Epiphyte Communities in Cupressus lusitanica Windbreaks and Myrsine coriacea , Monteverde, Costa Rica Casey Halstead University of Colorado at Boulder in conjunction with CIEE Monteverde ABSTRACT Monteverde, Costa Rica is home to a quintessential lower montane cloud forest that supports an amazing diversity of epiphytes. Windbreaks comprised of the exotic Cupressus lusitanica are common surrounding deforested areas. C. lusitanica appears to be poor epiphyte habitat when compared to a native species Myrs ine coriacea. This survey was conducted using species richness and occurrence data collected using transparent acetates. This survey examines the idea that C. lusitanica supports reduced epiphyte diversity. The survey determined that the apparent differenc e in epiphyte richness and community composition is significant. ANCOVA analysis revealed several negative correlations for M. coriacea but not C. lusitanica. Lichen and moss abundance were affected by each other s abundance as well as by CBH. -------------------------------------------------------------------------------------------------------------------------------RESUMEN Monteverde, Costa Rica alberga un bosque montane bajo n u boso que sostiene una diversidad asombrosa de epifitas . Los rompe viento s abarcados por la exÃ³tica Cupressus lusitanica son comunes en Ã¡reas deforestadas de los alrededores. Cupressus lusitanica aparece ser un hÃ¡bitat pobre en epifitas cuando se compara con la especie nativa Myrsine coriaceae . Este estudio fue conducido usando la riqueza de especies y los datos de la ocurrencia recogidos usando los acetatos transparentes. Este estudio examina ademÃ¡s la idea de que C. lusitanica disminuye la diversidad de epifitas . El estudio determinÃ³ que la diferencia evidente en riqueza de epifitas y la composiciÃ³n de la comunidad es significativa. El anÃ¡lisis de ANCOVA revelÃ³ varias correlaciones negativas para M. coriaceae pero no en C. lusitanica . La abundancia de lÃquenes y musgos se vio afectada por cada otras abundancia asÃ como por CB H. INTRODUCTION The lower montane cloud forest of Monteverde, Costa Rica may support the most diverse epiphyte communities in the world with approximately 250 vascular Ingram 1996 and 190 bryophyte Gradstein 2000 species identified from a single 4 he ctare plot. However, epiphyte diversity in the area s second growth forests, including native and planted non native trees, appears to be greatly reduced. Deforestation in Monteverde facilitated the negative effects of dry season winds by promoting soil l oss that reduced
pasture grass production thereby reducing milk yields Burlingame 2000. In the 1980 s local farmers were encouraged to plant windbreaks in order to protect soil and grasses, shield cattle, and restore forest coverage Burlingame 2000. Th is effort resulted in over 500,000 trees being planted Burlingame 2000 in approximately 1000 windbreaks that stretch a combined distance of 185 km Harvey 2000. A handful of studies have examined the ecology of windbreaks of the Monteverde area Harvey 1999, Harvey and Haber 1999, Harvey 2000a, b, however, no study has been conducted on a species specific basis or focused on epiphytic communities. Cupressus lusitanica Cupressaceae is among the most commonly used windbreak species Harvey 2000a, Harve y 2000b, Nielsen and DeRosier 2000, Burlingame 2000 and appear to be poor habitat for epiphytes personal observations . Cupressus lusitanica is an exotic tree used because of a fast growth rate, ease of cultivation on cleared land and extensive low branc hing Burlingame 2000, Zuchowski 2005. Myrsine coriacea is a native tree with similar growth patterns Haber et al. 2000 and, unlike C. lusitanica, appears to be good habitat for epiphytes personal observations . Myrsine coriacea is not planted in wind breaks although the common name tapa viento , Spanish for blocks wind, implies it has potential . If M. coriacea is better habitat for epiphytes it may provide a native alternative to C. lusitanica. This study compares epiphyte communities between the two species in an attempt to determine whether the apparent reduction in epiphyte richness is significant. Since epiphytes are a dominant component of cloud forest ecosystems, a study of this nature will increase understanding of the role C. lusitanica windbre aks play in the ecosystem and help establish how effective this species is as a reforestation tool. In addition studies
investigating why the richness is reduced cannot be undertaken until the difference is established to be significant. A variety of bio logical phenomena support the idea that epiphyte richness and occurrence should be the same between species. For instance, epiphytes are not dependent on their hosts for nutrients. In addition, epiphyte communities become more complex with age and the spec imens analyzed are of equal age potentially younger in the case of M. coriacea . Other phenomena support the idea C. lusitanica should have increased richness. For instance, Island Biogeography Theory IBT predicts that targets of an equal distance fro m propagule sources and equal size and equal age should harbor equal richness and abundance. The average size of C. lusitanica in the study area is nearly double that of M. coriacea . Therefore, IBT predicts increased richness and abundance on C. lusitanica . Personal observations as well as biological phenomena contradict the idea that richness and abundance is increased. Cupressus lusitanica has very fast growth rates and studies conducted by Gentry and Vasquez 1993 showed that epiphyte diversity is redu ced on species with rapid growth rates. In addition C. lusitanica lacks resources that might attract disperser further supporting the idea that C. lusitanica would have reduced richness and abundance. Finally, conifers have been known to alter their enviro nmental chemistry which may prevent epiphyte growth. Since biological phenomena exist to explain equal, reduced, or increased richness and abundance I conducted a survey to determine if the observed richness and abundance are reduced on C. lusitanica.
MET HODS This survey was conducted during the rainy season in a second growth lower montane forest adjacent to the EstaciÃ³n BiolÃ³gica, Monteverde, Costa Rica GPS. The site was chosen because succession on this property has progressed for a known amount of time allowing for assumptions to be made about tree ages. The C. lusitanica examined were planted in 1987; the M. coriacea examined grew as a result of natural succession that began in 1987. This led to the assumption that all M. coriacea were of the same age, or younger than the C. lusitanica . The area was paralleled, on one side, by intervened but intact forest. The remainder of the site surrounding consisted of second growth forest, a road and several residential establishments. This study analyzed ep iphyte communities on 36 individual trees 18 per species. Richness estimations were conducted on a morphospecies basis. Total richness was determined by visual analysis of tree boles in an area between 0 m and 3 m from the tree base. Occurrence data were collected using transparent acetates with 304, 1 cm 2 , cells and recorded on a presence or absence basis. For this presence or absence data was collected for lichen, moss, ferns, orchids and non orchid angiosperms. Bryophytes were considered present if the y occupied Â€ 50% of the area of a cell; vascular plants were considered present if found regardless of the percent of the cell they occupied. Acetates were pinned to the east side of trees at 30 cm and 80 cm above ground. Occurrence data was pooled. Tree size measure ments were taken using the circumference at breast height cbh method. In this study ÂBHÂ‚ is defined as eye level for an individual who 1.75 m tall. In the event that a tree forked before this height the larger of the two forks was measured. The data were analyzed using ANCOVA, linear regression, and T test.
RESULTS A total of 26 morphospecies were identified and consisted of seven orchids, six lichen, five mosses, five ferns and three non orchid angiosperms. However, in occurrence surveys, non orchid ang iosperms were absent and orchids were least abundant n = 47, followed by ferns n = 186. Lichens n = 4078 and mosses n = 13,329 were most abundant. Richness, total epiphyte abundance, and lichen abundance were found to be significantly higher on M. coriacea while moss abundance was found to be significantly greater on C. lusitanica Fig. 1 . ANCOVA revealed no significant relationship between variables for C. lusitanica Table 1. ANCOVA revealed several significant relationships for M. coriacea Tab le 2. All relationships reported by ANCOVA were negative Fig. 2. DISCUSSION The results support the impression that C. lusitanica harbors reduced epiphyte richness and abundance. A negative correlation goes against predictions that arise from IBT. Since hundreds of studies support IBT, I argue this is a result of a unique characteristic of C. lusitanica. The idea that C. lusitanica has unique characteristic is further supported by ANCOVA results since the correlations between moss, lichen, DBH, and richness that were found on M. coriacea were not found on C. lusitanica. Now that the apparent reduction in epiphyte growth has been established future studies can investigate ecological explanations for this result. This may be due to conifers ability to alter the surrounding chemistry. In addition I observed bark peeling off of C. lusitanica. A reduction of animal traffic in these areas may also be responsible for this
reduction. It is for these reasons I recommend future studies focus on bark characte ristics such as chemistry and adhesion as well as the presence of dispersers. Studies of Monteverde windbreaks have shown that they are a powerful reforestation tool by providing habitat for a variety of forest tree species, including those associated w ith primary growth forests Harvey 2000a. This is reportedly because of the shade they provide Harvey 2000a. This study has shown that M. coriacea supports greater biodiversity. For this reason I propose that it would be a better windbreak species. Beyo nd the increase in epiphyte richness M. coriacea has fruits that attract frugivorous birds Haber et al. 2000. Removal of C. lusitanica windbreaks might disturb characteristics that allow primary growth forest trees to grow. Therefore I argue that M. cori acea should be added to C. lusitanica windbreaks in order to increase the occurrence and variety of these species. Unfortunately, M. coriacea is a light loving species and current windbreak structure may prevent its proliferation. It is for this reason tha t I recommend future studies also investigate thinning effects on C. lusitanica windbreaks . If thinning of C. lusitanica windbreaks does not alter their ability to support primary growth trees and increases the ability to support M. coriacea then the comb ination of these two species may prove to be a more powerful reforestation tool. ACKNOWLEDGEMENTS I would like to thank CIEE for providing me with the opportunity to come to Costa Rica and conduct this study. This research would not have been possible wit hout economic assistance provided by the Ping and Stafford foundations. The EstaciÃ³n BiolÃ³gica provided housing as well as library and internet resources. Pablo Allen and JosÃ© Carlos Calderon provided much needed guidance during brainstorming and writing p rocess. Finally, I would like to thank Karen Masters for being a great inspiration as well as providing invaluable assistance in field identifications, experimental design and a suitable study site. LITERATURE CITED Burlingame, J. L. 2000. Conservation in the Monteverde area: Contributions of conservation organizations. In: Monteverde: Ecology and Conservation of a
Tropical Cloud Forest , Nadkarni, M. N., and N. T. Wheelwright, eds. Oxford University Press, New York, page 351 376 Gradstein, S. R., M. I. Morales, and N. M. Nadkarni. 2000. Diversity and habitat differentiation of mosses and liverworts in the cloud forests of Monteverde, Costa Rica. Caldasia 231: 203 212 Haber, A. W., W. Zuchowski , and E. Bello. 2000. An Introduction to Cloud Forest Tr ees of Monteverde , Costa Rica. Mountain Gem Publications, Monteverde, page 134 Harvey, A. C. 2000a. Colonization of agricultural windbreaks by forest trees: effects of connectivity and remnant trees. Ecol. Appl. 106: 1762 1773 Harvey, A. C. 2000b. Windb reaks enhance seed dispersal into agricultural landscapes in Monteverde, Costa Rica. Ecol. Appl. 101, 155 173 Nielsen, K., and D. DeRosier. 2000. Windbreaks as corridors for birds. In: Monteverde: Ecology and Conservation of a Tropical Cloud Forest , Nad karni, M. N., and N. T. Wheelwright, eds. Oxford University Press, New York, page 351 376 Ingram, W. S., K. F. Ingram, and N. M. Nadkarni. 1996. Floristic composition of vascular plants in a neotropical cloud forest, Monteverde, Costa Rica. Selbyana 17: 88 103 Zuchowski , W. 2005. A Guide to Tropical Plants of Costa Rica . A Zona Tropical Publications, San JosÃ© . page 223
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Comunidades de epfitas en el rompe vientos Cupressus lusitanica y Myrsine coriacea, Monteverde, Costa Rica
Epiphyte communities in Cupressus lusitanica windbreaks and Myrsine coriacea, Monteverde, Costa Rica
Monteverde, Costa Rica is home to a quintessential lower montane cloud forest that supports an amazing diversity of epiphytes. Windbreaks comprised of the exotic Cupressus lusitanica are common surrounding deforested areas. C. lusitanica appears to be poor epiphyte habitat when compared to a native species Myrsine coriacea. This survey was conducted using species richness and occurrence data collected using transparent acetates. This survey examines the idea that C. lusitanica supports reduced epiphyte diversity. The survey determined that the apparent difference in epiphyte richness and community composition is significant. ANCOVA analysis revealed several negative correlations for M. coriacea but not C. lusitanica. Lichen and moss abundance were affected by each others abundance as well as by CBH.
Monteverde, Costa Rica alberga un bosque nuboso montano bajo que sostiene una diversidad asombrosa de epfitas. Los rompe vientos abarcados por la extica Cupressus lusitanica son comunes en reas deforestadas de los alrededores. Cupressus lusitanica parece ser un hbitat pobre en epfitas cuando se compara con la especie nativa Myrsine coriacea. Este estudio fue conducido usando la riqueza de especies y los datos recogidos de la ocurrencia usando los acetatos transparentes. Este estudio examina adems la idea de que C. lusitanica disminuye la diversidad de epfitas.
Text in English.
Epiphytes--Costa Rica--Puntarenas--Monteverde Zone
Cupressus lusitanica--Costa Rica--Puntarenas--Monteverde Zone
Myrsine coriacea--Costa Rica--Puntarenas--Monteverde Zone
Windbreaks, shelterbelts, etc.
Cloud forest ecology--Costa Rica
Epfitas--Costa Rica--Puntarenas--Zona de Monteverde
Cupressus lusitanica--Costa Rica--Puntarenas--Zona de Monteverde
Rompe vientos, corta vientos, etc
Ecologa del bosque nuboso--Costa Rica
Tropical Ecology 2008
Ecologa Tropical 2008
t Monteverde Institute : Tropical Ecology