59 Five of the 11 inflorescences found to harbor mites also exhibited chew marks, possibly evidence of bat visitation. In addition, six of the eleven inflorescences found to be colonized by mites had Vaseline applied to the peduncle prior to inflorescence opening to exclude crawling mites. This indicates that the colonizing mites arrived by air or a phoretic carrier. Insect larvae Arthropoda: Insecta were found in 28% of the inflorescences surveyed w ith up to 5 larvae in one inflorescence. The presence of weevils Insecta:
Scaribidae feeding on flowers and surface worms Annelida: Oligochaeta feeding in floral pits were also observed. 0 10 20 30 40 50 60 5-Nov 8-Nov 10-Nov 16-Nov 19-Nov Date of Collection Percent of Mite-Positive Inflorescences FIGURE 1. Percentage of mite positiv e inflorescences for each day of collection. 0 5 10 15 20 25 30 35 40 0 1 2 3 4 5 6 7 8 9 10 > 10 Mites on one Inflorescence Number of Inflorescences FIGURE 2. Histogram of frequency of bat mites on each individual inflorescence.
Plant Phenology Phenology studies at both sites showed variation in frequency of C. brachystachyss inf lorescences and flowers present each day. Trends in flower phenologies at site 1 are depicted in Figure 1. Over the two week sampling period at site 1, the number of open inflorescences increases while the average number of flowers per inflorescence decrea ses. Site 2 showed an increase in frequency of open inflorescences and a corresponding increase in mean flower frequency per inflorescence Fig. 2. 0.00 10.00 20.00 30.00 40.00 50.00 60.00 70.00 80.00 90.00 100.00 5-Nov 8-Nov 10-Nov 14-Nov 19-Nov Date of Sample 2005 Percent of Individual Inflorescences -5.00 0.00 5.00 10.00 15.00 20.00 25.00 30.00 Flowers per Inflorescence Open Unopened Average Number of Flowers per Inflorescence FIGURE 3. Phenology study at site 1 n sample dates = 5, i inflorescences = 35 to 49. Frequency of open inflorescences increases slowly over a two week period with the reciprocal decrease of unopened inflorescences. Interestingly the trend of average flowers per open inflorescence does not follow the trend of open inflorescences.
0.00 10.00 20.00 30.00 40.00 50.00 60.00 70.00 80.00 9-Nov 16-Nov Date of Sample 2005 Percent of Individual Inflorescences -10.00 -5.00 0.00 5.00 10.00 15.00 20.00 Flowers per Inflorescence Open Unopened Average Number of Flowers per Inflorescence FIGURE 4. Phenology study at site 2 n sample dates = 2, n inflorescences = 13. Frequency of open inflorescences increases over a one week period with the reciprocal decrease of unopened inflorescences. Here the trend of average flowe rs per open inflorescence follows the trend of open inflorescences. Local site survey of reproductive stages in C. brachystachys presented disproportionate frequencies in plant stages. Both sites were found to have low frequencies of inflorescences in bud ding and flowering stages in addition to individual fruiting or post fruiting Figs. 3 and 4. Many individuals were found to have unopened inflorescences, suggesting a high frequency of flowering individuals within the population in the near future.
0 10 20 30 40 50 60 70 Non-Reproductive Unopened Budding Flowering Post-Flowering Fruiting Post-Fruiting Developmental Stage Frequency FIGURE 4. Site 1, Bullpen fragment inflorescence stage survey n = 145. Markedly more plants are in non reproductive stages or have unopened inflorescence than any other, with inflorescences past the fruiting stage next common in fr equency. 0 5 10 15 20 25 30 35 Non-Reproductive Unopened Budding Flowering Post-Flowering Fruiting Post-Fruiting Developmental Stage Frequency FIGURE 5. Site 2, behind La EstaciÃ³n BiolÃ³ gica de Monteverde n = 84. Similar to site 1 Fig. 2 an d disproportional amount of plants have inflorescences that are unopened, post fruiting or have no inflorescence stock a t all.
Bat Phenology Data taken from Dinerstein 1983 suggests seasonal phenology in Artibeus toltecus activity in the Monteverde Cloud Forest Fig. 5. Bat activity in the area during the months of November and December are notably low. 0 20 40 60 80 100 120 140 160 M J J A S O N D J F M A Month Artibeus toltechus Capture FIGURE 5. Phenology of Artibeus toltecus presence in Monteverde Cloud Forest. April 1981 to May 1982. Data from Dinerstein 1983. Weather Local weather in the Monteverde Cloud Forest for the fall of 2005, occurring over the duration in which t his study took place was exemplified by high winds, rain and mist Fig. 6. The mean rainfall for the month of October taken between 1952 and 1992 was 425 mm Haber, 2000, while October 2005, the month preceding this study had a mean rainfall of 908 mm, i ncluding 22 days with greater than 10 mm of rainfall a day and 17 days with more than 20 mm of rainfall a day. In addition to these data, personal observations document high occurrence of mist and gusts in the evenings of November 2005.
0 10 20 30 40 50 60 70 80 10/1/05 10/8/05 10/15/05 10/22/05 10/29/05 11/5/05 11/12/05 11/19/05 Date Rainfall mm cloudwater/mist rain FIGURE 6. Weather for Monteverde, Costa Rica. October 1 st through November 19 th , 2005 DISCUSSION Phoretic flower mites were found on Calyptrogyne brachystachys inflorescences in Monteverde that had Vaseline applied to the peduncle, confirming arr ival by air. Mites were found in lower frequencies and abundance than those found by Tschapka and Cunningham 2004 on La Selva Calyptrogyne ghiesbreghtiana inflorescences. The difference in mite populations between these two sites may be due to many reaso ns. Abiotic differences in sites such as altitude, seasonality and weather as well as biotic interactions between species may be responsible for differences in occurrence of this interaction mite, bat and plant. La Selva Biological Station is located on t he Atlantic slope, encompassing Holdridge Life zones of Tropical Wet Forest and Tropical Premontane Wet Forests, with an annual rainfall of 3,991 +/ 728 mm n = 22 years and ranging over elevations of 35 to 150 m Janzen 1983. Although La Selva experien ces a distinct dry season, every month receives a minimum of 150 mm of rainfall Lesica and Antibus 1990. In fact rainfall at La Selva, as in much of the Atlantic slope forests, is so well distributed throughout the year that it is considered to have an a seasonal climate Haber 2000. This study was conducted in Cloud Forest on the Pacific slope of Costa Rica. The Monteverde Cloud Forest Preserve is found in a lower montane rainforest that receives an average of 3,000 mm of precipitation per year Lesica and Antibus 1990. Site 1 in this study was located at 1520 m of elevation. Similarly, the EstaciÃ³n BiolÃ³ gica de Monteverde and site 2 are located at a higher elevation of 1600 m, and found in a lower montane wet forest life zone. Although clouds and mist are frequent throughout the dry
season, Pacific slope forests express higher seasonality than Atlantic forests Haber 2000. Elevational differences between sites is unlikely to be important to mite colonization considering hummingbird flower mites are fou nd in normal communities and frequencies in both Monteverde and Cerro de la Muerte Colwell 1983. The major differences between the study conducted by Tschapka and Cunningham 2001 and this study is the location of collection sites in relation to the co ntinental divide and seasonality differences that go along with this. In Monteverde minimal flowering activity occurs during the heaviest months of rainfall, September and October as well as during the windy misty season of November and December Haber et al. 1995. Inflorescence stage surveys in the month of November indicate that at the time of mite census gathering both survey sites experienced a local minimum in budding and flowering inflorescences Figures 3 and 4. Plant stage surveys over both sites indicate that most plants were not in flowering stage Figures 1 and 2 and were in fact not producing inflorescence stalks at all, or that inflorescences were still covered by bracts. The unavailability of flowers may impose a restriction on both coloniz ation and growth rate of mites. In addition, mature flowers may not occur at great enough abundances to cause fruit and/or nectar bats to devote foraging behavior to C. brachystachys. Koptur et al 1988 did a survey of seasonal penology of C. brachystach ys in the Monteverde, finding continuous leafing, different annual periodicity, unsynchronized individuals within species. In addition to peak and lag periods present in C. brachystachys phenology , Artibeus bat activity in the Monteverde region is also very seasonal. Absence of a phoretic carrier would prevent mites from accessing resources and therefore colonize. Heavy precipitation in both rain and mist are part of the local weather here in Monteverde and may affect local pollination and pollinator ac tivity. Two different populations of the mite, Proctolaelaps lobata, phoretic on Lepidoptera Insecta and specific to Lantana spp. Verbenaceae flowers were found by Boggs and Gilbert 1987 to have remarkably different annual population dynamics in response to differences in geographic seasonality. Lantana spp. are distributed from Central America to Texas and the mites are assumed to follow an analogous range. Populations of P. lobata were studied in Santa Rosa, Costa Rica, where Lantana spp. flower continuously and in Austin, Texas, USA where Lantana spp. are dormant during the winter. Due to monophagy of P. lobata , an annual local extinction occurs each winter in Texas. Although Lantana begins to flower in Texas in April, mites are not observed unt il June. Soon after, density and distribution grew quickly to equilibrium within a week Boggs and Gilbert 1987. This time of year coincides with both the year s first annual Southerly winds and the import of commercially cultivated Lantana flowers. Boggs and Gilbert 1987 concluded that populations in Austin are a result of annual re colonization of local phoresy on wide ranging insects, but could not rule out anthropogenic import. While this bat flower mite commensalisms relationship has been found to be widespread, more seasonal habitats are more difficult for steady populations to exist. Furthermore, seasonal populations may experience extreme minima due to seasonality of both host plant and phoretic carrier. A similar pattern to Lantana mites in Tex as of complete re colonization following temporary extinction each year is likely to explain
why densities and frequencies of phoretic bat mites were found to be so low on C. brachystachys in Monteverde. Far ranging Artibeus bats may annually introduce mit es from the nearby Atlantic slope. This would explain the presence of mites, however in very low numbers. Further studies concerning flowering phenologies need to be conducted over longer time periods. Foraging behavior of Artibeus bats and any differences of these between Pacific and Atlantic slope sites should also be explored. FIGURE 5. Phoretic mite on Calyptrogyne brachystachys inflorescence.
AC NOWLEDGEMENTS This study was made possible by my Grandfather, Don Smith; my parents and the Council for International Exchange Education. I thank the Vargas family, Richard LaVal, Alan and Karen Masters, Javier Mendez, Maria Jost and Oliver Hyman for all their help and guidance throughout this study. I am especially grateful for the time and energy expended by Maria and Ollie during attempted bat mist net sessions. Richard was also very helpful in suggestion and advice on collection sites and methods. I also would like to thank all my fellow classmates for encouragement, technical assistance and comradely . L ITERATURE CITED B OGGS , C ARROL AND L AWRENCE E. G ILBERT . 1987. Spatial and Temporal Distribution of Lantana Mites Phoretic on Butterflies. Biotropica 19: 301 305. C OLWELL , R OBERT K. AND S HAHID N AEEM . 1993. Life History Patterns of Hummingbird Flower Mites in Relation to Host Phenology and Morphology. In Houck, Marilyn A. Mites: Ecological and Evolutionary Analyses of Life History Patterns, pp 23 44. C OLWELL , R. K. 1983. Rhinoseius colwelli. In Janzen, Daniel H. 1983. Costa Rican Natural History, pp. 767 76 8. The University of Chicago Press. D INERSTEIN , E RIC . 1983. Reproductive Ecology of Fruit bats and Seasonality of Fruit Production in a Costa Rican Cloud Forest. University of Washington H ABER , W ILLIAM , W ILLOW Z UCHOWSKI AND E RIC B ELLO . An Introduction to the Trees of Monteverde. 1995. San Jose, Costa Rica. H ABER , W.A. 2000. Description of Permanent Forest Plots in Monteverde p. 90 Plants and Vegetation p. 41 Vascular plants of Monteverde p.503. In Nadkarni, Nalini M. and Nathaniel T. Wheelwright. 2000.Monteverde: Ecology and Conservation of a tropical Cloud Forest. H ENDERSON , A NDREW , G LORIA G ALEANO AND R ODRIGO B ERNAL . 1995. Field Guide to Palms of the Americas. Princeton University Press. H YNEMAN , A MY J O , R OBERT K. C OLWELL , S HAHID N AEEM , D AVID S. D OBKIN AND B ERNARD H ALLET . 1991. Host Plant Discrimination: Experiments with Hummingbird Flower Mites. In Price, Peter W., G. Wilson Fernandes, Thomas M. Lewinsohn and Woodruff W. Benson. Plant Animal Interactions, pp. 455 485. John Wiley & Sons, Inc. J ANZEN , D ANIEL H. 1983. Costa Rican Natural History. The University of Chicago Press. K OPTUR ET . AL . 1988. Phenological studies of shrub and treelet species in tropical cloud forests of Costa Rica. Department of Botany and Department of Entomology, Unive rsity of California, Berkeley. L ESICA , P ETER AND R OBERT K. A NTIBUS . The Occurrence of Vascular Epiphytes of Two Costa Rican Rainforests Mycorrhizae. Biotropica 223: 250 258 1990 T IMM , R OBERT M. AND R ICHARD K. L A V A l. 2005. Mammals of Monteverde. In Nadk arni, Nalini M. and Nathaniel T. Wheelwright. 2000.Monteverde: Ecology and Conservation of a tropical Cloud Forest. pp: 90 and 503. T SCHAPKA , M ARCO AND S AULA A. C UNNINGHAM . 2004. Flower Mites of Calyptrogyne ghiesbreghtiana Arecaceae: Evidence for Dispe rsal Using Pollinating Bats. Biotropica 36: 377 381.
W HEELWRIGHT , N ATHANIEL T. 2000. A Hypothesis About the timing of Flowering and Fruiting in Competing tropical Trees. In Nadkarni, Nalini M. and Nathaniel T. Wheelwright. 2000.Monteverde: Ecology and C onservation of a tropical Cloud Forest. pp: 281 283.