and pattern characters are analyzed for the adult members of the Leptodactylus pentadactylus species group. Available data on tadpoles, mating calls, and karyotypes are incorporated in the analyses. Results of the analyses lead to the recognition of 11 species comprising the group. For each species, the following are provided: synonymy, description of adult characteristics, a distribution map and list of localities, and specimens examined. The following are included if known: distinctive adult colors in life, larval characteristics, mating call description, and karyotype. A key is provided for the adult members of the complex. Larval adaptations appear to be most important in interpreting the evolutionary history within this species group. Adult morphology appears to be evolutionarily rather conservative, contrasting with the patterns found in other species groups so far studied within this genus. 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 Systematics of the pentadactylus species group of the frog genus Leptodactylus (Amphibia: Leptodactylidae) (Smithsonian contributions to zoology ; no.

Tadpoles involved in predator-prey interactions were studied in tropical wet forest in Costa Rica under laboratory and field conditions. Larvae of the frog Leptodactylus pentadactylas and naiads of the odonate Pantala flavescens are important predators on larvae of several species of frogs. The predators discriminate the prey on the basis of size and species, but not type of habitat in which predation occurs. A graphical model is proposed to illustrate the relationships between species diversity and habitat complexity as they affect the composition of tadpole communities. The model is used to evaluate the relative importance of abiotic and biotic factors in determining the use of specific kinds of aquatic habitats by frogs with larval stages. Predation by permanent aquatic predators (primarily fish) is considered to be the most important biotic factor influencing the temporal and spatial composition of tadpole communities. The development and maintenance of predatory feeding modes, including cannibalism, in certain tadpoles is examined in light of the model.MOST TADPOLES are generalized herbivores (Jenssen 1967) that either scrape plant material from a substrate or filter planktonic food directly from the water. Only a few species are known to be carnivorous although some instances of cannibalism among tadpoles have been reported (Bragg 1964). We were surprised, therefore, to find a well-developed predator-prey system in which tadpoles of one species were carnivores on other tadpoles. This finding was unusual not only because documented instances of tadpoles feeding on other tadpoles are relatively rare, but also because the carnivorous tadpoles were found in relatively small puddles.The predator-prey interactions studied occur in small rain-filled tropical puddles characteristically utilized by opportunistic breeders. Experiments were run to study the relative ability of predators to capture prey and their preference for certain prey species, to ascertain the influence of microhabitat on predator-prey interactions, and to evaluate the influence of prey size on their ability to escape predation.Results of these experiments led to a consideration of tadpole habitats and to those factors controlling the diversity of tadpole communities. A model is used to illustrate the relationship between pond complexity and predation as they affect the species composition of a tadpole community. The predatory feeding mode of tadpoles is considered in light of this model. METHODS AND MATERIALSObservations and experiments were made during two weeks in June 1973 (WRH and RWM) and two weeks in August 1973 (DLW and RWM) in the vicinity of the Osa Field Station, Rincon de Osa, Puntarenas Province, Costa Rica. The station is surrounded by the remnants of the lowland wet forests that once covered much of the Golfo Dulce region. This area receives more than 4000 mm of rain each year and is the only remaining large tract of wet forest on the Pacific coast of Middle America. Most of the work was done in the immediate vicinity of ...

Heyer, W. Ronald. A Preliminary Analysis of the Intergeneric Relationships of the Frog Family Leptodactylidae. Smithsonian Contributions to Zoology, number 199, 55 pages, appendix, 16 figures, 38 tables, 1975.-Thirty-seven characters of external morphology, myology, osteology, life history, and chromosome morphology are studied. For each character, the evolutionary directions of changes of states are inferred. This information is used to construct a phylogenetic hypothesis of the intergeneric relationships of the New World frog family Leptodactylidae. Five major groupings of leptodactylids are proposed: the telmatobines, ceratophrines, leptodactylines, grypiscines, and eleutherodactylines. Formal recognition of these groupings is delayed until more information becomes available which will likely modify the intra-and intergroup relationships. The phylogenetic analysis demonstrates that the five groups are robust units, however. Recognition of the five groups allows a reinterpretation of the historical zoogeography of the family. The family Leptodactylidae had its origins in the temperate beech forests of South America. The telmatobines represent a remnant of the original leptodactylid stock, which has remained in the beech forests. Two groups became adapted to drying conditions, the ceratophrines and leptodactylines. The grypiscines represent a forest-stream adaptational complex that centered in southeastern Brazil. The eleutherodactylines were probably derived from a grypiscine ancestor. Early attainment of direct development in the eleutherodactylines was a preadaptation which resulted in an explosive radiation of the Eleutherodactylus-complex, which is now represented by about 350 species which occupy diverse environmental situations. A leptodactylid-liopelmatid relationship is suggested, which has the advantages of an in situ evolution of the leptodactylids rather than a migration from North Temperate regions as previously proposed. An alternate leptodactylid-discoglossid relationship argument which was based in large part on tadpole evidence is countered by a consideration of the major functional adaptations of tadpoles. OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5309. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging in Publication Data Hcyer, W. Ronald A preliminary analysis of the intergeneric relationships of the frog family Leptodactylidae.

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