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The Ecology of Vascular Epiphytes on a Ficus L. Host (Moraceae) in a Peruvian Cloud Forest

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The Ecology of Vascular Epiphytes on a Ficus L. Host (Moraceae) in a Peruvian Cloud Forest The ecology of vascular epiphytes on a Ficus L. host (Moraceae) in a Peruvian cloud forest by Damien Catchpole BSc. (University of Tasmania) A thesis submitted in partial fulfilment of the requirements for an Honours Degree at the School of Geography and Environmental Studies, University of Tasmania (June, 2004). Declaration This thesis contains no material which has been accepted for the award of any other degree or diploma in any tertiary institution, and to the best of my knowledge and belief, contains no material previously published or written by another person, except where due reference is made in the text of the thesis. Signed Damien Catchpole BSc. ii Abstract A total of 190 holoepiphytic and 5 hemiepiphytic vascular plant species were collected from the canopy and the trunk of an emergent Ficus L. species host that is common to a Peruvian cloud forest. One hundred and fourteen of the vascular epiphyte species were orchids. A large majority of the vascular species were rare in occurrence. Vascular epiphyte diversity and density was highest in the outer canopy zone of the host crown. In the inner canopy zone there was a dearth of epiphytes, attributed to a high rate of epiphyte slumping on the smooth-barked branches. The trunk had a different suit of species to those found in the canopy. During the wet season, the thickening of epiphytic matter in the outer canopy zone appeared to contribute to a large amelioration of daytime and nocturnal temperatures through evaporation and heat retention respectively. Nocturnal temperatures were highest in the outer canopy zone, and were lowest on the forest floor. The latter was attributed to the downward flow of the product of radiative cooling as a result of the heterogeneous canopy on the steep slope. Epiphyte clumps appear to slump before competition causes the loss of early successional species. Some species showed a preference for more shaded epiphyte clumps and many were more frequent on smaller branch diameters. Most species showed a moderately high niche overlap with a large number of other species, which suggested a high degree of species coexistence. Epiphyte slumping is suggested to be the major driving mechanism for the maintenance of non-equilibrium in the community. Aspects of the phenologies of epiphytes, age-structure of the community and high environmental variation could be other mechanisms for the maintenance of a high degree of species coexistence. iii Acknowledgements This research was partly funded by the Andrew J. Mellon Foundation, the Australian Ecological Society and the Walter Guinness Charitable Trust. Generous donations of equipment were received from Macpac. The loan of a motorcycle during the study was given without charge by Benjamin Kroll of the Fundación Peruana para la Conservación de la Naturaleza (Pro Naturaleza). Logistical support was received from the Andean Amazon Research Station, the Missouri Botanical Garden, and the Universidad Nacional Agraria La Molina. I would like to thank Yoshie Yoshioka for her support, and for assistance with fieldwork and herbarium specimens. Field assistance was received from Guido Casimiro, Hana Eynon, Daniel Gómez, Benjamin Jones and Gaby Rivera. I would like to thank Rodolfo Vásquez, Rocío Rojas and Abel Monteagudo of the Missouri Botanical Garden for their friendly support and advice with all aspects of the plant collection. Assistance with plant identification was received from Rodolfo Vásquez, Luis Valenzuela (Orchidaceae), Jorge Lingan (Araceae) and Franco Mellado (Pteridophyta). I am indebted to my supervisor Prof Jamie Kirkpatrick for his support, advice and assistance, and for his initial support of my ambitious proposal as an honours project. I must also thank Assoc/Prof Manuel Nunez and Dr Emma Pharo for their much needed advice and assistance in many aspects of the project. Darren Turner was invaluable for data management and performed the masking of all hemispherical images. I thank Pedro Aguilar for his friendship and general helpfulness, and, Assoc/Prof Carlos Llerena, Assoc/Prof Michael McClain and Billy Huggard for facilitating institutional cooperation. Sonya Stallbaum helped with manuscript reviews. A special thanks goes to Yoav Barness who donated his time to teach me the rope techniques and procedures for canopy access. My deepest thanks are reserved for my family who have tolerated my selfish tunnel vision during the project, lessened the heavy financial burden and remained supportive throughout. iv Table of Contents List of Figures............................................................................viii List of Tables ...............................................................................xi Chapter 1 Introduction.............................................................1 1.1 Background......................................................................................1 1.2 Research Aims.................................................................................5 1.3 Structure of thesis...........................................................................5 Chapter 2 Methods ...................................................................7 2.1 Study Area........................................................................................7 2.2 Study Design & Field Data Collection..........................................10 2.2.1 Canopy Sampling ............................................................................ 10 2.2.2 Plant Identification .......................................................................... 11 2.2.3 Lifeform and Taxonomic Groups ................................................... 13 2.2.4 Environmental Data......................................................................... 14 2.3 PAR & Canopy Structure ..............................................................15 2.3.1 Output Descriptions ........................................................................ 17 Chapter 3 Structure and Composition..................................19 3.1 Aims................................................................................................19 3.2 Methods..........................................................................................19 3.3 Results............................................................................................20 3.3.1 Floristic Composition...................................................................... 20 3.3.2 Community Structure...................................................................... 22 3.3.3 Epiphyte Clumps ............................................................................. 24 3.4 Discussion .....................................................................................27 3.4.1 Floristic Composition...................................................................... 27 3.4.2 Community Structure...................................................................... 28 3.4.3 Epiphyte Clumps ............................................................................. 30 3.4.4 Community Comparison................................................................. 32 3.5 Conclusion .....................................................................................35 Chapter 4 Habitat Characteristics and Distribution ............36 4.1 Aims................................................................................................36 4.2 Methods..........................................................................................37 4.2.1 Environmental Habitat Variables ................................................... 37 4.2.2 Epiphyte Distribution by Environmental Habitat Variables......... 37 4.2.3 Epiphyte Distribution by Johansson Zones ................................. 38 4.2.4 Floristic Ordination ......................................................................... 39 4.3 Results............................................................................................40 4.3.1 Environmental Habitat Variables ................................................... 40 4.3.2 Epiphyte Distribution ...................................................................... 42 v 4.3.3 Johansson Zones ............................................................................ 47 4.3.3.1 Community Structure ............................................................................. 47 4.3.3.2 Clump and Environmental Habitat Variables ....................................... 49 4.3.3.3 Floristics .................................................................................................. 52 4.3.4 Floristic Ordination ......................................................................... 53 4.4 Discussion .....................................................................................54 4.4.1 Environmental Habitat Variables ................................................... 54 4.4.2 Epiphyte Distribution ...................................................................... 56 4.4.3 Johansson Zones ............................................................................ 59 4.4.3.1 Community Structure ............................................................................. 59 4.4.3.2 Clump and Environmental Habitat Variables ....................................... 60 4.4.3.3 Floristics .................................................................................................. 63 4.4.4 Floristic Ordination ......................................................................... 66 4.5 Conclusion .....................................................................................68
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