Bulletin of the Biological Society of Washington No. 2 THE PANAMIC BIOTA: SOME OBSERVATIONS PRIOR TO A SEA-LEVEL CANAL A Symposium Sponsored by The Biological Society of Washington The Conservation Foundation The National Museum of Natural History The Smithsonian Institution MEREDITH L. JONES, Editor September 28, 1972 CONTENTS Foreword The Editor - - - - - - - - - - Introduction Meredith L. Jones ____________ vi A Tribute to Waldo Lasalle Schmitt George A. Llano 1 Background for a New, Sea-Level, Panama Canal David Challinor - - - - - - - - - - - Observations on the Ecology of the Caribbean and Pacific Coasts of Panama - - - - Peter W. Glynn _ 13 Physical Characteristics of the Proposed Sea-Level Isthmian Canal John P. Sheffey - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 31 Exchange of Water through the Proposed Sea-Level Canal at Panama Donald R. F. Harleman - - - - - - - - - - - - - - - - - - - - - - - - - - - 41 Biological Results of the University of Miami Deep-Sea Expeditions. 93. Comments Concerning the University of Miami's Marine Biological Survey Related to the Panamanian Sea-Level Canal Gilbert L. Voss - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 49 Museums as Environmental Data Banks: Curatorial Problems Posed by an Extensive Biological Survey Richard S. Cowan - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 59 A Review of the Marine Plants of Panama Sylvia A. Earle - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 69 Ecology and Species Diversity of Coral Reefs on Opposite Sides of the Isthmus of Panama James W. Porter - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 89 Origin of the Existing Panamic Molluscan Biotas in Terms of Their Geologic History and Their Separation by the Isthmian Land Barrier Axel A. Olsson - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 117 Comparative Habitat Diversity and Faunal Relationships Between the Pacific and Caribbean Panamanian Decapod Crustacea: A Preliminary Report, with Some Remarks on the Crustacean Fauna of Panama Lawrence G. Abele 125 The Status of Knowledge of Panamanian Echinoids, 1971, with Comments on Other Echinoderms Richard H. Chesher 139 The State of Knowledge of the Coastal Fish Fauna of the Panamic Region Prior to the Construction of an Interoceanic Sea-Level Canal C. Richard Robins 159 The Marine Fisheries of Panama Harvey R. Bullis and Edward F. Klima 167 Terrestrial Plants of Panama Robert L. Dressler 179 The Entomology of Panama Neal A. Weber 187 The Status of Herpetology in Panama Charles W. Myers 199 The Birds of the Isthmus of Panama Alexander Wetmore 211 Mammalogy in Panama Charles 0. Handley, Jr. 217 The Anthropology of Eastern Panama Reina Torres de Arafiz 229 The National Academy of Science Committee on the Ecology of the Interoceanic Canal William A. Newman 247 Summary Daniel M. Cohen 261 Appendix 1. Pelamis platurus as a Potential Colonizer of the Caribbean Sea Chaim Kropach 267 FOREWORD The Biological Society of Washington is pleased to present the papers in this volume as a contribution toward the dissemination of fundamental knowledge of a controversial topic. The Society, as well as the other sponsoring groups and the Editor, must disclaim responsibility for the statements included herein. Each author is responsible for the content of his own paper. As Editor, I have been confronted, as all editors are, with as many individual writing styles as there are contributors and as many formats as there are disciplines. I have tried to edit with as light a hand and as pale a blue pencil as I felt possible. This was done in the interests of preserving individual style, discipline format, and the passions of the moment of the several authors who have shown their sincere and profound feelings concerning the sea-level canal. It should be noted that the term "Panamic", as used in the follow- ing papers, refers to the Panamanian region, in general, and not to the zoogeographic Panamic Province, unless otherwise indicated. As Convenor and Editor of the Symposium I am most grateful to the Conservation Foundation, Washington, D. C., and especially to Mr. John P. Milton, for defraying publication costs, to the Na- tional Museum of Natural History, Smithsonian Institution, for providing funds for the travel and expenses of the participants from beyond the Washington area, and to the Council of the Biological Society of Washington for permission to resurrect the Society's Bulletin series for the Symposium and for permission to seek travel and publication funds for the Symposium, as I could find them. Finally, I would thank Teresa Smith, Division of Worms, National Museum of Natural History, Smithsonian Institution, for cheerfully and competently putting up with the inherent grouchiness and perpetual demands for retyping which characterize an editor. MEREDITH L. JONES INTRODUCTION BY MEREDITH L. JONES National Museum of Natural History Smithsonian Institution, Washington, D. C. and Past President (1970), the Biological Society of Washington As a point of departure for this Symposium, it might be well to restate a few fundamental facts concerning Central America and the Panamanian isthmus. 1. Until the opening of the present canal in 1914, the Central American land mass constituted a barrier to the organisms of the Pacific and the Caribbean, and a land bridge for terrestrial organisms. 2. The marine species had been separated, and the ter- restrial species had been linked, for from one to five million years; in the case of the marine faunas and floras, there were classical textbook opportunities for the formation of geminate species through independent evolution. 3. With the building of the present canal, a fresh water barrier was formed with respect to the terrestrial forms, and a barrier, only slightly less formidable than the previous one, was established for the marine forms. 4. Presently, the building of a new canal, a sea-level canal, has been proposed. A number of rather basic questions are suggested by these facts: 1. In the case of geminate species, how far have they diverged? What are their morphological, their physiological, and their biochemical differences—in short, what are the genetic differences between species pairs? 2. How do the present faunas and floras, the marine, the terrestrial, and those of the northern and southern fresh water habitats, differ? vi 3. How much faunal and floral mixing of marine species has taken place? Is this due to shipping, as fouling organisms or as ballast-water survivors? Or is it due to normal dispersion? 4. What might be the effect of a sea-level canal on the Panamic biota—both as a continuum of salt water from ocean to ocean and as a salt water barrier to terrestrial and fresh water forms? All of these, as well as the many more which undoubtedly come to mind, are interesting, intriguing, and important, insofar as distributional, ecological, physiological, and evolu- tionary studies are concerned. However, the answers to all of these questions, and the foundation for all of these and other studies, no matter how far removed, ultimately devolve from one basic premise: To evaluate differences, to justify comparisons, to analyze ob- servations, one must have a valid point of departure. In order to compare floras and faunas, one must know what spe- cies are present. In order to study similarities and differences between geminate species, one must recognize their existence. In order to assess the role of ships in the dispersal of species, one must know the fouling organisms on the ships' hulls, as well as the potential fouling organisms of both sides of the isthmus, and one must know what organisms, if any, can survive a ballast- water environment. In order to hypothesize concerning the effects of a sea-level canal on the Panamic biota, even assum- ing that the hydrologic and hydrographic parameters are unequivocally known, the faunas and floras of both oceans, as well as the northern and southern terrestrial and fresh water floras and faunas, must be known. In short, we can not say what differences exist now, we can not access the independent evolution of geminate species, we can not comment on the amount of mixing between areas, we can not evaluate the effect of shipping, and we can not theorize about the effects of a sea-level canal, unless we have a basic knowledge of the organisms. As a first step toward this end, we are convened here to sum- marize the present knowledge of the Panamic biota. The importance of base line collections and observations, such as are called for here, long have been recognized, whether vii it was by Verrill in New England, by Wyville Thompson on the CHALLENGER, or by Agassiz on the BLAKE and ALBA- TROSS, to name but a few of the pioneers. There is such a man with us today; one who worked on then-recently-collected AL- BATROSS material in the early 1900's and was senior biologist on a cruise to the Antarctic forty years later—a man dedicated to base line collecting, to the working up of such collections, and to the zoogeographic analysis of the data—a man to whom we pay tribute today, albeit inadequately relative to the leader- ship he has shown, the inspiration he has given, and the contributions he has made to marine biology and marine biologists. Our next speaker will speak in detail of the contri- butions of Waldo Lasalle Schmitt. viii A TRIBUTE TO WALDO LASALLE SCHMITT BY GEORGE A. LLANO Acting Chief Scientist, Office of Polar Programs National Science Foundation, Washington, D. C. There are many present here who have known Waldo L. Schmitt longer than I; or who, during his fifty years of service with the Smithsonian Institution, have been professionally more closely associated with him. As a matter of record I have come to know Waldo best since 1959, after he retired; and, as a consequence, I can best address myself to those quiet years which, theoretically, come with retirement. I am most appreciative of being given the opportunity of rendering this small tribute to one who, by his name and through his works, has identified himself with this Institution and who, as a civil servant, has served his government in full measure.