Skip to main content

The functional roles of epiphytes and arboreal soils in tropical montane cloud forests

  • Sybil G. Gotsch (a1), Nalini Nadkarni (a2) and Autumn Amici (a2)

Epiphytes and their associated decomposing litter and arboreal soils (herein, epiphytic material, EM) are ubiquitous features of tropical montane cloud forests (TMCF) and play important roles in ecosystem function. EM intercepts water and nutrients from the atmosphere and from intercepted host tree sources, and may contribute significant inputs of these resources to the forest floor. Despite the importance of EM in the TMCF, a systematic review of the ecosystem roles of EM has not been compiled before. We have synthesized the literature that documents functions of EM in undisturbed TMCFs and discuss how these roles may be affected by disturbances, including changes in climate and land use. The range of EM biomass and water storage in the TMCF varies greatly across sites, with different amounts associated with stand age and microclimate. EM is important as habitat and food for birds and mammals, with over 200 species of birds documented as using EM in the Neotropics. Given its sensitivity to moisture, projected shifts in cloud base heights or precipitation due to changes in climate will likely have a large impact on this community and changes in EM diversity or abundance may have cascading impacts on the ecosystem function of the TMCF.

Corresponding author
1 Corresponding author. Email:
Hide All
ACKERMAN, J. D. 1986. Coping with the epiphytic existence: pollination strategies. Selbyana 9:5260.
AFFELD, K., SULLIVAN, J., WORNER, S. P. & DIDHAM, R. K. 2008. Can spatial variation in epiphyte diversity and community structure be predicted from sampling vascular epiphytes alone? Journal of Biogeography 35:22742288.
ALDRICH, M., BUBB, P., HOSTETTLER, S. & VAN DE WEIL, H. 2000. Tropical montane cloud forests: time for action. WWF International, IUCN, Gland. 28 pp.
BENZING, D. 1990. Vascular epiphytes. Cambridge University Press, Cambridge. 376 pp.
BOHLMAN, S., MATELSON, T. J. & NADKARNI, N. M. 1995. Moisture and temperature patterns of canopy humus and forest floor soils of a montane cloud forest, Costa Rica. Biotropica 27:1319.
BOUCHER, V. L. & NASH, T. H. 1990. The role of the fruiticose lichen Ramalina menziesii in the annual turnover of biomass and macronutrients in a Blue Oak woodland. Botanical Gazette 151:114118.
BRUIJNZEEL, L. A. & PROCTOR, J. 1995. Hydrology and biogeochemistry of tropical montane cloud forests: what do we really know? Ecological Studies 110:3878.
BRUIJNZEEL, L. A., SCATENA, F. & HAMILTON, L. (eds.). 2010. Tropical montane cloud forests: science for conservation and management. Cambridge University Press, Cambridge. 768 pp.
BUSH, S. P. & BEACH, J. H. 1995. Breeding systems of epiphytes in a tropical montane wet forest. Selbyana 16:155158.
CARDELÚS, C. L. & MACK, M. 2010. The nutrient status of epiphytes and their host trees along an elevational gradient in Costa Rica. Plant Ecology 207:2537.
CARROLL, G. 1980. Forest canopies: complex and independent subsystems. Pp. 87107 in Waring, R. H. (ed). Forests: fresh perspectives from ecosystem analysis: proceedings of the 40th annual biology colloquium. Oregon State University Press, Corvallis.
CASCANTE-MARÍN, A., OOSTERMEIJER, J. G. B., WOLF, J. H. D. & DEN NIJS, J. C. M. 2005. Reproductive biology of the epiphytic bromeliad Werauhia gladioliflora in a premontane tropical forest. Plant Biology 7:203209.
CAVELIER, J., JARAMILLO, M., SOLIS, D. & DE LEÓN, D. 1997. Water balance and nutrient inputs in bulk precipitation in tropical montane cloud forest in Panama. Journal of Hydrology 193:8396.
CÉRÉGHINO, R., LEROY, C., DEJEAN, A. & CORBARA, B. 2010. Ants mediate the structure of phytotelm communities in an ant-garden bromeliad. Ecology 91:15491556.
CHEN, L., LIU, W. & WANG, G. 2010. Estimation of epiphytic biomass and nutrient pools in the subtropical montane cloud forest in the Ailao Mountains, south-western China. Ecological Research 25:315325.
CLARK, K., NADKARNI, N., SCHAEFER, D. & GHOLZ, H. 1998a. Cloud water and precipitation chemistry in a tropical montane forest, Monteverde, Costa Rica. Atmospheric Environment 32:15951603.
CLARK, K. L., NADKARNI, N. & GHOLZ, H. L. 1998b. Atmospheric deposition and net retention of ions by the canopy in a tropical montane forest, Monteverde, Costa Rica. Journal of Tropical Ecology 14:2745.
CLARK, K., NADKARNI, N. & GHOLZ, H. 2005. Retention of inorganic nitrogen by epiphytic bryophytes in a tropical montane forest. Biotropica 37:328336.
COXSON, D. & NADKARNI, N. 1995. Ecological roles of epiphytes in nutrient cycles of forest ecosystems. Pp. 495543 in Lowman, M. & Nadkarni, N. (eds.). Forest canopies. Academic Press, San Diego.
FONTOURA, T., CAZETTA, E., NASCIMENTO, W. D., CATENACCI, L., DE VLEESCHOUWER, K. & RABOY, B. 2010. Diurnal frugivores on the Bromeliaceae Aechmea depressa L.B. Sm. from Northeastern Brazil: the prominent role taken by a small forest primate. Biota Neotropica 10:351354.
FORMAN, R. T. T. 1975. Canopy lichens with blue-green algae: a nitrogen source in a Colombian rain forest. Ecology 56:11761184.
GANNON, B. & MARTIN, P. H. 2014. Reconstructing hurricane disturbance in a tropical montane forest landscape in the Cordillera Central, Dominican Republic: implications for vegetation patterns and dynamics. Arctic, Antarctic, and Alpine Research 46:767776.
GENTRY, A. H. & DODSON, C. 1987a. Contribution of non-trees to species richness of a tropical rain forest. Biotropica 19:149156.
GENTRY, A. H. & DODSON, C. 1987b. Diversity and biogeography of neotropical vascular epiphytes. Annals of the Missouri Botanical Gardens 74:204233.
GOLLEY, F. B., MCGINNIS, J. T. & CLEMENTS, R. G. 1971. Biomass and mineral structure of some forest ecosystems in Darien, Panama. Turrialba 21:189196.
GOTSCH, S. G., NADKARNI, N. M., 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 85:393412.
GRADSTEIN, S. R., NADKARNI, N. M., KRÖMER, T., HOLZ, I. & NÖSKE, N. 2003. A protocol for rapid and representative sampling of vascular and non-vascular epiphyte diversity of tropical rain forests. Selbyana 24:105111.
HAGER, A. & DOHRENBUSCH, A. 2011. Hydrometeorology and structure of tropical montane cloud forests under contrasting biophysical conditions in north-western Costa Rica. Hydrological Processes 25:392401.
HERTEL, D., HOHLER, L. & RILLIG, M. C. 2011. Mycorrhizal, endophytic and ecomorphological status of tree roots in the canopy of a montane rain forest. Biotropica 43:401404.
HERWITZ, S. 1986. Episodic stemflow inputs of magnesium and potassium to a tropical forest floor during heavy rainfall events. Oecologia 70:423425.
HIETZ, P. & WINKLER, M. 2006. Breeding systems, fruit set, and flowering phenology of epiphytic bromeliads and orchids in a Mexican humid montane forest. Selbyana 27:156164.
HIETZ, P., WANEK, W., WANIA, R. & NADKARNI, N. M. 2002. Nitrogen-15 natural abundance in a montane cloud forest canopy as an indicator of nitrogen cycling and epiphyte nutrition. Oecologia 131:350355.
HOFSTEDE, R., WOLF, J. & BENZING, D. 1993. Epiphytic biomass and nutrient status of a Colombian upper montane rain forest. Selbyana 14:3745.
HÖLSCHER, D., KÖHLER, L., VAN DIJK, A. I. & BRUIJNZEEL, L. A. 2004. The importance of epiphytes to total rainfall interception by a tropical montane rain forest in Costa Rica. Journal of Hydrology 292:308322.
HOLWERDA, F., BRUIJNZEEL, L. A., MUÑOZ-VILLERS, L. E., EQUIHUA, M. & ASBJORNSEN, H. 2010. Rainfall and cloud water interception in mature and secondary lower montane cloud forests of central Veracruz, Mexico. Journal of Hydrology 384:8496.
HSU, C. C., HORNG, F. W. & KUO, C. M. 2002. Epiphyte biomass and nutrient capital of a moist subtropical forest in north-eastern Taiwan. Journal of Tropical Ecology 18:659670.
INGRAM, S. & NADKARNI, N. 1993. Composition and distribution of epiphytic organic matter in a neotropical cloud forest, Costa Rica. Biotropica 25:370383.
KNOPS, J., NASH, T., BOUCHER, V. L. & SCHLESINGER, W. 1991. Mineral cycling and epiphytic lichens: implications at the ecosystem level. Lichenologist 23:309321.
KÖHLER, L. 2002. Die Bedeutung der Epiphyten im ökosystemaren Wasser-und Nahrstoffumsatz verschiedener Altersstadien eines Bergregenwaldes in Costa Rica. Berichte des Forschungszentrums, Waldökosysteme Reihe A, Band 181. University of Göttingen, Gottingen, Germany. 134 pp.
KÖHLER, L., TOBÓN, C., FRUMAU, K. & BRUIJNZEEL, L. 2007. Biomass and water storage dynamics of epiphytes in old-growth and secondary montane cloud forest stands in Costa Rica. Plant Ecology 193:171184.
LAWTON, R., NAIR, U., PIELKES, R. & WELCH, R. 2001. Climatic impact of tropical lowland deforestation on nearby montane cloud forests. Science 294:584587.
LESICA, P. & ANTIBUS, R. 1991. Canopy soils and epiphyte richness. National Geographic Research and Exploration 7:156165.
MARTIN, P. H., SHERMAN, R. E. & FAHEY, T. J. 2004. Forty years of tropical forest recovery from agriculture: structure and floristics of secondary and old growth riparian forests in the Dominican Republic. Biotropica 36:297317.
MAFFIA, B., NADKARNI, N. M. & JANOS, D. P. 1993. Vesicular-arbuscular mycorrhizae of epiphytic and terrestrial Piperaceae under field and greenhouse conditions. Mycorrhiza 4:511.
MATELSON, T. J., NADKARNI, N. M. & LONGINO, J. T. 1993. Survivorship of fallen epiphytes in a neotropical cloud forest, Monteverde, Costa Rica. Ecology 74:265269.
MCCUNE, B. 1993. Gradients in epiphyte biomass in three Pseudotsuga-Tsuga forests of different ages in western Oregon and Washington. Bryologist 96:405411.
MULLIGAN, M. 2010. Modeling the tropics-wide extent and distribution of cloud forest and cloud forest loss, with implications for conservation priority. Pp. 1439 in Bruijnzeel, L. A., Scatena, F. N. & Hamilton, L. S. (eds.). Tropical montane cloud forests: science for conservation and management. Cambridge University Press, Cambridge.
MURCIA, C. 1997. Evaluation of Andean alder as a catalyst for the recovery of tropical cloud forests in Colombia. Forest Ecology and Management 99:163170.
NADKARNI, N. M. 1981. Canopy roots: convergent evolution in rainforest nutrient cycles. Science 213:10241025.
NADKARNI, N. M. 1984a. Epiphyte biomass and nutrient capital of a neotropical elfin forest. Biotropica 24:2430.
NADKARNI, N. M. 1984b. Biomass and mineral capital of epiphytes in an Acer macrophyllum community of a temperate moist coniferous forest, Olympic Peninsula, Washington State. Canadian Journal of Botany 62:22232228.
NADKARNI, N. M. 2000. Colonization of stripped branch surfaces by epiphytes in a lower montane cloud forest, Monteverde, Costa Rica. Biotropica 32:358363.
NADKARNI, N. M. & LONGINO, J. T. 1990. Macroinvertebrate communities in canopy and forest floor organic matter in a montane cloud forest, Costa Rica. Biotropica 22:286289.
NADKARNI, N. M. & MATELSON, T. J. 1989. Bird use of epiphyte resources in neotropical trees. The Condor 91:891907.
NADKARNI, N. M. & MATELSON, T. J. 1991. Dynamics of fine litterfall within the canopy of a tropical cloud forest, Monteverde. Ecology 72:20712082.
NADKARNI, N. M. & MATELSON, T. J. 1992. Biomass and nutrient dynamics of epiphyte litterfall in a neotropical cloud forest, Costa Rica. Biotropica 24:2430.
NADKARNI, N. & SOLANO, R. 2002. Potential effects of climate change on canopy communities in a tropical cloud forest: an experimental approach. Oecologia 131:580584.
NADKARNI, N. M. & WHEELWRIGHT, N. T. (eds.). 2000. Monteverde: ecology and conservation of a tropical cloud forest. Oxford University Press, New York. 573 pp.
NADKARNI, N., SCHAEFER, D., MATELSON, T. & SOLANO, R. 2002. Comparison of arboreal and terrestrial soil characteristics in a lower montane forest, Monteverde, Costa Rica. Pedobiologia 46:2433.
NADKARNI, N., SCHAEFER, D., MATELSON, T. & SOLANO, R. 2004. Biomass and nutrient pools of canopy and terrestrial components in a primary and a secondary montane cloud forest, Costa Rica. Forest Ecology and Management 198:223236.
OGBURN, R. M. & EDWARDS, E. J. 2010. The ecological water use strategies of succulent plants. Advances in Botanical Research 55:179225.
PATIÑO, J., GONZÁLEZ-MANCEBO, J. M., FERNÁNDEZ-PALACIOS, J. M., ARÉVALO, J. R. AND BERMÚDEZ, A. 2009. Short-term effects of clear-cutting on the biomass and richness of epiphytic bryophytes in managed subtropical cloud forests. Annals of Forest Science 66:113.
PÓCS, T. 1980. The epiphytic biomass and its effect on the water balance of two rainforest types in the Uluguru Mountains (Tanzania, East Africa). Acta Botanica Academiae Scientiarum Hungaricae 26:143167.
POUNDS, A., FOGDEN, M. P. L. & CAMPBELL, J. 1999. Biological responses to climate change on a tropical mountain. Nature 398:611615.
PYPKER, T. G., UNSWORTH, M. H. & BOND, B. J. 2006. The role of epiphytes in rainfall interception by forests in the Pacific Northwest. II. Field measurements at the branch and canopy scale. Canadian Journal of Forest Research 36:819832.
RAINS, K. C., NADKARNI, N. & BLEDSOE, C. 2003. Epiphytic and terrestrial mycorrhizas in a neotropical cloud forest, Costa Rica. Mycorrhiza 13:257264.
REMSEN, J. V. & PARKER, T. A. 1984. Arboreal dead-leaf-searching birds of the Neotropics. The Condor 86:3641.
RITTER, A., REGALADO, C. M. & ASCHAN, G. 2009. Fog reduces transpiration in tree species of the Canarian relict heath-laurel cloud forest (Garajonay National Park, Spain). Tree Physiology 29:517528.
SCHIMPER, A. W. 1888. Die epiphytische Vegetation Amerikas. G. Fisher, Jena. 162 pp.
SCOTT, M. G. & HUTCHINSON, T. C. 1987. Effects of a simulated acid rain episode on photosynthesis and recovery in the caribou-forage lichens, Cladina stellaris (Opiz.) Brodo and Cladina rangiferina (L.) Wigg. New Phytologist 107:567575.
SILLETT, T. S. 1994. Foraging ecology of epiphyte-searching insectivorous birds in Costa Rica. The Condor 96:863877.
SONGWE, N.C., FASEHUN, F.W. & OKALI, D. U. U. 1988. Litterfall and productivity in a tropical rainforest, Southern Bakundu Forest Reserve, Cameroon. Journal of Tropical Ecology 4:2537.
STEWART, G. S., SCHMIDT, L., TURNBULL, M., ERSKINE, P. & JOLY, C. 1995. 15N natural abundance of vascular rainforest epiphytes: implications for nitrogen source and acquisition. Plant, Cell & Environment 18:8590.
STRONG, D. 1977. Epiphyte loads, tree falls, and perennial forest disruption: a mechanism for maintaining higher tree species richness in the tropics without animals. Journal of Biogeography 4:215218.
SUGDEN, A. & ROBINS, R. 1979. Aspects of the ecology of vascular epiphytes in Colombian cloud forests, I. The distribution of the epiphytic flora. Biotropica 11:173188.
TANNER, E. V. J. 1980. Studies on the biomass and productivity in a series of montane rain forests in Jamaica. Journal of Ecology 68:573588.
TERBORGH, J. 1986. Keystone plant resources in the tropical forest. Pp. 330344 in Soule, M. E. (ed.). Conservation biology, the science of scarcity and diversity. Sinauer, Sunderland.
TOBÓN, C., KÖHLER, L., FRUMAU, K. F. A., BRUIJNZEEL, L. A., BURKARD, R. & SCHMID, S. 2010. Water dynamics of epiphytic vegetation in a lower montane cloud forest: fog interception, storage, and evaporation. Pp. 261267 in Bruijnzeel, L. A., Scatena, F. N. & Hamilton, L. S. (eds.). Tropical montane cloud forests: science for conservation and management. Cambridge University Press, Cambridge.
VANCE, E. & NADKARNI, N. 1990. Microbial biomass and activity in canopy organic matter and the forest floor of a tropical cloud forest. Soil Biology and Biochemistry 22:677684.
VANCE, E. & NADKARNI, N. 1992. Root biomass distribution in a moist tropical montane forest. Plant and Soil 142:3139.
VARESCHI, V. 1986. Cino breves ensayos ecologicos acerca de la selva virgen de Rancho Grande. Pp. 171187 in Huber, O. (ed.). La selva nublada de Rancho Grande parque nacional “Henri Pittier”. Fondo editorial Acta Cientifica Venezoalana, Seguros Anauco C.A., Caracas.
VENEKLAAS, E. 1990. Nutrient fluxes in bulk precipitation and throughfall in two montane tropical rain forests, Colombia. Journal of Ecology 78:974992.
VENEKLAAS, E. 1992. Litterfall and nutrient fluxes in two montane tropical rainforests, Colombia. Journal of Tropical Ecology 7:319336.
WAGNER, S., BADER, M. Y. & ZOTZ, G. 2014. Physiological ecology of tropical bryophytes. photosynthesis in bryophytes and early land plants. Advances in Photosynthesis and Respiration 37:269289.
WEAVER, P. L. 1972. Cloud moisture interception in the Luquillo Mountains of Puerto Rico. Caribbean Journal of Science 12:129144.
WERNER, F. A., HOMEIER, J., OESKER, M. & BOY, J. 2012. Epiphytic biomass of a tropical montane forest varies with topography. Journal of Tropical Ecology 28:2331.
WILSON, E. O. 1988. Biodiversity. Harvard University Press, Cambridge. 538 pp.
ZOTZ, G. & HIETZ, P. 2001. The physiological ecology of vascular epiphytes: current knowledge, open questions. Journal of Experimental Botany 364:20642078.
ZOTZ, G., BUDEL, B., MEYER, A., ZELLNER, H. & LANGE, O. L. 1997. Water relations and CO2 exchange of tropical bryophytes in a lower montane rain forest in Panama. Botanica Acta 110:917.
Recommend this journal

Email your librarian or administrator to recommend adding this journal to your organisation's collection.

Journal of Tropical Ecology
  • ISSN: 0266-4674
  • EISSN: 1469-7831
  • URL: /core/journals/journal-of-tropical-ecology
Please enter your name
Please enter a valid email address
Who would you like to send this to? *



Full text views

Total number of HTML views: 30
Total number of PDF views: 260 *
Loading metrics...

Abstract views

Total abstract views: 870 *
Loading metrics...

* Views captured on Cambridge Core between September 2016 - 17th July 2018. This data will be updated every 24 hours.