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Variation Within the Leptodactylus Podicipinus-Wagneri Complex of Frogs (Amphibia: Leptodactylidae)

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Variation Within the Leptodactylus Podicipinus-Wagneri Complex of Frogs (Amphibia: Leptodactylidae) Variation within the Leptodactylus podicipinus-wagneri Complex of Frogs (Amphibia: Leptodactylidae) W. RONALD HEYER SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 546 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs'submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 546 Variation within the Leptodactylus podicipinus-wagneri Complex of Frogs (Amphibia: Leptodactylidae) W. Ronald Heyer SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1994 ABSTRACT Heyer, W. Ronald. Variation within the Leptodactylus podicipinus-wagneri Complex of Frogs (Amphibia: Leptodactylidae). Smithsonian Contributions to Zoology, number 546, 124 pages, 46 figures, 55 tables, 1994.—Variation was studied in frogs identified as either Leptodactylus podicipinus or L. wagneri as diagnosed in my previous revision of the L. melanonotus species group. Over 6200 adults and juveniles were examined and variation analyses were performed on data from just over 3000 adult and near-adult specimens. The data set consists of a series of three kinds of morphological characters (patterns, structures, measurements), and, when available, advertisement call and habitat data. Intrapopulation variation indicates that variation among the characters studied is extensive within large samples. The intrapopulation variation results are used as a framework to interpret interpopulation and interspecific variation. Study of character variation among sympatric populations typically leads to the following results: (1) when compared side by side, most specimens can be sorted readily into distinct morphotypes representing distinct species; and (2) the variation between sympatric populations is usually distinctive, but not discrete. Advertisement calls in members of this complex appear to contain not only species-coding information, but other information that weakens the utility of calls in assessing species boundaries. The results of this study are used to delimit species for most of the specimens examined. Available material from most of Venezuela is inadequate to evaluate how many species occur there and which of them are conspecific with geographically adjacent species. Thirteen species are diagnosed as a result of this study, including the description of five new species. The recognition of 13 species is conservative and additional information likely will confirm that some of these species are composite. The distributions of most taxa within this complex are expected to be modified significantly by either newly collected specimens or data that will redefine currently recognized (composite) species. Only two distributions are considered robust as the result of this study, those of L. natalensis and L. podicipinus. The provenance of museum specimens collected by Borys Malkin purportedly from Igarape Bele"m, Amazonas, Brazil, is called into question. Unresolved problems are highlighted to encourage and focus further studies to understand the speciation processes and distribution patterns in this complex. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Heyer, W. Ronald Variation within the Leptodactylus podicipinus-wagneri complex of frogs (Amphibia : Leptodactylidae) / W. Ronald Heyer. p. cm. — (Smithsonian contributions to zoology ; no. 546) Includes bibliographical references. I. Leptodactylus—Classification. 2. Leptodactylus—Variation. I. Title. II. Title: Leptodactylus podicipinus- wagneri complex of frogs. III. Series. QL1.S54 no. 546 [QL668.E257] 591 s-dc20 [597.8'9] 93-6156 ® The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984. Contents Page Introduction 1 Acknowledgments 2 Methods and Materials 2 Analysis of Single-Taxon Populations 3 Individual OTUs 4 Small Size, Dark Belly, Porto Velho, Brazil OTU 4 Moderate Size, Light Posterior Belly, Limoncocha, Ecuador OTU 4 Small Size, Dark Belly, Rurrenabaque, Bolivia OTU 5 Small/Moderate Size, Dark Belly, Alejandria, Bolivia OTU 5 Small/Moderate Size, Light Thigh Stripe, Buenavista, Bolivia OTU 6 Small Size, Light Posterior Lip Stripe, Kartabo, Guyana OTU 7 Large Size, Boldly Mottled Belly, Santa Cecilia, Ecuador OTU 7 Small Size, Dark Belly, Curuca\ Brazil OTU 8 Small Size, Light Posterior Lip Stripe, Langaman Kondre, Surinam OTU .... 9 Discussion 10 Analysis of Sympatric Species Populations 12 Peru, Madre de Dios, Tambopata OTUs 12 Peru, Madre de Dios, Cuzco Amazonico OTUs 14 Peru, Loreto, Estir6n, Rio Apiyacu OTUs 14 Peru, Huanuco, Divisoria OTUs 16 Bolivia, Beni, Tumi Chucua OTUs 16 Ecuador, Napo, Santa Cecilia OTUs 18 Brazil, Acre, Iquiri OTUs 18 Brazil, Rondonia, Santa Cruz da Serra OTUs 18 Brazil, Amazonas, Boca do Acre OTUs 19 Brazil, Amazonas, Borba OTUs 19 Brazil, Amazonas, Igarapd Bele"m, Rio Solimoes OTUs 19 Brazil, Amazonas, Lago Amana OTUs 21 Brazil, Par£, Cachoeira do Espelho, Rio Xingu OTUs 21 Surinam, Marowijne, Loekreek OTUs 21 Surinam, Nickerie, Amotopo OTUs 21 Surinam, Marowijne, Paloemeu OTUs 22 Guyana, Kartabo OTUs 22 Colombia, Amazonas, Leticia OTUs 22 Colombia, Amazonas, Quebrada Tucuchira OTUs 22 Variation within Taxa/Regions 23 Distinct Taxa 23 Core Region Analyses 24 Region 1—South 24 Region 2—East Coast Brazil 25 Region 3—Interior Brazil 26 Region 4—Amazonia 32 Region 5—Guiana Shield 45 Region 6—Trinidad, Tobago, Lesser Antilles 56 Region 7—Andean Slopes 62 Interregional Synthesis 72 ui iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY Comparison of Certain Colombian and Venezuelan OTUs 74 Region 3 Light Belly OTU 76 Island OTUs 77 Summary and Identification of Specimens Previously Not Assigned to a Region 77 Nomenclature 77 Species Accounts 82 Leptodactylus colombiensis, new species 82 Leptodactylus diedrus, new species 86 Leptodactylus griseigularis (Henle, 1981) 87 Leptodactylus leptodactyloides (Andersson, 1945) 88 Leptodactylus natalensis Lutz, 1930 89 Leptodactylus nesiotus, new species 91 Leptodactylus pallidirostris Lutz, 1930 93 Leptodactylus pascoensis, new species 94 Leptodactylus petersii (Steindachner, 1864) 96 Leptodactylus podicipinus (Cope, 1862) 97 Leptodactylus sabanensis, new species 99 Leptodactylus validus Garman, 1887 102 Leptodactylus wagneri (Peters, 1862) 104 Distributions 105 Differentiation 106 Comment on Relationships 108 Unresolved Problems and Their Consequences in Understanding Evolutionary Processes 108 Appendix 1: OTU (Operational Taxonomic Unit) Assignments 110 Appendix 2: Specimens Examined and Locality Data 113 Literature Cited 123 Variation within the Leptodactylus podicipinus-wagneri Complex of Frogs (Amphibia: Leptodactylidae) W. Ronald Heyer Introduction First, the study materials have increased in two critical ways. Now there are some large single-taxa samples from specific In 1970,1 published the results of a species-level analysis of localities so that intrapopulation variation can be characterized frogs of the Leptodactylus melanonotus species group (Heyer, much more adequately. There also are many more localities 1970), which included the taxa that are the topic of this paper. from which more than a single taxon of this complex have been In that paper, six species were recognized: Leptodactylus collected. In the previous study only two such localities were dantasi, discodactylus, melanonotus, podicipinus, pustulatus, identified. Study of variation in these sympatric samples allows and wagneri. Leptodactylus dantasi, a very distinct species, is clarification of species differences and similarities. still known from only the holotype. Leptodactylus
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