Diversity and richness of epiphytes on strangler fig trees Sullivan 1 Diversity and r ichness of n on v ascular e piphytes on s trangler f igs : effects of elevation on emergent trees Kimberlie M. Sullivan Department of Social Sciences, Environmental Studies, University of California, Santa Cruz EAP Tropical Biology and Conservation Spring 2019 7 June 2018 ABSTRACT The Monteverde cloud forest is characterized by its high precipitation levels, frequency of cloud immersion, and extraction of cloud water , but global climate models indicate a reduction in low level cloudiness, suggesting cloud forests may soon be replac ed by forests that occur at lower altitud es . Studies found that n on vascular epiphytes are very sensitive to changes in moisture and temperature, resulting in a substantial deterioration in health, su ch as reduced growth rates, water holding capacity, and ability to fix nitrogen, which makes non vascular epiphytes a compelling indicator of climate change . I conducted a survey that contrasts epiphyte diversity and species richness on emergent strangler fig tree s at two elevations as well as compares diversity and species richness in the u nderstory to the upper canopy. My survey showed lichen have the highest species richness and that there is significantly greater abundance of lichen at 1470 m compared to 1200m . My study indicated that different morphospecies of lichen grow at specific elevations, with only 28% of the same morphospecies of lichen prevalent at both elevations. This study supports previous studies that found differences in epiphyte communities due to variances in microclimates along emergent trees and at different elevations. D iversid ad y riqueza de epÃfitas no vasculares en higuerones : efecto de la elevaci Ã³ n . RESUME N El bosque nuboso de Monteverde se caracteriza por sus altos niveles de precipitaciÃ³n, la frecuente nubosidad y el goteo de agua, pero los modelos climÃ¡ticos globales indican una reducciÃ³n en la nubosidad, lo que sugiere que los bosques nubosos eventualment e serÃ¡n reemplazados por bosques similares a los de altitudes mÃ¡s bajas. Los estudios han encontrado que las epÃfitas no vasculares son muy sensibles a los cambios en la humedad y la temperatura, lo que produce un deterioro sustancial en su salud, como la reducciÃ³n de las tasas de crecimiento, la capacidad de retenciÃ³n de agua y la capacidad de fijar nitrÃ³geno, lo que hace que las epÃfitas no vasculares sean un indicador convincente del cambio climÃ¡tico. RealicÃ© un muestreo de la diversidad de epÃfitas y la riqueza de especies en higuerones estranguladores emergente en dos elevaciones, y comparÃ© la diversidad y la riqueza de especies en el sotobosque con el dosel superior. Mi estudio mostrÃ³ que los lÃquenes tienen la mayor riqueza de especies y que existe un a abundancia significati vamente mayor de lÃquenes a 1470 m de elevaciÃ³n
Diversity and richness of epiphytes on strangler fig trees Sullivan 2 en comparaciÃ³n con los 1200 m de elevaciÃ³n. Mi estudio indicÃ³ que las diferentes morfoespecies de los lÃquenes crecen a elevaciones especÃficas, con solo el 28% de las mismas morfoespe cies de lÃquenes ocurriendo en ambas elevaciones. Este estudio apoya estudios previos que encontraron diferencias en las comunidades epÃfitas debido a las variaciones en los microclimas entre Ã¡rboles emergentes y en diferentes elevaciones. Tropical montane cloud forests are regions of high biodiversity (Nadkarni & Wheelwright, 2000) . Cloud forests are characterized by their high precipitation levels, frequency of cloud immersion, and extraction of cloud water that only occurs in a narrow altitudinal range (Ray, 2013). Monteverde encompasses a unique variety of topography, containing six Holdridge life zones ( Haber 2000 & BolaÃ±os , et al ., 2005 cited by Nadkarni &Wheelwright, 2000 ). Many species found in Monteverde have notable responses to the diverse microclimates that exist along the elevational gradient , creating species stratification in a relatively small area (HÃ¤ger 2010 cited by Nadkarni & Wheelwright, 2000). Ficus tuerckheimii , s trangler figs, are one of the five most common canopy trees in the Monteverde cloud forest (Nadkarni & Solano 2002) and are also a keystone species because they provide ample food and shelter for other organisms as well as host many different epiphytes (Lamen, 1995). Epiphytes ar e a significant constituent of plant diversity in the Monteverde cloud forest because they consist of many different species, such as bromeliads, orchids, ferns, and are immensely abundant. Epiphytes play a crucial role in ecosystem processes, such as the hydrological cycle (Ah Peng, et al., 2017). For example, dense areas of epiphytes on trees in cloud forests capture and preserve large amounts of rainwater that help maintain hig h humidity that is crucial for this ecosystem (Nadkarn i & Wheelwright, 2000 ). Global climate models indicate a reduction in low level cloudiness, suggesting cloud forests may be replaced by forests that occur at lower elevations , and the Monteverde cloud forest appears to be enduring this ecological transition (Foster, 2001). Epiphytes are tightly connected with atmospheric inputs and are endangered by climatic change s , especially non vascular epiphytes (mosses, liverworts and lichen). There are known microclimates along emergent trees in tropical forests due to variances in shade coverage, temperature, moisture, and vapor pressure vary along a single tree, which affects what epiphytes grow on different areas of the tree (Nadkarni & Solano 2002). The upper canopy of an emergent tree (defined in Figure 1 ) is purportedly the optimal area for epiphyte diversity and species richness (Bohlman, Matelson, & Nadkarni, 1995), with section two being the primary zone for epiphytes covering boughs and section three being a drier area typically amass with lichen (Nadkarni & Wheelwright, 2000 ). Epiphytes that exist in the microhabitat of the upper canopy undergo a more frequent cycling of wet and dry conditions, more sunlight, and more air movement compared to u nderstory epiphytes (Nad karni & Wheelwright, 200 ), while the understory is consistently much moister than the upper canopy, especially during the dry season (Bohlman, et al., 1995 ). Studies found that non vascular epiphytes are very sensitive to changes in moisture and temperature, resulting in a substantial deterioration in health, such as reduced growth rates, water holding capacity, and ability to fix nitrogen , if their microclimate undergoes even a slight chang e (Song, Liu, &
Diversity and richness of epiphytes on strangler fig trees Sullivan 3 Nadkarni, 2012). This extreme sensitivity to microclimatic factors makes non vascular epiphytes a compelling indicator of climate change because even a slight alteration to their ecosystem can result in severe reverberations for this community. It is crucial to study the upper canopy because canopy plants, such as epiphytes, play a significant role in carbon, nutrient, and water cycles ( Song, Liu, & Nadkarni, 2012 & Ah Peng, et al., 2017 ) . Since epiphytes have extremely limited to no access to resources on the ground, they are extremely vulnerable to changes in climate, such as increasing canopy temperatures, decreasing precipitation levels, and declining cloud i mmersion ( Ah Peng, et al., 2017 & Liang, et al.). Previous studies found that presence of certain epiphyte species are restricted to specific areas on emergent trees; for example, one study juxtaposed bryophyte species diversity in the upper canopy to th e understory, with 52% of bryophytes species occurring only in the upper canopy and 20% existing solely in the understory (Gradstein , Griffin, Morales, & Nadkarni , 2000 ) . It is crucial to study this community in order to fully understand the Monteverde cloud forest and to more accurately cognize their diversity . The Monteverde cloud forest is extremely susceptible to the negative ramifications of climate change and may already be suffering from these consequences, so it is imperative to conduct a survey of epiphytes in the upper canopy to note current diversity and establish a reference for future studies before this ecosystem is severely altered. I conducted a survey that contrasts epiphyte diversity and species richness on an emergent strangler fig tree at two elevations as well as collates diversity and species richness in the understory to the upper canopy, and I aimed to answer the central question, does non vascular epiphyte diversity differ with elevation and heigh t on strangler figs? METHODS I surveyed epiphyte diversity and species richness on stranger fig trees in the Monteverde cloud forest at two elevations : behind the Monteverde Institute ( 1470 m ) and at Santuario Ecologico ( 1200 ). I surveyed three trees behind Monteverde Institute, elevations 1466, 1473, and 1477, and three trees at Santuario Ecologico, 1200. The Monteverde Institute is primary growth forest , while Santuario Ecologico is mostly secondary growth. I also sampled si x areas on a single emergent tree behind Monteverde Institute (Tree 1). I used a GPS to determine the coordinates and elevation of each tree, a compass to note the cardinal direction of the area sampled both of the area around the trunk of the tree as we ll as of each distinct branch sampled in the canopy . On each strangler, I observed epiphytes in the understory and upper canopy at the three areas with the highest concentration of epiphytes on the trunk of t he tree . I surveyed three branches and three vertical areas in the canopy . I used a 50cm by 50cm grid to standardize the area surveyed and to assess the total percent coverage using a 1 5 scale (1: 0 15%; 2: 15 29%; 3: 30 64%; 4:65 99%; 5:100%). In the understory, I sampled areas within a range if 60 to 180m , and I measured the exact height range of each area sampled. I measured the height of each branch and area sampled in the canopy. I noted general diversity of vascular epiphytes by counting the number of stems and counting the number of
Diversity and richness of epiphytes on strangler fig trees Sullivan 4 different morpho species. I surveyed with a more detailed focus on non vascul ar epiphytes, specifically moss and lichen, by counting the number of morpho species of moss and lichen and measuring the area of moss and lichen in square centimeters. I took photos of the distinct morpho species of lichen I observed so that I could com pare and distinguish between morpho species of lichen I found amongst stranglers and among elevations. I use d Coefficient of Sorenson to calculate similar ity of moss and lichen between 1470 m and 1200m . I used JMP to preform statistical tests, and I calculated correlation coefficient using https://www.socscistatistics.com. RESULTS I found 51 morphospecies of lichen , six morphospecies of moss , and three species of vascular epiphytes around the trunk of strangler figs in the understory. Tree 1 has the greatest species richness of lichen and moss, with 30 morphospecies of lichen and 6 morphospecies of moss. Tree 2 and 4 contained the least morphospecies of lichen, with a total of 17 morphospecies of lichen observed, and Tree 3 and 4 contain the least morphospecies of moss, with a total of 3 morphospecies of moss observed (Figure 1 ). I found lichen to be the most abundant epiphyte for all stranglers but Tree 5 (Figure 2 ). Morphospecies of moss and lichen as well as species of vascular epiphytes observed also varied among the areas sampled on each tree. For example, the greatest number of morphospecies of lichen I found in a 50cm by 50cm sampled area on an emergent tree is 18, which I observed on T1A1, T1A3, and T5A3, while the least number morpho species of lichen I found in a single quadrat is six on T4A3 (Figure 3 ). I found that the correlation between abundance of moss and lichen are highly negative ly correlated (Figure 4 . R 2 significant relationship between abundance of non vascular epiphytes and vascular epiphytes (Figure 5 and Figure 6 ).
Diversity and richness of epiphytes on strangler fig trees Sullivan 5 Figure 1. Total species richness per tree in the understory shown as the total number of morpho species of moss and lichen found on each emergent strangler fig. Trees 1 3 are located behind Monteverde Institute, and trees 4 6 are in Santuario Ecol Ã³ gico. Figure 2. Total abundance of moss a nd lichen per tree in the understory shown as the total area of moss and lichen in square centimeters on each emergent strangler fig. 0 5 10 15 20 25 30 35 Tree 1 Tree 2 Tree 3 Tree 4 Tree 5 Tree 6 Nunmber of morpho species Morphospecies of moss Morpho-species of lichen 0 1000 2000 3000 4000 5000 6000 7000 8000 Tree 1 Tree 2 Tree 3 Tree 4 Tree 5 Tree 6 Area Total square cm of moss Total square cm of lichen
Diversity and richness of epiphytes on strangler fig trees Sullivan 6 Figure 3. Total species richness per area on an emergent tree in the understory (T=Tree, A= Specific area sampled on e ach strangler fig; T1A1= Tree 1, Area 1, etc.) shown as the total number of morpho species of moss and lichen found at each area sampled on an emergent strangler fig. Trees 1 3 are located behind Monteverde Institute, and trees 4 6 are in Santuario Ecol Ã³ g ico. Figure 4. Negative c orrelation between cm 2 of moss and lichen cover in the understory. ( R 2 = .82189, p<.0005 ) . 0 2 4 6 8 10 12 14 16 18 20 T1A1 T1A2 T1A3 T2A1 T2A2 T2A3 T3A1 T3A2 T3A3 T4A1 T4A2 T4A3 T5A1 T5A2 T5A3 T6A1 T6A2 T6A3 Number of morpho species vascualr species morphospecies of moss morphospecies of lichen 0 500 1000 1500 2000 2500 3000 0 500 1000 1500 2000 Squared cm lichen Squared cm of moss R 2 = .82189
Diversity and richness of epiphytes on strangler fig trees Sullivan 7 Figure 5. No significant correlation between abundance of lichen a nd abundance of vascular stems in the understory ( R 2 = 0.12322 ) . Figure 6. No significant correlation between number of vascular stems and squared cm of moss in the understory , R 2 =0.15548 . 0 500 1000 1500 2000 2500 3000 0 2 4 6 8 10 12 Squared cm of lichen Number of Vascular Epiphyte Stems 0 200 400 600 800 1000 1200 1400 1600 1800 2000 0 2 4 6 8 10 12 Squared cm of moss Number of Vascular Stems
Diversity and richness of epiphytes on strangler fig trees Sullivan 8 I found the abundance of lichen to be significantly greater at 1470m than at 1200m (Figure 7 . t test, n=18, p= .0232) , but lichen was still more prevalent than moss on average at each elevation. I observed greater average abundance of mos s at elevation 2 than e levation 1 (Figure 8 ). I noted greater average species richness of lichen on emergent strangler figs a t 1470 m than 1200m , and I found no substantial variation of species richness of moss and vascular epiphytes between elevations (Figure 8 ). At e levation 1, I found less variance in morpho species of lichen amongst Trees 1,2,3, with each tree having 52 72% of morpho speci es of lichen in common (Table 1 ), but a greater variance in moss found on all trees, with each tree having 29 40% of morpho species of moss foun d on all trees (Table 2 ). At e levation 2, I noted a greater variance in both lichen and moss amongst trees 4,5,6, with 23 30% similarity of morpho species of lichen (Table 3 ) and 25 40% similarity of morpho species of moss found on all trees (Table 4 ). I observed the same morpho species of moss at e levatio n 1 and 2, but I found significant variance in the morpho species of lichen between the two elevations, finding only 28% of morpho species of lic hen at both elevations (Table 5 & Table 6 ) . Figure 7. Average abundance of moss and lichen observed on strangler figs in the understory at 1470 and elevation 1200 . Error bars indicate standard deviation. 0 500 1000 1500 2000 2500 3000 elevation 1 elevation 2 Area avg cm squared moss avg cm squared lichen 1470m 1200m
Diversity and richness of epiphytes on strangler fig trees Sullivan 9 Figure 8 . Average species richness of vascular epiphytes, moss, lichen observed on strangler figs i n the understory at elevation 1 and elevation 2. Error bars indicate standard deviation. Table 1 : Similarity in morpho species of lichen amongst strangler fig trees 1,2,3 at 1470m using Coefficient of Sorenson. T ree 1 T ree 2 T ree 3 T ree 1 ---T ree 2 .66 --T ree 3 .72 .52 -Table 2 : Similarity in morpho species of moss amongst strangler fig trees 1,2,3 at 1470m using Coefficient of Sorenson . T1 T2 T3 T1 ---T2 .4 --T3 .33 .29 -Table 3 : Similarity in morpho species of lichen amongst strangler fig trees 4,5,6 at 1200m using Coefficient of Sorenson . T4 T5 T6 T4 ---T5 .26 --T6 .23 .30 -0 2 4 6 8 10 12 14 16 18 20 1470m 1200m Average number of morpho species avg morphospecies vascular morphospecies moss avg mosphospecies lichen
Diversity and richness of epiphytes on strangler fig trees Sullivan 10 Table 4 : Similarity in morpho species of moss amongst strangler fig trees 4,5,6 at 1200m using Coefficient of Sorenson . T4 T5 T6 T4 ---T5 .25 --T6 .38 .40 -Table 5 : Similarity in morpho species of lichen and moss between all trees at 1470m and 1200m using Coefficient of Sorenson. Table 6 : Presence or absence of lichen in strangler fig trees at 1470m and1200m. Morphospecies of lichen 1200m 1470m JJ 1 0 KK 1 0 LL 1 0 MM 1 0 NN 1 0 OO 1 0 PP 1 0 QQ 1 0 RR 1 0 SS 1 0 TT 1 0 UU 1 0 VV 1 0 WW 1 0 XX 1 0 YY 1 0 B 1 1 C 1 1 D 1 1 E 1 1 G 1 1 J 1 1 O 1 1 P 1 1 R 1 1 U 1 1 V 1 1 W 1 1 Y 1 1 1470m and 1200m Lichen .28 Moss 1
Diversity and richness of epiphytes on strangler fig trees Sullivan 11 Z 1 1 AA 1 1 BB 1 1 CC 1 1 BB 1 1 DD 1 1 EE 1 1 II 1 1 F 0 1 FF 0 1 GG 0 1 H 0 1 HH 0 1 I 0 1 K 0 1 L 0 1 M 0 1 N 0 1 Q 0 1 S 0 1 T 0 1 X 0 1 I found both the number of vascular epiphyte stems and number of morpho species of vascular plants to be significantly greater in the canopy than in the understory (Figure 10 . T test, n=6,3, p=.034 & Figure 11 , t test, n=6,3, p=0.285 ). I found the abunda nce and species richness of lichen to be significantly greater in the understor y than in the canopy (Figure 12 , t test , n=6,3, p<.05 & Figure 13 , t test, n=6,3, p<.05). Figure 9. V ascular epiphytes abundance in the understory and the canopy ( t test, n=6,3, p=.034 ) . 0 5 10 15 20 25 30 35 40 0 0,5 1 1,5 2 2,5 Number pf vascualr stems Understory Upper canopy
Diversity and richness of epiphytes on strangler fig trees Sullivan 12 Figure 1 0. V ascular epiphytes species richness in the understory and upper canopy ( t test, n=6,3, p=0.285 ) Figure 1 1. L ichen abundance in the understory and canopy in squared cm ( t test, n=6,3, p<.05 ) 0 2 4 6 8 10 12 14 16 18 20 0 0,5 1 1,5 2 2,5 Understory Canopy 0 500 1000 1500 2000 2500 0 0,5 1 1,5 2 2,5 Area Tree Understory Canopy
Diversity and richness of epiphytes on strangler fig trees Sullivan 13 Fig ure 1 2. Lichen species richness in the understory and canopy ( t test, n=6,3, p<.05 ) DISCUSSION My survey showed lichen have the highest species richness and that there is significantly greater abundance of lichen a t the higher elevation compared to the lower elevation . My study indicated that different morpho species of li chen grow at specific elevations: only 28% of the same morpho species of lichen were prevalent at both elevations . Numerous studies stated that Monteverde is enduring a n ecological transition from a cloud forest to forests that traditionally occupied a lower altitudinal region due to a reduction of low level cloudiness , which may eliminate epiphyte communities that are only found in the Monteverde cloud forest ( Foster, 2007; Nalini & Solano 2002; Ray, 2013 ) . Nadkarni and Solano (2002) conducted an experiment that exemplifies the effects of distinct microclimates on epiphyte communities. They found that when epiphytes were transferred from an elevation of 1480m to trees that were 70 declined by either substantially decreas ing in size or experienc ing season al dependent mortality. This shows that t he morpho species of epiphytes that grow along an elevational gradient are effected by the amount of cloud immersion each microclimate experiences. This transition and its ramifications for epiphyte species richness explain s the discrepancies in lichen co mmunities I found between the higher elevation and the lower e levation . A substantially greater abundance of lichen at 1470m and s uch a low similarity rate between elevations indicates that t he 28% of lichen morpho species that are also found at the lower elevation are currently being phased out for morpho species of lichen that traditionally grew at lower elevations. 0 2 4 6 8 10 12 14 16 18 20 0 0,5 1 1,5 2 2,5 Number of morpho species Tree Understory Canopy
Diversity and richness of epiphytes on strangler fig trees Sullivan 14 I did not find significant difference in specie s richness between elevations, which is likely due to the fact that when areas of forest are cleared, strangler figs are left intact. A study done by Nadkarni and Haber ( 2009 ) surveyed the seed banks of remnant pasture trees, trees that remained while the majority of the forest was cleared for pasture, and found that soils and epiphyte mats that accumulate in the upper canopy also accumulate seed banks, like soils on the forest floors. These seed banks are dense and diverse, containing many species that occ ur predominately on the forest floor, and help regenerate forest s , which emphasizes the importance of epiphyte communities in forest regeneration and explai ns why strang lers in the primarily secondary growth forest are dense with epiphytes . This clarifies the lack of variation I found in spe cies richness among elevations. Lichen are known to trap seeds (Favero & Piervittori, 2010) , and b ecause strangler figs are left intact, the seed mats of remnant pasture trees remain , thereby preserving epiphyte species richness in secondary growth forests . Another study completed by Sheldon and Nadkarni (2013) deposited in the forest floor to the canopy and found significant differences in the composition of seed rain but found that seeds dispersed by birds were most common overall epiphyte seeds composed of most seed in the canopy while larger , independent plants accounted for most seeds in the understory . Sheldon and Nadkarni c oncluded that dispersal agents deposit the seeds to the site that fosters the most growth for each plant . This finding of greater vascular seed abundance in canopy seed banks provides insight as to why I found more diversity of vascular epiphytes in the canopy than in the understory. Dispersal agents deposit more vascular epiphyte seeds to the canopy because canopy conditions are better for plant growth, resulting in a greater abundance of vascular epiphytes in the canopy. I found a substantial negative correlation between moss and lichen abundance in the understory , and I observed a greater species richness and abundance of lichen in the understory tha n in the upper canopy . Lichen are composed of the symbiotic relationship between a filamentous fungus and minimally one photosynthetic organism . They are dispersed by wind and require very specific conditions to reproduce. In order to reproduce asexually , lichen fungal and photosynthetic partners must transmit from one generation to the next. To reproduce sexually, lichen require the germination of fungal spores with a compatible photobiont, which resynthesize s the lichen symbiosis (Lutzoni & Miadlikowaska , n.d. ) . Once lichen develop, it can severely alter the microclimate it inhabits. Previous studies found that lichen can kill neighboring moss or even overgrow the substrate and modify microclimatic factors, such as light irradiance, creating a more favorable habitat for lichen (Favero Longo & Piervittori 2010). U nderstory conditions are substantially more constant than the canopy microclimate, so perhaps lichen in the understory have been able to more effec tively alter understory microclimate conditions, which explains why I found such a higher species richness in lichen than in moss and other vascular epiphytes in the understory. This also provides a possible explanation as to why I observed the same si x morpho species of moss at 1470 m and 1200m. Since I found lichen to be more abundant than moss but observed the same six morpho species of moss at both elevations, this suggests that these morpho species of moss are able to grow alongside lichen and can withstand differences in microclimate factors that lichen create .
Diversity and richness of epiphytes on strangler fig trees Sullivan 15 Epiphytes are immensely diverse and abundant in cloud forests, with species variance depending on location in the forest and position on emergent strangler figs. Epiphytes are imperative to the Monteverde cloud forest , but studying their effects on cloud forest ecological processes is a fairly recent endeavor. These studies ameliorate our knowledge of the crucial role in hydrological, nutrient, and carbon cycles , helping us understand the intricacies of cloud forests. Long term studies should be done to compare and track the abundance of the lichen that grows at b oth elevations to prove whether it is decr easing in abundance at the lower elevation . Further research should investigate the correlation between the abundance of mos s and lichen to understand ACKNOWLEDGEMENTS This project could not have been done without the knowledge, support, and patience of biologist (not liar) Federico Chinchilla whose humor mitigated the vicissitude of my project . Thanks to Frank Joyce for making my tree climbing dreams come true , teaching me how to run stats, and offering crucial insight . Special than ks to the wonderful Cruz family for their warmth and hospitality. Additional thanks to Santuario Ecologico , the staff at EstaciÃ³n BiÃ³logica, and Monteverde Institute. LITERATURE CITED Ah Peng, C., Cardoso, A. W., Flores, O., West, A., Wilding, N., Strasberg, D., & Hedderson, T. A. 2017. The role of epiphytic bryophytes in inte rception, storage, and the regulated release of atmospheric moisture in a tropical montane cloud forest. Journal of Hydrology,548 : 665 673. Bohlman, S. A., Matelson, T. J., & Nadkarni, N. M. 1995. Moisture and Temperature Patterns of Canopy Humus and Forest Floor Soil of a Montane Cloud Forest, Costa Rica. Biotropica,27(1) : 13. Favero Longo, S. E., & Piervittori, R. 2010. Lichen plant interactions. Journal of Plant Interactions,5 (3) : 163 177. Foster, P. 2001. The potential negative impacts of global climate change on tropical montane cloud forests. Earth Science Reviews,55(1 2) : 73 106. Gotsch , S. G., Nadkarni, N., Darby, A., Glunk, A., Dix, M., Davidson, K., & Dawson, T. E. 2015. Life in the treetops: Ecophysiological strategies of canopy epiphytes in a tropical montane cloud forest. Ecological Monographs,85(3) : 393 412. Gradstein, S. R., Griffin, D., III, Morales, M. I., & Nadkarni, N. M. 2000. Diversity and habitat differentiation of mosses and liverworts in the cloud forest of Monteverde, Costa Rica. Caldasia, 203 212. Retrieved May 29, 2019, from http://www.bdigital.una l.edu.co/ Laman, T. 1995. The ecology of strangler fig seedling establishment . Selbyana, 16(2) : 223 229. Retrieved from http://www.jstor.org/stable/41759910
Diversity and richness of epiphytes on strangler fig trees Sullivan 16 Longman, K. A., & JenÃk, J. 1987. Tropical forest and its environment (2nd ed.). New York, NY: Wiley. Lutzoni, F., & Miadlikowaska, J. (n.d.). Lichens. Current Biology,19 (1 3). Retrieved June 7, 2019, from https://www.cell.com/current biology/pdf/. Nadkarni, N. M., & Haber, W. A. 2009. Canopy Seed Banks as Time Capsules of Biodiversity in Pasture Rem nant Tree Crowns. Conservation Biology,23 (5) : 1117 1126. Nadkarni, N. M., & R. Solano . 2002. Potential effects of climate change on canopy communities in a tropical cloud forest: An experimental approach. Oecologia,131( 4) : 580 586. Nadkarni, N. M., & Kohl, K. D. 2018. Elements of disturbance that affect epiphyte vitality in a temperate rainforest: An experimental approach. Journal of Plant Ecology,12(2) : 306 313. Nadka rni, N. M., & Wheelwright, N. T. (Eds.). 2000. Monteverde: Ecology and conservation of a tropical cloud forest. New York: Oxford Univ. Press. Sheldon, K. S., & Nadkarni, N. M. 2013. Spatial and t emporal v ariation of s eed r ain in the c anopy and on the g round of a t ropical c loud f orest. Biotropica . 45 (5), 549 556. Song, L., Liu, W., & Nadkarni, N. M. 2012. Response of non vascular epiphytes to simulated climate change in a montane moist evergreen broad leaved forest in southwest Chin a. Biological Conservation,152, 127 135.