Bioacoustics of Nymphargus mariae

In order to provide future workers with data useful in characterizing Cochranella puyoensis , a numbered diagnosis is provided: ( 1 ) vomerine teeth present; ( 2 ) bones green in life; white or whitish green in preservative; ( 3 ) parietal peritoneum covered by guanophores (white), extending posteriorly to the level of the liver and covering the upper half of the stomach; visceral peritoneum clear; ( 4 ) coloration in life dark to light green dorsally with small yellow to cream spots/specks, venter bluish-green; in preservative dark to pale purplish-grey with light spots; ( 5 ) webbing formula of hand: webbing absent between I and II and vestigal between II and III, III ( 3 -– 3 ) – ( 2 1 / 2 – 2 2 / 3 ) IV; ( 6 ) webbing formula of foot: I ( 2 – 2 -) – ( 2 1 / 3 – 2 1 / 2 ) II 1 1 / 2 – 2 – III ( 1 1 / 2 – 1 2 / 3 ) – 2 2 / 3 IV ( 2 +– 2 2 / 3 ) – ( 1 2 / 3 – 2 -) V; ( 7 ) snout truncate in dorsal view and truncate to rounded in profile; nostrils slightly elevated and separated by a shallow depression; loreal region concave; ( 8 ) all skin surfaces with minute granulations, dorsal skin with scattered low warts or spicules; ( 9 ) low nonenameled ridge on forearms and hand; unscaloped fold on tarsus; ( 10 ) humeral spine absent in males; ( 11 ) tympanum evident ventrally, upper 1 / 4 to 1 / 3 concealed, directed dorsolaterally with slight posterior inclination; supratympanic fold moderately developed to barely evident; ( 12 ) 21.2–25.4 mm SVL in males, 27.2–30.0 mm SVL in females; ( 13 ) prepollical spine not protruding externally; unpigmented nuptial pad Type I; ( 14 ) anal ornamentation with at least two paired tubercles below vent and some additional tubercles and folds; ( 15 ) first finger longer than second; ( 16 ) lobate liver without guanophores; ( 17 ) eye diameter larger than width of disc on finger III; ( 18 ) iris in life light blue-green with a transverse green-brown bar ( holotype , DFCH-USFQ D 285 , QCAZ 7104 ) or white with grey-green fine reticulations and a transverse gray bar ( QCAZ 7499 ), iris in preservative grey with some fine dark reticulations, minute dark punctuations, and a transverse dark grey bar; ( 19 ) some melanophores on fingers and toes; ( 20 ) advertisement call unknown. The absence of a humeral spine in C. puyoensis would present a contradiction for the hypothesized relationships with C. mariae ( Flores & McDiarmid 1989 ), because C. mariae was reported to have humeral spines (M. Hensel, pers. comm., in Duellman and Schulte 1993 : 31 ). Unfortunately, no voucher specimens of males of mariae have been studied or reported to be deposited in collections, and the state of the humeral crista ventralis (either with or without a humeral spine) has not been corroborated. The coloration patterns of puyoensis , mariae and azulae are similar and, to the best of our knownledge, unique among glassfrogs (only C. ocellata has a similar coloration pattern, and we think that it is probably related to C. puyoensis ), but without a phylogeny of Centrolenidae any conclusion about their relationships is speculative at best and dependent on the collection and examination of male specimens of mariae and azulae . If mariae and puyoensis are closely related, their placement in different genera would be a consequence of the non-monophyly nature of the centrolenid genera Cochranella and Centrolene (and the non-informative nature of the character used to separate them: humeral spines) or to C. mariae actually lacking humeral spines, and Hensel's comments based on a male of a different species. The conclusions by Flores and McDiarmid ( 1989 ) on the relationships between puyoensis , mariae and azulae (= mariae species group) were based on several characters considered synapomorphies (e.g., microgranulations on the skin surfaces, reduced interdigital webbing). Most subsequent authors have followed their hypothesis ( Ruiz-Carranza & Lynch 1991 ; Duellman & Schulte 1993 ). Ruiz-Carranza and Lynch ( 1995 a) were the first authors to suggest that the proposed mariae group was based on phenetic rather than derived characters, and we concur. The skin microgranulations were considered a unique character uniting C. azulae , C. mariae , and C. puyoensis , but we have subsequently found them present also in C. cochranae ( Goin, 1961 ) , C. posadae ( Ruiz-Carranza & Lynch, 1995 a ), C. luminosa ( Ruiz-Carranza & Lynch, 1995 b ), C. luteopunctata ( Ruiz-Carranza & Lynch, 1996 ) , and C. chami ( Ruiz-Carranza & Lynch, 1995 b )—and they probably occur in more centrolenid species. Reduced webbing between the outer fingers is a condition present in several species of Cochranella . The particular combination of reduced hand webbing and presence of vomerine teeth is present in Cochranella chami , C. cochranae (some specimens), C. cristinae ( Ruiz-Carranza & Lynch, 1995 b ), C. luminosa , C. luteopunctata , C. prasina ( Duellman, 1981 ) , C. puyoensis , and C. spilota ( Ruiz-Carranza & Lynch, 1997 ) .