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Molecular Systematics of the Frog Genus Leptodactylus (Amphibia: Leptodactylidae)

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Molecular Systematics of the Frog Genus Leptodactylus (Amphibia: Leptodactylidae) 1 UNIVERSITY OF I1L1NOIS LIBRAR" AT SSS5-CHAMPAWM BIOLOGY APR 91992 oology W SERIES, NO. 41 olecular Systematics of the Frog Genus Leptodaetylus (Amphibia: Leptodactylidae) Linda R. Maxson I19BB W. Ronald Heyer biology i\mm 101 BURWLL HALl A Contribution in Celebration of the Distinguished Scholarship of Robert F. Inger on the Occasion of His Sixty-Fifth Birthday 29, 1988 "'blication 1384 BLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: Fieldiana is primarily a journal for Field Museum staff members and research associates, although manuscripts from nonaffiliated authors may be considered as space permits. The Journal carries a page charge of $65 research and invited authors will con- per printed page or fraction thereof. 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References should be typed in the following form: Croat. T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp. Grubb, P. J., J. R. Lloyd, andT. D. PENNINGTON. 1963. A comparison of montane and lowland rain forest in Ecuador. I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601. ... J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L . and R. A. Schwarz, eds., Spirits. Shamans, and Stars. Mouton Publishers, The Hague. Netherlands. Mlrra, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American Indians. Vol. 2, The Andean Civilizations. Bulletin 143. Bureau of American Ethnology, Smithsonian Institution. Washington, D.C. >LZE, R. G. 1981. Ferns and fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana: Botany, n.s.. 6: 1-522. Illustrations: Illustrations are referred to in the text as "figures" (not as "plates"). Figures must be accompanied by some indication of scale, normally a reference bar. Statements in figure captions alone, such as "x 0.8," are not Fieldiana for details o\' acceptable. Captions should be typed double-spaced and consecutively. See recent issues of style. ires as submitted should, whenever practicable, be 8'/2 by 11 inches (22 x 28 cm) and may not exceed 11 Vi by 16'/2 inches (30 x 42 cm). Illustrations should be mounted on boards in the arrangement you wish to obtain in the ink printed work. This original set should be suitable for transmission to the printer as follows: Pen and drawings may the size and be originals (preferred) or photostats; shaded drawings should be originals, but within limitation; photo- the reverse with stats should be high-quality, glossy, black and white prints. All illustrations should be marked on the author's name, figure number(s), and "top." Original illustrations will be returned to author upon publication or color unless otherwise specified. Authors who wish to publish figures that require costly special paper reproduction must make prior arrangements with the Scientific Editor. Page Proofs: Fieldiana employs a two-step correction system. Each author will normally receive a copy of the edited manuscript on which deletions, additions, and changes can be made and queries answered. Only one set of page proofs will be sent. All desired corrections of type must be made on the single set of page proofs. Changes in page proofs (as opposed to corrections) are very expensive. Author-generated changes in page proofs can only be made if the author tdvance to pay for them. FIELDIANA Zoology NEW SERIES, NO. 41 Molecular Systematics of the Frog Genus Leptodactylus (Amphibia: Leptodactylidae) Linda R. Maxson Department of Ecology, Ethology, and Evolution University of Illinois at Urbana-Champaign Urbana. Illinois 61801 W. Ronald Heyer Department of Vertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC. 20560 A Contribution in Celebration of the Distinguished Scholarship of Robert F. Inger on the Occasion of His Sixty-Fifth Birthday Accepted for publication February 10, 1986 February 29, 1988 Publication 1384 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1988 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Tables Abstract 1 1. Matrix of reciprocal immunological dis- Introduction 1 tances among 1 species of Leptodactylus Materials and Methods 2 3 Results and Discussion 2 2. One-way immunological distances in the Reciprocal Reactions 3 Leptodactylus melanonotus group 5 Phylogenetic Considerations Based on Re- 3. One-way tests in the Leptodactylus ocel- ciprocal Data 4 latus group 5 One-Way Reactions 4 4. One-way tests in the Leptodactylus penta- Divergence Times in Leptodactylus 8 dactylus group 6 Conclusions 9 5. One-way tests in the Leptodactylus fuscus Acknowledgments 10 group 7 Literature Cited 10 6. One-way tests comparing Leptodactylus Appendix 12 riveroi and L. silvinambus to other species of Leptodactylus 7 List of Illustrations Phylogenetic relationships among mem- bers of the Leptodactylus pentadactylus group using L. labrosus as an outgroup Histograms showing frequency of pair- wise immunological distance compari- sons indicative of lineages diverging in the indicated geological epochs ui Molecular Systematics of the Frog Genus Leptodactylus (Amphibia: Leptodactylidae) Abstract More than three-quarters of the described species eastern Brazil and Amazonia are specifically dis- of the frog genus Leptodactylus were sampled and tinct from L. ocellatus from southeastern Brazil analyzed using the quantitative immunological and Uruguay. technique of micro-complement fixation. Eleven albumin antisera to representatives of the four de- scribed species groups in this genus were compared Introduction to one another and to albumins of all available species of Leptodactylus. The Neotropical frog genus Leptodactylus con- The results of this analysis indicated enormous sists of over 45 species. Comparative morpholog- albumin differentiation within the genus, suggest- ical and behavioral data (Heyer, 1969, 1979, and ing that most species of Leptodactylus have been other revisions cited therein) indicate that these established since the Paleocene with modest spe- species divide into four lineages. To test this hy- ciation occurring throughout the Eocene, Oligo- pothesis we have been gathering micro-comple- cene, and Miocene. No evidence for Pleistocene ment fixation (mc'f) data on albumin evolution speciation has been found. among Leptodactylus species since 1974. We ini- The four species groups of Leptodactylus defined tially anticipated that a few representative species on morphological and behavioral criteria are not samples would establish a molecular framework as clearly defined by this biochemical analysis. Al- to determine the relationships among the major bumins of species within each of the L. pentadac- lineages within the genus. Our early results indi- tylus, L. melanonotus, and L. ocellatus groups are cated that the problem of defining relationships more similar to one another than to members of within Leptodactylus was more complex than an- other groups. However, there is little evidence of ticipated. It is only now that we have sufficient close relationships among those members of the data from mc'f analyses to evaluate the utility of L. fuscus group available for study. Leptodactylus this approach for delineating relationships within riveroi is not genetically close to any of the refer- the genus. Because of Robert F. Inger's interest in ence species and appears to represent yet another frog systematics, we offer this summary, warts and lineage in this genus. Leptodactylus silvinambus all, as a token of our appreciation for his influence has its closest relatives among members of the L.
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