HARVARD UNIVERSITY Library of the Museum of Comparative Zoology OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas NUMBER 124, PAGES 1-19 APRIL 18. 1988 LOWER TELMATOBIINAE (ANURA: LEPTODACTYLIDAE): GENERIC DIAGNOSES BASED ON LARVAL CHARACTERS by /5PR 0 9 E. O. LAVILLA^ HARV UNIVERSITY As currcnlly recognized, the lower Telmatobiinae includes three tribes and 14 genera (Lynch, 1978; Laurent, 1983; Lavilla, 1983 ). These are, as follows: Batrachylini {Batrachyla and Thoropd), Calyptocephalellini {Cau- diverbera and Telmatobufo), and Telmatobiini (Alsodes, Atelognathus, Batrachophrynus, Eupsophiis, Hylorina, Insuetophrynus, Limnomedusa, Lynchophrys, Somuncuria, and Telmatobius). Larval characters have been used, often with differing results, in the taxonomic arrangements of anurans. Among the lower Telmatobiinae, for example, there have been some attempts to define genera using features of tadpole morphology (e.g., Lynch [1971] for leptodactyloid genera, Heyer [1975] for leptodactylid genera, and Formas [1975, 1976] for some lower telmatobiine genera). The following diagnoses include nine morphological units, with 16 characters relating to larval morphology, and two non-morphological units, with four characters relating to reproductive mode and larval behavior. The larvae of all genera except Lynchophrys are diagnosed in the accounts that follow. 'PRHERP-CONICET, Fundacion Miguel Lillo, 4000Tucuman, Argentina. Associ- ate in Herpetology, Museum of Natural History, The University of Kansas, Lawrence, KS 66045. 2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY MATERIALS AND METHODS Descriptions are based on tadpoles in developmental stages 31-35 of Gosner's (1960) table. Characters employed in the generic diagnoses are those that vary intergenerically but show little or no intrageneric variation. Each character is numbered (1-20) and the character states are designated as "A," "B," "C," etc. The rank of sequential character states in no way implies polarities. A data matrix of characters is provided in Table 1 . Table L — Character states of lower telmatobiine anuran larvae. See Morphologi- cal and Non-Morphological Character States for explanation of characters 1-20 and character states. Taxonomic abbreviations: THOR = Thoropa; BCHY = Batrachyla; CDVB = Caudiverbera; TMBF = Telmatobufo; ALSO = Alsodes; ATEL = Ate- lognathus; BTCO = Batrachophrynus; EUPS = Eupsophus; HYLO = Hylorina; INSU = Insuetophrynus; LIMN ^Limnomedusa; SOMU =Somuncuria; TELM = Tel- matobius. * = variable character. ( Character Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 THOR A A A F B B A A A B A A A B A B* B B B A BCHY A A A C B A A A A A A B* B* = A A A B B B A CDVB A A A E B A A A B A A A B A A A A AAA TMBF B A B F A A A A A A A A B B B A A AAA ALSO A A*A C A A A A A A A A B A A A A AAA ATEL A A A C B A A B A B A A B A A A A AAA BTCO A B A A B A A A A A A A B A A A A AAA EUPS A A A F B A B A A A A A A A A A B B B B HYLO A A A C B A A A A A A A B A A A B AAA INSU A B A F B A B A A A B A B A A A A AAA LIMN A A A D A A A A A A A A A A A A A AAA SOMU A A A E B A A A A A A A B A A A A AAA TELM A B A B B^ = A A A A A A A B A A A A AAA MORPHOLOGICAL CHARACTER STATES Oral Disc. Known also as "lips," the oral disc (Fig. 1) is the fleshy structure that surrounds the mouth in nearly all Type III and IV larvae. Laterally, it is possible to recognize three regions, important in the definition of papillae position. The oral angle region is the midlateral area on either side of the disc along which the oral disc can be folded; the area above the oral angle is termed the supraangular region, and the area below, the infraangular. The upper margin of the oral disc is designated the rostral region and the lower margin, the mental region. The most important characters related to the oral disc are its size and the presence or absence of constrictions. TELMATOBIINAE GENERIC DIAGNOSES ■*^ ..i»»- ..i\\\\ x^""'"""7/,„ A Fig. 1. — Oral discs of lower telmatobiine tadpoles. A. Telmatobius, showing transangular margin, intramarginal lateral papillae in angular area, and absence of intramarginal mental row of papillae. B . Limnomedusa, showing intranagular margin, intramarginal lateral papillae in supra- and infraangular regions and absence of papillae in angular area, and presence of intramarginal mental row of papillae. 1. Width of Oral Disc. A small oral disc has a width less than two- thirds that of maximum body width, whereas the width of a large oral disc is greater than two-thirds maximum body width. A. Less than two-thirds maximum body width. B. Greater than two-thirds maximum body width. 2. Lateral Margins of Oral Disc. If constrictions are present in the oral angle of the lateral margin, the disc is defined as intraangular, whereas if they are absent, the disc is termed transangular. A. Intraangular. B. Transangular. 4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Oral Papillae. Depending upon the distribution of these fleshy projec- tions from the oral disc (Fig. 1), it is possible to recognize two different patterns of papillae. Marginal papillae lie along the periphery of the disc; their presence frequently gives the disc an indented appearance. If marginal papillae are absent in part or all of the rostral region, Uie gap is known as the rostral gap. Intramarginal papillae are the inner papillae of the oral disc; these may be subdivided regionally into rostral, mental, and lateral intramarginal papillae. The most important characters related to oral papillae are the presence or absence of a rostral gap and the disposition of the intramarginal rows. 3. Rostral Gap. A. Present. B. Absent. 4. Intramarginal Lateral Papillae. A. Uninterrupted laterally. B. Present in the angular region and present or absent in the supra- and infraangular areas; if present in the latter areas, there are fewer papillae than when the intramarginal row is uninterrupted. C. Papillae present in the infra- and supraangular regions, but absent in the angular area. D. Papillae present only in the supraangular region. E. Papillae present only in the infraangular area. F. Papillae absent. 5. Intramarginal Mental Papillae. A. Present. B. Absent. Rostrodonts. These keratinized structures (Fig. 2C, D) surrounding the mouth are also known as "homy beaks." I follow the terminology of Van Dijk (1966), who recognized a suprarostrodont (=upper beak) associated with the suprarostral cartilage, and an infrarostrodont (=lower beak) associated with the infrarostral cartilage. 6. Rostrodont Shape. A. Wider than high. B. Higher than wide. Keratodonts. Van Dijk (1966) applied the name keratodont to the keratinized structures occurring in rows on the oral disc (Fig. 1). They are also known as "homy teeth," "odontoids," and "denticles." I consider a normal complement of keratodonts to be two upper and three lower rows. Animals having fewer rows have a reduced keratodont formula. 7. Keratodont Formula. A. 2/3. B. Reduced (i.e., fewer than 2/3). Nostrils. 8. Level of Nostril Aperture (Fig. 2A, B). A. Not raised (i.e., flush, or partially or completely depressed). B. Raised. TELMATOBIINAE GENERIC DIAGNOSES D Fig. 2. — Larval characters of lower telmatobiine tadpoles. A. General view of Atelognathus, showing raised nostrils. B. Detail of nasal chamber (shaded area) and nasal tube of Atelognathus. C. Rostrodont that is wider than high, the general condition of lower telmatobiine larvae. D. Rostrodont that is higher than wide, a feature characterizing Thoropa. The rostronasal position is defined by the ratio between the frontonasal distance (i.e., the distance from tip of snout to anterior border of nostrils; FN) and nasoocular distance (i.e., the distance from the posterior border of nostrils to anterior border of eye; NO). 9. Value ofFN/NO Ratio. A. Greater than 1 .0. B. Less than 1.0. Spiracle. 10. Position of Sinistral Spiracle. A. Lateral. B. Ventral. The location of the spiracle (Fig. 3) along the longitudinal axis of the body is defined by the ratio between the rostro-spiracular distance (i.e., the distance from the tip of the snout to the midpoint of the spiracular opening; RSD), and the spiracular-posterior position (i.e., the distance from the midpoint of the spiracular opening to the base of the proctodeal tube if it is present. If a proctodeal tube is not present, the posterior limit of the body is interpreted as the border between the body and the caudal musculature, ventrally; SPD). 11. Value of RSD I SPD Ratio. A. Greater than 1 .0. B. Less than 1.0. OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY A B D E Fig. 3. — Larval characters of lower telmatobiine tadpoles. A. Telmatobius; spir- acle lateral and sinistral, tail fins not reduced. B. Telmatobufo; reduced tail fins, oral disc modified as sucker. C. Thoropa; spiracle ventral and sinistral, tail fins reduced, oral disc normal, abdominal sucker present. Bars = 1 cm. TELMATOBIINAE GENERIC DIAGNOSES 7 Vent. 12. Proctodeal Tube. A. Present. B. Absent. 13. Orientation of Vent (Fig. 4). A. Opening at midline. B. Opening dextrally. Tail. Usually, the tail fins (Fig. 3) originate on the anterior third of the caudal musculature. If fins are reduced, they originate on the posterior two- thirds of the length of the caudal musculature. 14. Size of Tail Fins. A. Normal. B. Reduced. Special Features. 15. Oral Disc Modified as Sucker (Fig. 3). A, Absent. B. Present, 16. Abdominal Sucker (Fig. 3). A. Absent. B. Present. NON-MORPHOLOGICAL CHARACTER STATES Reproductive Strategies. 17. Egg Deposition. A. In water. B. Out of water, or in special situations (described in each case). 18. Larval Hatching. A. In water, B. Out of water, or in special situations (described in each case). 19. Larval Development. A. In water. B. Out of water, or in special situations (described in each case). Larval Behavior. 20. Larval Feeding. A. Active, B, Non-feeding larvae. RESULTS GENERIC DIAGNOSES OF LOWER TELMATOBIINAE Thoropa Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae absent; intramarginal mental papillae 8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Fig. 4. — Variation in the protodeal tube among lower telmatobiine larvae. A. Tube present and not reduced (most genera). B. Tube reduced {Hylorina). C. Tube absent {Batrachyla). TELMATOBIINAE GENERIC DIAGNOSES 9 absent. Rostrodonts deeper than wide; keratodont formula normal [(1) (1-1)/ (1-1) (2)] . Level of nostril aperture not raised; FN/NO > 1 .0. Spiracle position ventrally sinistral; RSD/SPD > 1.0. Spiracle position lateral sinistral; RSD/ SPD > 1.0. Proctodeal tube present; vent opening at midline. Tail fins reduced. Oral disc sucker absent; abdominal sucker present (except in T. miliaris). Eggs, hatching, and larval development on wet stones covered by a thin film of water. Larvae active feeders. Content. Four species: T. lutzi, T. miliaris, T. megatytnpanum, and T. petropolitana. Distribution. Mountain ranges of SE Brazil. Total length. 24.7-35. 1 mm. The lowest values are shown by T. miliaris, and the highest by T. petropolitana; T. megatympanum was not considered. Batrachyla Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae present in infra- and supraangular regions, but not in angular area; intramarginal mental papillae absent. Rostrodonts wider than deep; keratodont formula normal [(1) (1-1)/(1-1) (2)]. Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube absent (except in B. leptopus); vent opening dextrally (except in B. leptopus). Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs laid in vegetation out of water; hatching and first larval stages out of water; larvae reaching water by active movement. Larvae active feeders. Content. Three species: B. antartandica, B. leptopus, and B. taeniata. Distribution. Nothofagus forest in Chile and Argentina. Batrachyla taeniata also occurs in nonforested habitat south to Valparaiso (approx. 32° S). Total length. 2 1 .3-36.6 mm. The lowest values are shown by B. leptopus and the highest by B. antartandica. Caudiverbera Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae present only in infiraangular region; intramarginal mental papillae absent. Rostrodonts wider than deep; kerato- dont formula normal [(1) (1-1)/(1-1) (2)], poorly developed. Level of nostril aperture not raised; FN/NO < 1.0. Spiracle position lateral sinistral; RSD/ SPD > 1 .0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in water. Larvae active feeders. Content. Monotypic: Caudiverbera caudiverbera. Distribution. In Chile, from Aconcagua Province to Puerto Mont. Fossils known from Oligocene of Patagonia (Argentina). Total length. 60.5-73.7 mm. 1 0 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Telmatobufo Diagnosis. Oral disc large; lateral margins intraangular; rostral gap absent; intramarginal lateral papillae absent; intramarginal mental papillae present. Rostrodonts wider than deep; keratodont formula normal, fused [(2)/ (1-1) (2)]. Level of nostril aperture notraised; FN/NO > 1 .0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins reduced. Oral disc sucker present; abdominal sucker absent. Eggs, hatching, and larval development in water. Larvae active feeders. Content. Three species: T. australis, T. bullocki, and T. venustus. Distribution. Restricted populations in Nothofagus forest in southern Chile. Total length. 52.1-58.8 mm for T. australis. Alsodes Diagnosis. Oral disc small, lateral margins intraangular (except A. mon- ticola, with transangular margin); rostral gap present; intramarginal lateral papillae present in infra- and supraangular regions, but not in angular area; intramarginal mental papillae present. Rostrodonts wider than deep; kerato- dont formula normal [(1)(1-1)/(1- 1) (2)]. Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in water. Larvae active feeders. Content. Ten species: A. barrioi, A. gargola (with two subspecies), A. laevis,A. monticola,A. montanus,A. nodosus,A.pehuenche,A. tumultuosus, A. vanzolini, and A. verrucosus. Distribution. In Nothofagus forest of Chile and Argentina, central Chile, and northern Patagonia (Argentina). Total length. 39.8-77.1 mm. The lowest value is shown by A. gargola neuquensis, and the highest by A. tumultuosus. The larvae of A. laevis, A. vanzolini, and A. verrucosus were not considered. Atelognathus Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae in supra- and infraangular regions but not in angular area; intramarginal mental papillae absent. Rostrodonts wider than deep; keratodont formula normal [(1) (1-1)/(1-1) (2)]. Level of nostril aperture raised, opening at tip of a short nasal tube; FN/NO > 1.0. Spiracle position abdominal sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in water. Larvae active feeders. Content. Seven species: A. grandisonae, A. nitoi, A. patagonicus, A. TELMATOBIINAE GENERIC DIAGNOSES 1 1 praebasalticus (four subspecies), A. reberverii, A. salai, and A. solitarius. Distribution. Nothofagus forest of Chile and Argentina, Patagonia, and Meseta de Somuncura (Argentina). Total length. 38.8-53.3 mm. The lowest value is shown by A. patagoni- cus and the highest by A. reberverii. Only these two taxa were considered. Batrachophrynus Diagnosis. Oral disc small; lateral margins transangular; rostral gap present; intramarginal lateral papillae present uninterrupted in lateral region; intramarginal mental papillae absent. Rostrodonts wider than deep; kerato- dont f orm ula normal [( 1 ) ( 1 - 1 )/( 1 - 1 ) (2)] . Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in water. Larvae active feeders. Content. Monotypic: Batrachophrynus macrostomus. Distribution. Departamento Junin, Peru. Total length. 98.6-127.6 mm. Eupsophus Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae absent; intramarginal mental papillae absent. Rostrodonts wider than deep; keratodont formula reduced [(1) (1-1)/ (1-1) (1)], poorly developed. Level of nostril aperture not raised; FN/NO > LO. Spriracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening medially. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in tiny reservoirs of water. Non-feeding larvae. Content. Five species: E. calcaratus, E. insularis, E. miqueli, E. roseus, and E. vitattus. Distribution. In Nothofagus forest of Chile and Argentina. Total length. 16.7-21.5 mm for £". roseus (the only species considered). Hylorina Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae present in supra- and infi-aangular regions but not in angular area; intramarginal mental papillae absent. Rostt^o- donts wider than deep; keratodont formula normal [( 1 ) ( 1 - 1 )/( 1 - 1 ) ( 1 )] . Level of nostril aperture not raised; FN/NO > 1 .0. Spiracle position lateral sinistral; RSD/SPD > 1 .0. Proctodeal tube present, small; vent opening dextrally. Tail fins not reduced. Oral disc sucker absent; abdominal sucker absent. Eggs laid in vegetation out of water; hatching and larval development in water. Larvae active feeders. Content. Monotypic: Hylorina sylvatica. 12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY Distribution. Nothofagus forest of Chile and Argentina. Total length. 48.1-53.0 mm. Insuetophrynus Diagnosis. Oral disc small, lateral margins transangular; rostral gap present; intramarginal lateral papillae absent; intramarginal mental papillae absent. Rostrodonts wider than deep; keratodont formula reduced [(1) (1-1)/ (2-2), with infraangular rows vestigial or absent]. Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD < 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval develop- ment in water. Larvae active feeders. Content. Monotypic: Insuetophrynus acarpicus. Distribution. Nothofagus forest of Valdivia Province, Chile. Total length. 48.7-49.6 mm. Limnomedusa Diagnosis. Oral disc small; margins intraangular; rostral gap present; intramarginal lateral papillae present only in supraangular region; intra- marginal mental papillae absent (marginal papillae in multiple rows men- tally). Rostrodonts wider than deep; keratodont formula normal [(1) (1 - 1)/(1 - 1) (2)]. Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening medially. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development out of water. Larvae active feeders. Content. Monotypic: Limnomedusa macroglossa. Distribution. Uruguay, NE Argentina, SE Brazil, and Paraguay (?). Total length. 40.7-42.5 mm. Somuncuria Diagnosis. Oral disc small; lateral margins intraangular; rostral gap present; intramarginal lateral papillae present only in infraangular region; intramarginal mental papillae absent. Rostrodonts wider than deep; kerato- dont formula normal [( 1 )( 1 - 1 )/( 1 - 1 ) (2)] . Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval development in water. Content. Monotypic: Somuncuria somuncurensis. Distribution. Meseta de Somuncura, Rio Negro Province, Argentina. Total length. 35.1-41.8 mm. Telmatobius Diagnosis. Oral disc small; lateral margins intraangular; rostral gap TELMATOBIINAE GENERIC DIAGNOSES 13 present; intramarginal lateral papillae present in angular area; present or not in supra- and infraangular regions, but if present, fewer than in Batra- chophrynus; intramarginal mental papillae following two patterns — ^present in the most austral taxa (i.e., Argentinian forms), and absent in septentrional taxa (i.e., in Bolivia, Chile, Peru, and Ecuador). Rostrodonts wider than deep; keratodont formula normal [(1) (1-1)/(1-1) (2)]. Level of nostril aperture not raised; FN/NO > 1.0. Spiracle position lateral sinistral; RSD/SPD > 1.0. Proctodeal tube present; vent opening dextrally. Tail fins normal. Oral disc sucker absent; abdominal sucker absent. Eggs, hatching, and larval develop- ment in water. Larvae active feeders. Content. About 28 species, and 55 subspecies. Distribution. Highlands of Argentina, Bolivia, Chile, Peru, and Ecuador, from San Juan Province (Argentina) to Imbabura Province (Ecuador). Total length. Extreme measurements for tadpoles of 36 taxa: 46.7-95.9 mm. The lowest value is shown by T. culeus culeus, and the highest by T. marmoratus angustipes. COMMENTS The genera of lower telmatobiines characteristically have been defined by features of the adults. Larval characters are not concordant with the generic grouping of species in three recognized genera. Telmatobius. In a previous paper (Lavilla, 1985), I proposed the recogni- tion of two "species groups" in Telmatobius. The southern group includes those taxa from the highlands of Argentina, characterized by the presence of an intramarginal mental row of papillae in the oral disc (a character shared with Alsodes, Limnomedusa, and Telmatobufo, but not with the northern taxa). The northern group is characterized by the absence of the aforemen- tioned row of papillae and includes the known larvae from Chile, Bolivia, Peru, and Ecuador. Alsodes. Tadpoles o( Alsodes monticola are morphometrically different from the other species of the genus (Lavilla and Scrocchi, 1986). Moreover, larvae of this taxon have a transangular oral disc, a character shared with Batrachophrynus, Insuetophrynus, and Telmatobius, but not with the other Alsodes. Batrachyla. Two species of this genus (B. antartandica and fi. taeniata) are characterized by the absence of a proctodeal tube and presence of a dextral vent. This condition is not shared by Batrachyla leptopus which has a short proctodeal tube with median aperture. KEY TO THE GENERA OF THE LOWER TELMATOBHNAE BASED ON LARVAL CHARACTERS la. Oral disc large (>2/3 body width), modified as a sucker; rostral gap absent; keratodont formula [(2)/(l-l) (2)] Telmatobufo 14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY lb. Oral disc small (<2/3 body width), not modified as a sucker; rostral gap present; keratodont formula [(1) (1-1)/(1-1) (2)], or reduced 2 2a. Rostrodonts wider than high; without abdominal sucker; tail fins origin normal; without rupicole habits 3 2b. Rostrodonts higher than wide; generally with abdominal sucker; tail fins origin reduced; with rupicole habits Thoropa 3a.Keratodontformula[(l)(l-l)/(l-l)(2)] 5 3b. Keratodont formula reduced 4 4a. Oral angle constrictions present; ratio RSD/SPD > 1 ; vent opens medially; eggs, hatching, and larval development in tiny water reservoirs; non- feeding larvae Eupsophus 4b. Oral angle constrictions absent; ratio RSD/SPD < 1; vent opens dextrally; eggs, hatching, and larval development in normal situations; actively feeding larvae Jnsuetophrynus 5a. Abdominal spiracle; nostrils raised, at the end of a short nasal tube Atelognathus 5b. Lateral spiracle; nostrils not raised, open totally or partially flush or in a little depression 6 6a. Proctodeal tube absent £atrachyla(pan) 6b. Proctodeal tube present 7 7a. Ratio FN/NO < 1 Caudiverbera 7b. Rauo FN/NO > 1 8 8a. Oral disc intraangular 9 8b. Oral disc transangular 13 9a. Intramarginal mental row of papillae absent 10 9b. Intramarginal row of papillae present Alsodes (part) 10a. Vent median Matrachyla (part) 10b. Vent dextral 11 11a. Intramarginal lateral papillae present in supra- and infraangular regions, but not in angular area; eggs laid out of water Mylorina lib. Intramarginal lateral papillae present only in one of the two regions; eggs laid in water 12 12a. Intramarginal lateral papillae present only in supraangular region Limnomedusa 12b. Intramarginal lateral papillae present only in infraangular region Somuncuria TELMATOBIINAE GENERIC DIAGNOSES 15 13a. Intramarginal mental row of papillae present 14 13b. Intramarginal mental row of papillae absent 15 14a. Intramarginal lateral papillae only in supra-angular region (infraangular papillae mixed with mental row); angular area without papillae Abodes (psTt) 14b. Intramarginal lateral papillae in angular area; may be present, in low numbers, in supra- and infraangular regions Telmatobius (part) 15a. Intramarginal lateral papillae uninterrupted in lateral region Satrachophrynus 15b. Intramarginal lateral papillae in angular area; may be present, in low numbers, in supra- and infraangular regions Telmatobius (part) LITERATURE ON TELMATOBIINE TADPOLES The following are references to papers dealing with Telmatobiinae tad- poles. BATRACHYLINI Thoropa. — lutzi: Bokermann (1965), Lavilla (1983); megatympanum: Caramaschi and Sazima(l 984): w/Z/amiBokerm ann( 1965), Lavilla( 1983), Wassersug and Heyer {\9?>T)\ pelropolitana: Bokermann (1965), Heyer and Crombie (1979), Lavilla (1983), Wassersug and Heyer (1983). Batrachyla.—antartandica: Barrio (1967a), Formas (1976), Cei (1980), Lavilla (1983); leptopus: Busse (1971), Formas (1976), Cei (1980), Lavilla (1983); taeniata: Cei and Capurro (1958), Cei (1962), Formas (1976), Cei (1980), LavUla (1983). CALYPTOCEPHALELLINI Telmatobufo. — australis: Formas (1972), Lavilla (1983); venustus: Diaz etal.(1983). Caudiverbera. — caudiverbera: Krieg (1924), Cei (1962), Jorquera and Izquierdo (1964), Lavilla (1983), Diaz and Valencia (1985). TELMATOBIINI Alsodes. — barrioi: Velosoetal. (1981), Lavilla (1983); gargola: Gallardo (1970), Cei (1980), Lavilla (1983); monticola: Formas (1975), Lavilla (1983); montanus: Busse (1980), Lavilla (1983), Lavilla and Scrocchi (1986); nodosus: Cei (1962), Formas (1975), Lavilla (1983); pehuenche: Cei (1980), Lavilla (1983); tumultuosus: Lavilla (1983), Diaz and Valencia (1985). Atelognathus.—patagonicus: Cei (1965), Cei (1980), Lavilla (1983); 16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY reberverii: Cei (1980), Lavilla (1983). Batrachophrynus. — macrostomus: Lavilla (1983). Eupsophus. — roseus: Formas and Pugin (1978a), Cei (1980), Formas and Vera (1980), Lavilla (1983); vittatus: Formas and Pugin (1978b), Formas and Vera (1980). Hylorina. — sylvatica: Barrio (1967b), Formas and Pugin (1978b), Cei (1980), Formas and Vera (1980), Lavilla (1983). Insuetophrynus. — acarpicus: Formas et al. (1980), Lavilla (1983). Limnomedusa. — macroglossa: Cei (1980), Gehrau and de Sa (1980), Gudynas and Gerhau (1981), Lavilla (1983). Somuncuria. — somuncurensis: Cei (1980), Lavilla (1983). Telmatobius.—albiventris: Vellard (1951), Lavilla (1983, 1985); arequi- pensis: Lavilla (1983, 1985); atacamensis: Gallardo (1962), Cei (1980), Lavilla (1983, 1985); ceiorum: Cei (1980), Lavilla (1983, 1984b, 1985); crawfordi: Vellard (1953), Lavilla (1983, 1985); culeus: Lavilla (1983, 1985); halli: Cei (1962), Lavilla (1983, 1985); hauthali: Koslowsky (1895), Fernandez (1926), Lavilla (1983, 1984a, 1985); hauthali pisanoi: LaviUa (1983, 1984b, 19^5); jelski: Vellard (1951), Lavilla (1983, 1985); laticeps: Cei (1980), Lavilla (1983, 1984b); marmoratus: Vellard (1951), Cei (1980), Lavilla (1983, 1985); niger. Trueb (1979); oxycephalus: Cei (1980), Lavilla (1983, 1985); pefauri: Lavilla (1983, 1985), Diaz and Valencia (1985); peruvianas: Schmidt (1928), Vellard (1951), Cei (1962), Lavilla (1983, 1985); rimac: Lavilla (1983, 1985); schreiteri: Vellard (1946), Cei (1980), Lavilla (1985); stephani: Lavilla (1983, 1985); vellardi: Trueb (1979). ACKNOWLEDGMENTS Material for this study was available thanks to the kindness of Nelly Carrillo de Espinosa (Museo Javier Prado, Lima, Peru), William E. Duellman (Museum of Natural History, University of Kansas), Ramon Formas (Insti- tuto de Zoologia, Universidad Austral, Valdivia, Chile), Raymond F. Laurent (PRHERP-CONICET, Fundacion Miguel Lillo, Tucuman, Argentina), Jose Valencia (Facultad de Ciencias, Santiago, Chile), Paulo Emilio Vanzolini (Museu de 2^ologia, Universidade de Sao Paulo, SP, Brasil), and Alberto Veloso (Instituto de Biologia Celular y Genetica, Universidad de Chile, Santiago, Chile). LITERATURE CITED Barrio, A. 1967a. Batrachyla antartandica n. sp. (Anura: Leptodactylidae). Descripcion y estudio comparativo con la especie genotipica B. leptopus Bell. Physis (Bs.As.) 27(74):101-109. Barrio, A. 1967b. Observaciones eco-etologicas sobre Hylorina sylvatica Bell (Anura: Leptodactylidae). Physis (Bs.As.) 27(74):153-157. Bokermann, W. C. A. 1965. 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Biologia y desarrollo de Eupsophus taeniatus Girard. Inv. Zool. Chil. 4:77-82. Diaz, N., M. Sallaberry, and H. Nunez. 1983. The tadpole oiTelmatobufo venustus (Anura: Leptodactylidae) with considerations of generic relationships. Herpeto- logica39(2):lll-113. Diaz, N. and J. Valencia. 1985. Larval morphology and phenetic relationships of the C\u\.Q.dinAlsodes, Telmatobius, Caudiverbera and Insuetophrynus (Anura: Lepto- dactylidae). Ccpeia 1985:175-181. Fernandez, K. 1926. Sobre la biologia y reproduccion de batracios argentinos. Bol. Acad. Nac. Cienc. Cordoba 29:271-320. Formas, J. R. 1972. A second species of Chilean frog genus Telmatobufo (Anura: Leptodactylidae). J. Herpetol. 6:1-3. Formas, J. R. 1975. Las larvas de las esp)ecies chilenas pertenecientes al genero Eupsophus grupo nodosus (Anura: Leptodactylidae). Bol. Soc. Biol. Concepcion 49:231-237. Formas, J. R. 1976. Descriptions oi Batrachyla tadpoles. J. Herpetol. 10:221-225. Formas, J. R., N. F. Diaz, and J. Valencia. 1980. 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Biologische Reisestudien in Sudamerika. 11. Rhinoderma und Calyp- tocephalus. Zeitschr. Morph. Okologie der Tiere 3(1):150-168. Laurent, R. F. 1983. Heterogeneidad del genero Batrachophrynus Peters. Acta Zool. Lillona37(l):107-113. LaviUa,E.0. 1983. Sistematicadelarv as deTelmatobiinae(Anura: LeptodactyUdae). Ph.D. Dissertation. Fac. Cs. Nat. Univ. Nac. Tucuman, Argentina, v + 354 pp. LaviUa, E. O. 1984a. Redescubrimiento de Telmatobius hauthali Koslowsky, 1895, y descripcion de su larva. Acta Zool. Lilloana 38:51-57. LaviUa, E. 0. 1984b. Larvas de Telmatobius (Anura: LeptodactyUdae) de la Provincia de Tucuman (Argentina). Acta Zool. Lilloana 38:69-79. LaviUa, E. O. 1985. Diagnosis genericay agrupacion de las especies de Telmatobius (Anura: LeptodactyUdae) en base a caracteres larvales. Physis (Bs.As.) B 43(105):63-67. LaviUa, E. O. and G. J. Scrocchi. 1986. Morfometria larval de los generos de TeUnatobiinae (Anura: LeptodactyUdae) de .^Tgentinay ChUe. Physis (Bs.As.) B 44(106):39^3. Lynch, J. D. 1971. Evolutionary relationships, osteology and zoogeography of Leptodactyloid frogs. Univ. Kansas Mus. Nat. Hist. Misc. Publ. 53:1-238. Lynch, J. D. 1978. A re-assessment of the Telmatobiinae leptodactylid frogs of Patagonia. Occ. Pap. Mus. Nat. Hist. Univ. Kansas 72:1-57. PhiUppi, R. A. 1902. Suplemento a los Batraquios chilenos descritos en la Historia Fisica y Politica de Chile de don Claudio Gay. Santiago, ChUe. 161 pp. Schmidt, K. P. 1928. The Chilean frogs of the genus Telmatobius. Rev. Chil. Hist. Nat. 32:98-105. Trueb, L. 1979. LeptodactyUd frogs of the genus Telmatobius in Ecuador, with the description of a new species. Copeia 1979:714-733. Van Dijk, D. E. 1966. Systematics and field keys to the famiUes, genera and described species of southern Africa anuran tadpoles. Ann. Natal Mus. 18(2):23 1-286. VeUard, J. 1946. El genero Telmatobius en la Republica Argentina. Acta Zool. LUloana 3:313-326. VeUard, J. 1951. Estudios sobre batracios andinos. I. El grupo Telmatobius y formas afines. Mem. Mus. Hist. Nat. Javier Prado 1:89 pp. VeUard, J. 1953. Estudios sobre batracios andinos. II. El grupo Marmoratus y formas afmes. Mem. Mus. Hist. Nat. Javier Prado 2:53 pp. + 4 pi. Veloso, A. M., N. P. Diaz, P. C. Iturra, and M. V. Penna. 1981. Descripcion de una nueva especie de telmatobino del gcnexo Alsodes (Amphibia: LeptodactyUdae) de la Cordillera de Nahuelbuta (sur de Chile). Medio Ambicnte 51(2):72-77. TELMATOBIINAE GENERIC DIAGNOSES 19 Wassersug, R. and W. R. Heyer. 1983. Morphological correlates of subaereal existence in Leptodactylid tadpoles associated with flowing water. Can. J. Zool 61(4):761-769. University of Kansas publications MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum of Natural History. 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