INTRODUCTION Anurans exhibit a diversity o\' reproduc-tive modes, many of which are elTective means of escaping partly or completely from the constraints imposed by aquatic eggs and larvae. Direct development of terrestrial eggs and the concomitant omission of aquatic larvae has evolved independently in many lineages, as witnessed by anurans having this reproductive mode in such di-verse families as the Leiopelmatidac, Lep-todactylidae, Bufonidae, Hylidae, Ranidae, and Microhylidae. Thus, direct development of terrestrial eggs is a convergence in many anuran lineages. Studies on the embryology of diverse species having direct development have shown different developmental patterns and, especially important, different embry-onic respiratory mechanisms, such as caudal tissue in leptodactylids, lateral folds in platy-mantine ranids, and large gills in hylids. The only hylids that have direct develop-ment brood their eggs on the dorsum or in a dorsal pouch of the female. All of these have large, bell-shaped gills that partly or com-pletely envelop the developing embryo. This type of gill is unique to these frogs. There-fore, we may assume that the egg-brooding hylid frogs represent a monophyletic group. These frogs have been recognized as com-prising the subfamily Amphignathodontinae (Duellman, 1970), with the exception that the osteologically bizarre, carnivorous frogs of the genus Hemiphractiis have been ac-corded subfamilial rank— Hemiphractinae (Trueb, 1974). The Amphignathodontinae contains six genera. Two of these— Cryptobatrachus (3 species) and Stefania (7 species) carry the eggs on the dorsum and have direct develop-ment; in these features they are like the five species of Hemiphractus. In the two species of Fritziana the eggs are carried in an open basin on the dorsum; they hatch into large, nonfeeding tadpoles. Females of the other genera of egg-brooding hylids have dorsal pouches in which the eggs develop into feeding tadpoles (some Gastrorheca), non-feeding tadpoles (Flectonotus) , or froglets (Amphignathodon and some Gastrotheca). In the absence of a brooding basin or pouch, three genera can be considered <^ex\-crdViAcd—Cryptohatrachus, Stefania, and Hemiphractus— -dhhough each is charac-terized by the absence of a free larval stage. Hemiphractus has been reviewed systemat-ically (Trueb, 1974), and it is morphologi-cally and behaviorally quite distinct from the other two genera. Cryptohatrachus currently is being studied by Pedro M. Ruiz-C. and Maria C. Ardila M. of Bogota, Colombia. The purposes of the present paper are to: 1) define the genus Stefania; 2) review the taxonomy of the species; 3) present accumu-lated information on the life history, ecol-ogy, and distribution of the species; and 4) assess the phylogenetic relationships of the species and the genus among amphignatho-dontine hylids. The first mention of any of these gener-alized egg-brooding hylids is Boulenger's (1904) description of Hyla evansi from Guyana. The single female was carrying 22 eggs on the dorsum. Ruthven (1915) noted this brooding behavior in a Colombian frog referred to Hyla fuhrmanni Peracca. Ruthven (1916) named the genus Cryp-tohatrachus for these same specimens, for which he recognized a new species— C. boulengeri. In a review of the morphology and life history of egg-brooding hylids, Ruthven (1922) placed Hyla evansi Boulen-ger and H. fuhrmanni Perracca in the genus Cryptohatrachus. With the exception of No-ble (1925), who placed these species in Hyloscirtus Peters { = Hyla; Duellman, 1970) but who recognized Cryptohatrachus in 1927, subsequent workers recognized the three species of Cryptohatrachus. Rivero (1968) concluded that C. evansi was gener-ically distinct from the Colombian species and erected the genus Stefania; he also described three new species in the genus and transferred a species named as a Hyla to the genus. Rivero (1970) added another species to Stefania, and Cochran and Coin (1970) named a third species of Cryptohatrachus. Thus, at the present time Cryptohatrachus contains three species in the Andes of north-ern Colombia, whereas Stefania contains seven species in the Guianan region.
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