Structure of soil arthropod and plant communities in light gaps and continuous forest in Monteverde, Costa Rica


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Structure of soil arthropod and plant communities in light gaps and continuous forest in Monteverde, Costa Rica

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Title:
Structure of soil arthropod and plant communities in light gaps and continuous forest in Monteverde, Costa Rica
Translated Title:
Estructura de los artrópodos del suelo y las comunidades de plantas en los claros del bosque y un bosque continúo en Monteverde, Costa Rica
Creator:
Breza, Lauren
Publication Date:
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Text in English

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Subjects / Keywords:
Arthropods ( lcsh )
Artropodos ( lcsh )
Species diversity ( lcsh )
Diversidad de especies ( lcsh )
Cloud forest ecology ( lcsh )
Ecologia del bosque nuboso ( lcsh )
Monteverde Biological Station (Costa Rica)
Estacion Biologica de Monteverde (Costa Rica)
CIEE Fall 2008
CIEE otoño 2008
Genre:
Reports

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Abstract:
Several studies have confirmed that higher plant community richness facilitates greater arthropod community richness. The purpose of this study is to examine how plant community structure in continuous forest and light gaps impact soil arthropod community structure. In this study plant surveys were conducted for twenty plots, ten located in continuous forest and ten located within a forest gap. Along with plant surveys, a soil sample from each plot was collected in order to obtain soil dwelling arthropods. Arthropods were extracted from the soil first using a Burlese funnel, and then manually picked out of the soil. After all arthropods were obtained, they were identified to order and when possible to family. It was hypothesized that there would be a correlation between plant community structure and soil arthropod community structure, as well as a difference in diversity in continuous forest and forest gaps. The richness, evenness, and Smarg were calculated for plants in closed forest and in forest gap (H1' = 2.892, H2' = 2.406, t = 4.72, p < 0.0005). In addition, the richness, evenness and Smarg were calculated for soil arthropods in closed forest and in forest gap (H1' = 2.052, H2'= 1.896, t = 1.44, p = 0.10). There was no difference between soil arthropod diversity in continuous forest and forest gap and plant diversity did vary between continuous forest and forest gap. Regression analyses were also conducted, and the results suggest that there is no correlation between plant and soil arthropod community composition. However, there were obvious trends derived from the regressions that should be taken into consideration for future studies. ( ,, )
Abstract:
Algunos estudios han confirmado que una gran riqueza en las comunidades de plantas facilita una gran diversidad en la comunidad de artrópodos. El propósito de este estudio es examinar cómo la estructura de la comunidad de plantas en un bosque continuo y en los claros del bosque impacta a la comunidad de artrópodos del suelo. En este estudio se llevaron a cabo encuestas sobre plantas en 20 parcelas, diez ubicadas en el bosque continuo y diez ubicadas en los claros del bosque. Junto con las encuestas sobre plantas, también se llevaron a cabo encuestas sobre el suelo para obtener una muestra de los artrópodos que se encuentran en el mismo.
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Student affiliation : Department of Ecology and Evolutionary Biology, University of Tennessee
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Born Digital

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Monteverde Institute
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Monteverde Institute
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M39-00485 ( USFLDC DOI )
m39.485 ( USFLDC Handle )

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Estructura de los artrpodos del suelo y las comunidades de plantas en los claros del bosque y un bosque contino en Monteverde, Costa Rica
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Structure of soil arthropod and plant communities in light gaps and continuous forest in Monteverde, Costa Rica
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Several studies have confirmed that higher plant community richness facilitates greater arthropod community richness. The purpose of this study is to examine how plant community structure in continuous forest and light gaps impact soil arthropod community structure. In this study plant surveys were conducted for twenty plots, ten located in continuous forest and ten located within a forest gap. Along with plant surveys, a soil sample from each plot was collected in order to obtain soil dwelling arthropods. Arthropods were extracted from the soil first using a Burlese funnel, and then manually picked out of the soil. After all arthropods were obtained, they were identified to order and when possible to family. It was hypothesized that there would be a correlation between plant community structure and soil arthropod community structure, as well as a difference in diversity in continuous forest and forest gaps. The richness, evenness, and Smarg were calculated for plants in closed forest and in forest gap (H1' = 2.892, H2' = 2.406, t = 4.72, p < 0.0005). In addition, the richness, evenness and Smarg were calculated for soil arthropods in closed forest and in forest gap (H1' = 2.052, H2'= 1.896, t = 1.44, p = 0.10). There was no difference between soil arthropod diversity in continuous forest and forest gap and plant diversity did vary between continuous forest and forest gap. Regression analyses were also conducted, and the results suggest that there is no correlation between plant and soil arthropod community composition. However, there were obvious trends derived from the regressions that should be taken into consideration for future studies.
Algunos estudios han confirmado que una gran riqueza en las comunidades de plantas facilita una gran diversidad en la comunidad de artrpodos. El propsito de este estudio es examinar cmo la estructura de la comunidad de plantas en un bosque continuo y en los claros del bosque impacta a la comunidad de artrpodos del suelo. En este estudio se llevaron a cabo encuestas sobre plantas en 20 parcelas, diez ubicadas en el bosque continuo y diez ubicadas en los claros del bosque. Junto con las encuestas sobre plantas, tambin se llevaron a cabo encuestas sobre el suelo para obtener una muestra de los artrpodos que se encuentran en el mismo.
546
Text in English.
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Arthropod surveys--Costa Rica--Monteverde-- Biological Station
Species diversity--Costa Rica--Puntarenas--Monteverde Zone
Cloud forest ecology--Costa Rica
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Encuestas de Artrpodos--Costa Rica--Monteverde--Estacin biolgica
Diversidad de especies--Costa Rica--Puntarenas--Zona de Monteverde
Ecologa del bosque nuboso--Costa Rica
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Tropical Ecology 2008
Ecologa Tropical 2008
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Reports
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CIEE
773
t Monteverde Institute : Tropical Ecology
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u http://digital.lib.usf.edu/?m39.46



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Structure of soil arthropod and plant communities in light gaps and continuous forest in Monteverde, Costa Rica Lauren Breza Department of Ecology and Evolutionary Biology, University of Tennessee ABSTRACT Several studies have confirmed that higher pl ant community richness facilitates greater arthropod community richness. The purpose of this study is to examine how plant community structure in continuous forest and light gaps impact soil arthropod community structure. In this study plant surveys were conducted for twenty plots, ten located in continuous forest and ten located within a forest gap. Along with plant surveys, a soil sample from each plot was collected in order to obtain soil dwelling arthropods. Arthropods were extracted from the soil f irst using a Burlese funnel, and then manually picked out of the soil. After all arthropods were obtained, they were identified to order and when possible to family. It was hypothesized that there would be a correlation between plant community structure and soil arthropod community structure, as well as a difference in diversity in continuous forest and forest gaps. The richness, evenness, and Smarg were calculated for plants in closed forest and in forest gap H 1 ' = 2.892, H 2 ' = 2.406, t = 4.72, p < 0.0 005. In addition, the richness, evenness and Smarg were calculated for soil arthropods in closed forest and in forest gap H 1 ' = 2.052, H 2 ' = 1.896, t = 1.44, p = 0.10. There was no difference between soil arthropod diversity in continuous forest and fo rest gap and plant diversity did var y between continuous forest and forest gap. Regression analyses were also conducted, and the results suggest that there is no correlation between plant and soil arthropod community composition. However, there were obvi ous trends derived from the regressions that should be taken into consideration for future studies. RESUMEN Algunos estudios han confirmado que una gran riqueza en las comunidades de plantas facilita una gran diversidad en la comunidad de artrópodos. El propósito de este estudio es examinar como la estructura de la comunidad de plantas en un bosque continuo y en claros de bosque impacta la comunidad de artrópodos del suelo. En este estudio se muestrearon plantas en 20 parcelas, diez ubicadas en el bosqu e y diez ubicadas en claros de bosque. A lo largo de las plantas se tomaron muestras de suelo para obtener una muestra de los artrópodos que se encuentran en el mismo. Artrópodos fueron extraídos del suelo utilizando embudos Burlese, y luego extraídos ma nualmente del sustrato. Después de que se obtuvieron todos los artrópodos, se identificaron a nivel de orden o familia en los que fue posible. La hipótesis del estudio es que existe una correlación entre la estructura de la comunidad de plantas y la est ructura de la comunidad de artrópodos, así como la diferencia en diversidad entre el bosque continuo y claros de bosque. La riqueza, equidad y Smarg fueron calculados para bosque cerrado y claros de bosque H 1 ' = 2.892, H 2 ' = 2.406, t = 4.72, p < 0.0005. Además se calculó lo mismo para los artrópodos del sustrato en bosque cerrado y claros de bosque H 1 ' = 2.052, H 2 '= 1.896, t = 1.44, p = 0.10. No hay diferencia entre la diversidad de artrópodos del suelo entre el bosque cerrado y los claros de bosque, mientras que la diversidad de plantas si varía entre estos. No hay correlación en cuanto a la composición de la comunidad de plantas y artrópodos; sin embargo, existen tendencias obvias que pueden ser tomadas en consideración para futuros estudios.

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IN TRODUCTION Knowing and understating the relationships that soil dwelling arthropods have within a community is essential to understanding the processes of any food web. A majority of soil arthropods usually fall into the category of detritivores, organis ms that play a crucial role in the decomposition of detritus, however other trophic levels are found in the soil as well predators, herbivores, and parasites Siemann 1998. Detritus is made up of dead organic material, which generally includes dead fal len leaves and the dead branches, stems, and roots of plants which can be found above or below ground Whittaker 1975. With out these organisms, dead organic matter would continue to build up into unfathomable amounts. By taking part in the decompositio n processes, along with microbes and carrion feeders, detritivores are completing the necessary sequence of energy flow through a food web. It has been found through several studies that energy flow through ecosystems is controlled by a combination of top down and bottom up interactions Hättenschwiler 2005, Siemann et al. 1999. Top down interactions are facilitated by the top trophic level, where as bottom up interactions are initiated by plants, the primary producers. Both processes are a crucial part of any ecosystem, but each ecosystem is highly variable in which process, top down or bottom up, is the dominating force. In tropical premontane forests the soil is mostly comprised of nutrient poor clay and sand. All of the nutrients found in tropical rainforests reside within the canopy and only a thin layer of rich decaying organic matter is found on the forest floor. Depending on the type of forest, whether it is pristine continuous forest or secondary growth facilitated by a tree fall gap, the type s of succession found in a tropical forest will determine the types of flora and fauna species found there. Succession is the process by which one type of forest age gives way to the next stage until a stable climax is reached. The modern view of success ion says that only by bottom up interactions drive succession, however studies have shown that ecological succession is driven by both bottom up and top down interactions Schmitz 2006. Within the successsional process of forest regeneration it is import ant to know what kind of impact both top down and bottom up interactions have on the species richness. In general, it has been established that fauna diversity will increase with plant diversity Lagerlöf et. al. 1993, Siemann 1998 Wardel et. al. 1999. Previous studies have also found that detritivore communities are positively correlated with an increase in plant species in old fields Siemann et al. 1999. Though this is significant for old fields, it leaves plausible opportunities to investigate whet her or not detritivore diversity is positively correlated with an increase in plant species in other ecosystems. This same study also acknowledges that arthropod diversity herbivores, predators, parasites, and detritivores increases in diversity with an increase in succession. The purpose of this study is to investigate whether or not plant species richness and abundance, in both continuous forest systems and in naturally disturbed areas like forest gaps, impact the soil arthropod community composition and abundance. It was hypothesized that there would be a correlation between soil dwelling arthropods and plant community structure. I predicted that soil dwelling arthropods would be positively correlated in species richness and abundance with plant spe cies richness and abundance. It was also hypothesized that overall species richness of plants and soil arthropods would

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be different in different types of forest growth. I predicted that there would be the greatest diversity in pristine closed forest and less diverse in naturally disturbed forest. METHODS This study was conducted in the premontane wet forest of la Estación Biológi c a de Monteverde, Puntarenas, Costa Rica . Four 30 m transects were marked, two within continuous forest and two in a naturall y disturbed area light gap. On each transect five randomly selected 1 m x 1 m quadrates were placed. In each quadrate a plant survey was conducted by counting the abundance of each species present within the 1 m x 1 m quadrat e . Any unidentifiable plan t species was collected. In addition to plant surveys, 0.5 L of soil was collected from each quadrat e to determine the type and abundance of soil dwelling arthropods. At each quadrat e , the Braun Blanquet cover classes were used to estimate the canopy cov erage 1 = 1% € 5%; 2 = 6% € 25%;. 3 = 26% € 50%; 4 = 51% € 75%; and 5 = > 75% . The collected soil was taken back to the biological station and placed in Burlese funnels for a maximum of two days to collect soil dwelling arthropods. The soil was then hand proce ssed for 30 minutes per soil sample. The arthropods were stored in containers with 95% ethanol and were identified to order, if possible family. Collected uniden ti fiable plant species were taken back to the biological station, pressed in a plant press, and then identifie d . Statistical Analysis The Shannon Weiner index for diversity was calculated along with evenness and Margalef's index in order to compare plant diversity in a closed forest system and an open forest system. These same statistics were con ducted for soil arthropod diversity in both the closed forest system and an open forest system. R egression analyses were conducted to examine if there were any relations between the following: the number of arthropod taxa and the number of plant taxa; the arthropod abundance and the number of plant taxa; the number of arthropod taxa and plant abundance , and finally the arthropod ta x a and plant abundance. Then Chi squared tests were performed to d etermine if there was any difference between sites in arthro pod abundanc e, number of arthropod taxa , plant abundance and the number of plant taxa and finally the degree of canopy coverage. RESULTS In this study it was observed that the largest taxon of soil dwelling arthropods were coleopterans beetles and memb ers of the family Formicidae ants. The large numbers in Coleopteran abundance is not abnormal because Coleoptera is the largest order of insects. The large numbers of individuals from the family Formicidae are not representative of the abundance in soi l habitats; these numbers are derived from only two plots out of 20 . In the continuous forest the plants with the highest abundance was Icanthaceae I, Monstera adensoni , and Begonia involucrata . In the forest gap the plants that dominated in abundance we re Unknown 32, Pouzolia perasitica , and Viola

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stipularis . In both continuous forest and forest gaps Begonia involucrata was extremely common; this plant is a succulent understory plant that has a wide distribution and is found anywhere in primary and seco ndary forest, forest edges, and along side trails Zuchowski. Because of B. involucrata's ability to survive and persist in various habitats, it is not unusual that this was one of the most common plants in both forest systems. The results of the Shannon Wiener test showed that the closed forest H' = 2.892 is more diverse in plants than open naturally disturbed forest H' = 2.406 t = 4.72, p < 0005 Appendix, Table 2. However, there were no differences indicating that soil arthropod communities ar e more diverse in closed forest H' = 2.085 or in naturally disturbed forest H' = 1.945 t = 1.22, p = 0.10 Appendix, Table 1. The regression analyses showed that there were no statistically significant relationships between plant diversity and abund ance and soil arthropod diversity and abundance Fig. 1. In spite of this, there are obvious trends that suggest that there could be a significant correlation with further investigation. A 0 1 2 3 4 5 6 7 8 9 0 5 10 15 20 Number of Plant Taxa Number of Arthropod Taxa

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B C C 0 10 20 30 40 50 60 0 10 20 30 40 50 60 Plant Abundance Arthropod Abundance 0 1 2 3 4 5 6 7 8 9 0 10 20 30 40 50 60 Plant Abundance Number of Arthropod Taxa

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D FIGURE 1. Closed circles represent plots found in closed forest and open circles represent plots found in naturally disturbed areas. The solid lines are the regression lines for closed circles and the dashed lines are the re gressions for open circles. a A strong trend between the number of plant taxa and the number of arthropod taxa in both closed and open forest R2 = 0.029, F = 0.55, df = 1, 18, p = 0.46. b When arthropod abundance was correlated with plant abundance the re was not a significantly strong correlation to suggest any pattern R2 = 0.074, F = 0.134, df = 1, 18, p = 0.7185. c Shows a correlation between the number of arthropod taxa and plant abundance, but the correlation is not strong enough to pursue furthe r investigation R2 = 0.049, 0.925, df = 1, 18, p = 0.349. d a visible correlation within an open forest system, but not in the closed forest system R2 = 0.077, f = 1.496, df = 1, 18, p = 0.2370. The Chi squared analyses showed that arthropod abundan ce ðc 2 = 53.8, p < 0.0005 and plant abundance ðc 2 = 71.9, p < 0.0005 are impacted by the amount of canopy coverage. The Chi squared test also determined that canopy coverage does not affect the number of arthropod taxa ðc 2 = 5.07, p = 0.27 and the nu mber of plant taxa in a premontane wet forest ðc 2 = 2.57, p = 0.63. 0 10 20 30 40 50 60 0 5 10 15 20 Number of Plant Taxa Arthropod Abundance

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A 0 10 20 30 40 50 60 1 2 3 4 5 Canopy Coverage Class Number of Plant taxa B 0 5 10 15 20 25 30 35 1 2 3 4 5 Canopy Coverage Class Number of Arthropod Taxa

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C D 0 50 100 150 200 250 300 1 2 3 4 5 Canopy Coverage Class Plant Abundnace Number of Individuals 0 20 40 60 80 100 120 1 2 3 4 5 Canopy Coverage Class Arthropod Abundance Number of Individuals

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FIGURE 2. Frequency distributions of A number of plant taxa and amount of canopy coverage, B number o f arthropod taxa and amount of canopy coverage, C plant abundance and the amount of canopy coverage D and arthropod abundance and canopy coverage. DISCUSSION It is not surprising that continuous forest has a higher diversity of plant species than a naturally disturbed forest area; it is a widely accepted idea that forest age is an indicator of the amount of flora diversity Siemann et al. 1999. In this study it was determined that there was a higher amount of diversity found in the continuous for est system than in the younger forest gap. However, it is interesting to note that there was not a different in the diversity between soil arthropods in continuous forest and naturally disturbed areas. Several succession studies have shown that arthropod diversity in general increases with forest age. These studies included herbivores, predators, parasites, and detritivores. Even thought the overall arthropod diversity changes, it has been found that the largest of arthropods that change are herbivores that colonize individual plants rather in the soil. Detritivores in previous studies are shown to not change in diversity or relative abundance Schmitz 2006, Seimann et al. 1999 One aspect that is critical for future studies examining soil arthropod di versity in light gaps, it is important to take note of which pioneer plant species use chemical defenses to deter soil dwelling herbivores. In the premontane wet forest of Monteverde there are several species of plants that use alkaloids and other toxins to make themselves less desirable to herbivores, of these plants the species Bocconia frutescens Papaveraceae has been shown to have chemicals that are the most effective at deterring soil arthropods, specifically brine shrimp Veldman et al. 2007. Cano py coverage is factor that plays an important role in limiting resources such as light, moisture, and subcanopy temperatures. The amount of coverage impacts plant and soil arthropod abundance. It is not surprising that with greater canopy coverage, the a bundance of plants decreases Figure 2c, p < 0.0005; resources are limited and the plants with the greatest advantage will succeed in growth. However, in the frequency distribution of soil arthropod abundance and canopy coverage, it appears that there is an optimal amount of canopy coverage, which peaks at class 3 26 50% of coverage Figure 2d, p < 0.0005. This could be due to bottom up interactions; bottom up limitations, in this case canopy coverage, are the determinant factor in the amount of leaf litter that will make its way to the forest floor. Canopy coverage is a resulting variable of how productive the canopy elements are, and up to 90% of the aboveground net primary productivity soon to become leaf litter enters the detritus food web in fo rest ecosystems Chen and Wise 1999. Depending on what the optimal conditions are for soil arthropods thrive, it is possible that a certain level of light is needed as well as a specific amount of detritus. A study conducted by Siemann 1999 documented that increased plant productivity increases the total arthropod abundance and species richness, as well as species richness and abundance of each trophic group within an old field ecosystem. Detritivores, herbivores, parasites, and predators were shown to have been greater in

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plots with higher levels of modern methods of fertilization. This study suggests that the diversity and composition of plants control the diversity of consumers through both bottom up and top down interactions. However, in this stud y the data indicates that there is no correlation between plant abundance and richness to soil arthropod abundance and richness. Overall the hypotheses were not supported by the data; the prediction that plant diversity is greater in continuous forest than in gap forest was supported. However, the results did not support the prediction that soil arthropod diversity would be greater in continuous forest rather than in gap forest. Though many studies have concluded that detritivore diversity and abundance a re not correlated with plant richness and abundance, they have been shown to increase in diversity with increased plant productivity. A second study examining this trend especially in a tropical forest would provide more insight into the plant detritivo re relationship, which is usually overlooked in many cases. ACKNOWLEDGEMENTS First and foremost I would like to thank my parents, Pam Breza and Jim Breza for being supportive of my love and appreciation of biology. Without them I would have never respec ted the Earth that I love to study. I would like to thank both of my grandparents, Jesse Poore and Sandy Sartin; they made funding my trip to Costa Rica possible. Thank you Allan and Karen Masters for teaching me the knowledge I used to conduct this expe riment. Thanks you Tania Chavarría for providing me all the guidance through the good and difficult times of data collection and statistical analyses. Thank you Willow Zuchowski and Bill Haber for helping me identify the eighty or so plant species. Than k you Pablo Allen for letting me know that I didnt actually identify a new species of beetle and that it was just an odd looking beetle larvae, and thank you Moncho Calderón for always providing me with project supplies. You guys are the greatest TA's on earth ! Last but not least I would like to say thank you to all off my fellow students; my experience in Costa Rica would be dull with out every single one of you!

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LITERATURE CITED Chen, B. and D. H. Wise. 1999. Bottom Up Limitation of Predaceous Arthr opods in a Detritus Based Terrestrial Food Web. Ecology 803: 761 772 . Hättenschwiler, S., P. Gasser, C.B. Soil Animals Alter Plant Litter Diversity Effects on Decomposition. 2005. Proceedings of the National Academy of Sciences of the United States of A merica 1025: 1519 1524 Lagerlöf, J., H. Wallin. 1993. The abundance of arthropods along two field margins with different types of vegetation composition: an experimental study. Agriculture, Ecosystems and Environment 43: 141 154 Siemann, E . 1998. Exper imental Tests of Effects of Plant Productivity and Diversity on Grassland. Ecology 796: 2057 2070 . Siemann, E, J . Haarstad and D . Tilman. 1999. Dynamics of plant and arthropod diversity during old fied succession. Ecography 22: 406 414. Schmitz, O . J., E . L. Kalies and M . G. Booth. 2006. Alternative Dynamic Regimes and Trophic Control of Plant Succession. Ecosystems 9: 659 672 . Veldman, J . W., K. G . Murray, A . L. Hull, J. M . Garcia, W . S. Mungall, G . B. Rotman, M . P. Plosz and L . K. McNamara. 2007. Ch emical Defense and the Persistence of Plant Seeds in the Soil of a Tropical Cloud Forest. BIOTROPICA 391: 87 € 93 . Wardle, D.A., K.I. Bonner, G.M. Barker, G.W. Yeates, K.S. Nicholson, R.D Bardgett, R.N., Watson, A. Ghani. 1999. Plant Removals in Perennial Grassland: Vegetation Dynamics, Decomposers, Soil Biodiversity,and Ecosystem. Ecological Monographs 694: 535 568 Whittaker, R . H. Production. 1975. Communities and Ecosystems . MacMillan Publishing, New York. Pages 218 119. Zuchowski, W. 2005. Special Habitats: Tropical Montane Cloud Forests. A Guide to Tropical Plants of Costa Rica . David Featherstone, ed. Distribuidores Zona Tropical, S.A., Miami. Page 187

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APPENDIX Closed Forest Taxon Number of Individuals Taxon Number of Individuals Aracaceae 1 2 Pteridophyta H 1 Aracaceae 2 1 Pteridophyta I 6 Ardisia sp. 1 Pteridophyta J 3 Asteraceae 1 4 Pteridophyta K 3 Begonia involuorata 14 Pteridophyta L 4 Cercropia 1 1 Pteridophyta M 1 Cucurbitaceae 1 2 Rubiaceae 1 7 Eugenia sp. 2 Sanicula liberta 1 Geonoma sp. 1 6 Sideroxylon portoricense 1 Hampea appendiculata 10 Symplucos sp. 1 Hoffmannia laxa 5 Tibouchina longifolia 1 Icanthaceae 1 16 Unknown 1 3 Inga punctata 1 Unknown 2 2 Justicia sp. 12 Unknown 3 5 Melastome 1 5 Unknown 4 1 Mollinedia sp. 2 Unknown 5 2 Monstera adensoni 15 Unknown 6 1 Monstera sp. 1 Unknown 7 1 Paulinia sp. 3 Unknown 8 3 Philodendron sp. 1 3 Unknown 9 1 Piper hispidium 3 Unknown 10 4 Piper sp. 1 3 Unknown 11 1 Pouteria exforliata 2 Unknown 12 2 Pteridophyta A 4 Unknown 13 1 Pteridophyta B 10 Unknown 14 6 Pteridophyta C 1 Unknown 15 2 Pteridophyta D 1 Unknown 16 1 Pteridophyta E 2 Unknown 17 2 Pteridophyta F 2 Unknown 18 3 Pteridophyta G 2 Unkown Vine 1 1 Witheringia sp. 1 TABLE 1. List of all plant taxa found in the continuous forest transects

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Gap Forest Taxon Number of Individuals Taxon Number of Individuals Begonia involvorata 15 Pteridophyta Q 4 Besleria princeps 2 Pteridophyta R 2 Cissus mortiniana 1 Pteridophyta S 3 Conostegia pitteri 5 Pteridophyta T 5 Costus sp. 1 Rondeletia amoena 1 Cucurbitaceae 2 1 Rubiaceae 2 11 Geonoma sp. 1 Senecio cooperi 1 Hampea appendicula 3 Smilax sp. 5 Heliconia monteverdensis 8 Smilax sp. 2 1 Hydrocotyle leucocephala 13 Solanace sp. ? 3 Hydrocotyle mexicana 2 Ucinia hamata 2 Icanthaceae 3 15 Unknown 19 2 Justicia sp. 6 Unknown 20 11 Lasicis sp. 1 Unknown 21 1 Metastelma spp. 12 Unknown 22 1 Myrsina coriacea 7 Unknown 23 2 Neomirandia angularis 1 Unknown 24 2 Notopleura uliginosa 3 Unknown 25 4 Ophismenus borbannii 9 Unknown 26 5 Passiflora biflora 3 Unknown 27 1 Peperomia hernandiafolia 12 Unknown 28 2 Philodendron aurantiifolium 1 Unknown 29 3 Pilea auriculata 2 Unknown 30 13 Pilea pitteri 5 Unknown 31 3 Piper sp. 1 13 Unknown 32 39 Po uzolia perasitica 33 Unknown 33 2 Pteridophyta N 6 Verronia arborescens 3 Pteridophyta O 6 Viola stipularis 24 Pteridophyta P 7 Xanthosoma undipes 2 TABLE 2. List of all plant taxa found in the gap forest transects

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Closed Forest Gap Forest Tax on Number of Individuals Taxon Number of Individuals Acari 9 Acari 4 Blattodea 4 Chilopoda 13 Chilopoda 15 Coleoptera 24 Coleoptera 44 Collembola 3 Collembola 1 Copeopoda 1 Copepoda 1 Diplopoda 9 Diplopoda 20 Diptera 7 Diptera 4 Formicid ae 40 Forficulidae 1 Hemiptera 5 Formicidae 37 Isopoda 2 Hemiptera 8 Lepodoptera 1 Hymenoptera 1 Aracnidae 4 Isopoda 7 Lepodoptera 3 Aracnidae 2 TABLE 3. List of all arthropod taxa found in continuous and gap forest transects


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