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In these essays that survey the burgeoning field of tropical herpetology, former students and associates pay tribute to Jay Savage's four decades of mentoring. The result is a book unlike any other available in tropical herpetology. Covering a wide array of subjects, Ecology and Evolution in the Tropics is the first book in more than two decades to broadly review research on tropical amphibians and reptiles. A tribute to Savage and an invaluable addition to the herpetological literature, this work will be cited for years to come.

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Ecology & Evolution in the Tropics

The University of Chicago Press

Contents

Foreword by Luis D. Gmez........................................................................................................................................................................xiPreface..........................................................................................................................................................................................xiiiPart I: Evolution and Biogeography1. Taxonomy in Theory and Practice, with Arguments for a New Phylogenetic System of Taxonomy Arnold G. Kluge....................................................................................72. Biogeography and Molecular Phylogeny of Certain New World Caecilians Marvalee H. Wake, Gabriela Parra-Olea, and Judy P.-Y. Sheen.............................................................483. Diversity of Costa Rican Salamanders David B. Wake...........................................................................................................................................654. On the Enigmatic Distribution of the Honduran Endemic Leptodactylus silvanimbus (Amphibia: Anura: Leptodactylidae) W. Ronald Heyer, Rafael O. de S, and Sarah Muller.........................815. Chromosomal Variation in the rhodopis Group of the Southern Central American Eleutherodactyline Frogs (Leptodactylidae: Eleutherodactylus) Shyh-Hwang Chen...................................1026. The Physiological Basis of Sexual Dimorphism: Theoretical Implications for Evolutionary Patterns of Secondary Sexual Characteristics in Tropical Frogs Sharon B. Emerson.....................1457. Higher-Level Snake Phylogeny as Inferred from 28S Ribosomal DNA and Morphology Mary E. White, Maria Kelly-Smith, and Brian I. Crother.........................................................1568. Elapid Relationships Joseph B. Slowinski and Robin Lawson....................................................................................................................................1749. Wallace and Savage: Heroes, Theories, and Venomous Snake Mimicry Harry W. Greene and Roy W. McDiarmid.........................................................................................190Part II: Ecology, Biogeography, and Faunal Studies10. Quantification of Selection and Male Reproductive Success in Hyla calypsa, a Neotropical Treefrog Karen R. Lips.............................................................................21511. Patterns of Co-occurrence of Hylid Frogs at a Temporary Wetland in Costa Rica Craig Guyer and Maureen A. Donnelly...........................................................................22712. It's a Frog-Eat-Frog World in the Paraguayan Chaco: Food Habits, Anatomy, and Behavior of the Frog-Eating Anurans Norman J. Scott Jr. and A. Luz Aquino......................................24313. Long-Term Frog Monitoring by Local People in Papua New Guinea and the 1997-98 El Nio Southern Oscillation David P. Bickford................................................................26014. Historical Biogeographic Relationships within the Tropical Lizard Genus Norops Kirsten E. Nicholson.........................................................................................28415. Hypotheses on the Historical Biogeography of Bothropoid Pitvipers and Related Genera of the Neotropics Steven D. Werman......................................................................30616. The Herpetofauna of the Rincn Area, Pennsula de Osa, Costa Rica, a Central American Lowland Evergreen Forest Site Roy W. McDiarmid and Jay M. Savage.......................................36617. The Iwokrama Herpetofauna: An Exploration of Diversity in a Guyanan Rainforest Maureen A. Donnelly, Megan H. Chen, and Graham G. Watkins.....................................................42818. The Herpetofauna of the Guayana Highlands: Amphibians and Reptiles of the Lost World Roy W. McDiarmid and Maureen A. Donnelly................................................................461References.......................................................................................................................................................................................561Contributors.....................................................................................................................................................................................627Subject Index....................................................................................................................................................................................631Taxonomic Index..................................................................................................................................................................................655

Chapter One

Arnold G. Kluge

Theory without fact is fantasy, but facts without theory is chaos. CHARLES OTIS WHITMAN

Taxonomy as it is currently practiced is nearly 250 years old, and it is only now undergoing its first major revolution (Pennisi 1996). There is widespread agreement that categorical ranks must go, and with this change alone there is repudiation of what has been to date the venerable Linnaean approach to classification (Linnaeus 1758; see review in Ereshefsky 2001) as well as the rejection of the well-entrenched International Codes of Nomenclature (ICBN 1994; ICNB 1992; ICZN 1999). Making explicit connections to evolutionary theory and phylogenetic hypotheses is also considered important to the modernization of taxonomy. The issues underlying these attacks on the Linnaean tradition are especially complex because they involve the whole of science, from the philosophical to the pragmatic (Ereshefsky 2001). My attempt to understand the theory and practice of taxonomy, including the current revolution, is founded largely on the distinction between classification and systematization (Griffiths 1974; see also Griffiths 1973, 1976). Most of the discussion to follow explores the role science plays in systematization, and with that understanding I propose and exemplify a new phylogenetic system of taxonomy.

Definitions and Distinctions

There are several different kinds of definitions, which unfortunately are conflated in ordinary discourse, and whose record of use in science is not much better. Particularly important terms in distinguishing the classification of Linnaeus from the systematization of G. C. D. Griffiths are extension, intension, and ostension (table 1.1). Intension is used to define universals. Ostension and extension are used to define particulars. Extensional definitions, however, may look like intensional definitions, thus opening the door for confusion such that particulars seem to be defined in universal terms. Also potentially confusing are those things that may be judged abstract (table 1.1), such as the Equator and the North and South Poles. Suffice it to say that these kinds of particulars are not so abstract as to keep us from knowing when we have crossed the Equator or have arrived at either of the Poles.

Intension (connotation) is the principle according to which a class, or set, of things is picked out; it is the condition a thing must satisfy to be precisely described by the predicate (Blackburn 1994). Intension involves a defining rule whereby members are included, or excluded, from the set. The...