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Copyright by Rafe Marion Brown 2004 Copyright By Rafe Marion Brown 2004 The dissertation committee for Rafe Marion Brown certifies that this is the approved version of the following dissertation: Evolution of Ecomorphological Variation and Acoustic Diversity in Mate-Recognition Signals of Southeast Asian Forest Frogs (subfamily Platymantinae) Committee: ____________________________________ David Cannatella, Supervisor ____________________________________ David Hillis, Co-Supervisor ____________________________________ Michael Ryan ____________________________________ Walter Wilczynski ____________________________________ Robert Dudley Evolution of Ecomorphological Variation and Acoustic Diversity in Mate-Recognition Signals of Southeast Asian Forest Frogs (subfamily Platymantinae) by Rafe Marion Brown, B.A.; M.Sc. Dissertation Presented to the Faculty of the Graduate School of The University of Texas at Austin in Partial Fulfillment for the Degree of Doctor of Philosophy The University of Texas at Austin May 2004 Dedication This one goes out to all my homies, the fallen rappers—cut down in the streets, taken out in the drive-bys, ganked by their own brothers, beat down by The Man. Acknowledgements A great number of people have assisted with logistical and material support of my dissertation work. First, I thank the Protected Areas and Wildlife Bureau of the Philippine Department of Environment and Natural Resources for facilitating collecting and export permits and the following DENR officers for their efforts on my behalf: J. DeLeone, M. Mendoza, A. Lota, M. Lim, M. Maranan, L. Bacosa, J. Alisuag, A. Sandalo, R. Villegas, and W. Pollisco, and especially A. Tagtag and C. Custodio. While I was developing ideas in pursuit of a dissertation project, I benefittted immensely from consversations with W. Brown, A. Alcala, A. Diesmos, D. Cannatella, L. Heaney, and J. McGuire. The logistical support of of A. Alcala, D. Felicitas, R. Flores (Silliman University), and R. Sison, (National Museum of the Philippines) is greatly appreciated. In Manila I was hosted, entertained, and put back on track many times by A. Diesmos and his family, and for their support I will always be grateful. The dedicated field assistance of V. Yngente and R. Fernandez is greatly appreciated, as is the assistance of the provincial DENR authorities, too numerous to name here. Thanks are due to W. Olliver, L. Dans, and J. C. Gonzales for their kindness in letting me wisk Enteng away from his post on Polillo. I thank N. Mallari, R. Fernandez, J. Redor, C. Torno, A. C. and M. L. Diesmos, Kabanahaw, Inc., and Tayabas Mountaineers for help on Mt. Banahao. For assistance in Sorsogon I thank Bulusan Natural Park PASu L. Soriano, and for help in Mt. Isarog National Park PASu’s C. Rivero and park guard R. Buenviaje. I thank the PAMB boards of both National Parks v for their support of my research. I thank C. N. Dolino (Dumaguete) for guidance and logistical help in accessing Mt. Talinis, Cuernos de Negros; I also thank the NAPCOR power company of southern Negros and Malinao, Albay for access to their private land. I thank the Subic Bay Metropolitan Area Ecology Center for logistical support and assistance while working in Olongapo and Subic Bay and the University of the Philippines at Los Baños for supporting work on Mt. Makiling. I am especially grateful to L. Heaney for many helpful conversations, thoughtful guidance, invaluable advice, and for taking me to the Cordillera; fieldwork in Balbalasang was made possible through the cooperation of Haribon Foundation, the Field Museum of Natural History, the people of Balbalasang, and my friends G. Gee, D. Balete, B. Tabaranza, R. Fernandez, N. Malliari, and PASu C. Aquino. I thank R. Crombie, A. Alcala, A. Diesmos, S. Richards and the late W. Brown for expanding my understanding of platymantine frog taxonomy and J. McGuire and D. Zwickl for many helpful discussions about phylogenetics. For the generous loans of specimens and provision of working space during museum visits I thank J. Vindum (CAS), R. Crombie, K. de Queiroz, G. Zug (USNM), R. Sison (PNM), A. Resetar, H. Voris, R. Inger (FMNH), J. Rosales, and D. Cannatella (TNHC). Financial support for visits to CAS was generously provided by the Charles Stearns Fellowship. Financial support for fieldwork was provided by the Society of Systematic Biologists, the US National Science Foundation (Dissertation Improvement Grant to myself and NSF 9981631 and 0206729 to D. Cannatella, M. Ryan, D. Hillis, and R. Dudley), the Society for the Study of Amphibians and Reptiles, the American Society vi of Ichthyologists and Herpetologists, The Explorers Club, the Section of Integrative Biology and Texas Memorial Museum of the University of Texas at Austin. I am intensely grateful to Jen Weghorst for her help throughout: her untiring assistance with digitizing frog calls, her enthusiastic help scanning slides, the many days she spent working on my research rather than her own, the monumental task of proofreading multiple copies of all chapters, and for providing the support and motivation to follow through and finish up. Lastly, I thank my good friend Arvin Diesmos for paving the way before and during all phases of my Philippine field work, for keeping me out of trouble in Manila, for protecting me from many dangers (including myself) during my travels, for his patience with my eccentricities, and for always being willing to stay up late to catch frogs. vii Evolution of Ecomorphological Variation and Acoustic Diversity in Mate-Recognition Signals of Southeast Asian Forest Frogs (subfamily Platymantinae) Publication No._______ Rafe Marion Brown, Ph.D. The University of Texas at Austin, 2004 Supervisors: David Cannatella and David Hillis I estimated evolutionary relationships of frogs of the subfamily Platymantinae (genera Platymantis, Batrachylodes, Palmatorappia, Discodeles, Ceratobatrachus) and their relatives (genus Ingerana and various SE Asian ranids) using mitochondrial gene sequences. Various methods of phylogenetic inference all suggested that platymantines consist of two reciprocally monophyletic clades, one composed of Philippine Platymantis and the other containing all Papuan-Solomon-Bismarck archipelago taxa. Non- Platymantis genera are nested within Platymantis and fall basal to SW Pacific island archipelago species. I recognized 51 evolutionary lineages of Philippine platymantines and attempted to statistically define morphological, ecological, and acoustic classes of species. viii Multivatiate analyses of continuous morphological characters suggested that species diversity falls into five classes: (1) canopy frogs, (2) shrub frogs, (3) ground frogs, (4) giants, and (5) miniaturized species. Analyses of 10 acoustic characters similarly distinguished five call types: (1) simple single pulse (“tink”) calls, (2) pure tone calls, (3) frequency sweeps, (4) amplitude-modulated pulsed calls, and (5) complex calls. Although the correlation of morphological and call types was not perfect, their general association with each other and with specific microhabitat preferences suggests selection for suites of associated morphological, ecological, and behavioral traits. I also tested the Acoustic Adaptation Hypothesis (AAH) and the prediction that species should produce acoustic signals in the environments where those calls transmit best. I used playback experiments in which I broadcasted calls into different forest types, re-recorded calls along distance transects and statistically compared these experimentally degraded calls to non-degraded exemplars. I found moderate support for the predictions of the AAH and the general prediction that species may evolve advertisement calls that maximize transmission efficacy in preferred microhabitats. Finally, I mapped call characters on the phylogeny and asked whether “morphological” call characters (e.g., dominant frequency) and “behavioral” call characters (e.g., call rate) have evolved at different rates. Results suggested that some characters evolved more rapidly than others, but there was no clear-cut distinction between morphological and behavioral classes. Nevertheless, patterns of repeated evolution of call types and differential performance of calls in various environments ix suggest repeated evolutionary convergence on morphological, ecological, and behavioral traits across replicated radiations of platymantine frogs. x Table of Contents Chapter 1: Cryptic species diversity in a large oceanic island frog radiation: partial taxonomic review of Philippine forest frogs of the genus Platymantis (Amphibia: Ranidae) …………………………………………………………….1 Chapter 2: Ecological morphology of platymantine frogs (Ranidae) of SE Asia and the SW Pacific……………………………………………………………………..104 Chapter 3: Characterization of advertisement call variation in platymantine ranid frogs of the Philippines……………………………………………..……………122 Chapter 4: Phylogenetic systematics and biogeography of platymantine ranid frogs of SE Asia and the SW Pacific ……….………………………………………….132 Chapter 5: Transmission performance of advertisement calls of Philippine platymantine frogs in complex forest environments …………...…...…………………170 Chapter 6: A comparative analysis of mate recognition signals: platymantine ranids, call character diversity, and the tempo of advertisement callevolution…..……197 Tables…………………………………………………………………………………..225 Figures………………………………………………………………………………….234 Appendix……………………………………………………………………………….290 Bibliography……………………………………………………………………………297 Vita………………………………………………..……………………………………352 xi Chapter 1: Cryptic species
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