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Rivera-Correa, Mauricio, Baldo, Diego, Candioti, Florencia Vera, Orrico, Victor Goyannes Dill, Blackburn, David C., Castroviejo-Fisher, Santiago, Chan, Kin Onn, Gambale, Priscilla, Gower, David J., Quah, Evan S.H., Rowley, Jodi J. L., Twomey, Evan, Vences, Miguel (2021): Amphibians in Zootaxa: 20 salamanders, and caecilians years documenting the global diversity of frogs. Zootaxa 4979 (1): 57-69, DOI: https://doi.org/10.11646/zootaxa.4979.1.9

Hyalinobatrachium carlesvilai Castroviejo-Fisher, Padial, Chaparro, Aguayo, and De la Riva 2009 Figure 23 Background information. Castroviejo-Fisher et al. (2009) described this species from the Amazon slopes of the Andes of Bolivia and Peru, with the northernmost locality in the west slope of Abra Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06�� 12 ��� S, 76 �� 44 ��� W, 1080 m), Provincia Lamas, Region San Mart��n, Peru. This latter locality was based on a re-examination of the type series of H. lemur Duellman and Schulte, 1993, in which the only paratype (KU 211769) was assigned to H. carlesvilai (Castroviejo-Fisher et al. 2009). New data. We report five new localities for Hyalinobatrachium carlesvilai from northern Peru. Specimen MHNC 13958 (adult male) was collected on 28 March 2010 by M. Guerra Panaijo and Santiago Castroviejo-Fisher at Puente Chino on the Tocache-Pucallpa road, approximately 29 km NE from Tingo Maria (9 �� 8 ' 9.03 "S, 75 �� 47 ' 15.78 "W, 1147 m), Ucayali, Peru. To the best of our knowledge, this is the first report of this species for Ucayali. The frog was calling on the underside of a tree leaf overhanging a stream, at ~ 2 m above the water level and shows all the phenotypic characteristics described by Castroviejo-Fisher et al. (2009) for H. carlesvilai, except for pericardium color. All specimens of H. carlesvilai examined in the original description (19 individuals from 12 different localities of Peru and Bolivia) have a completely white pericardium due to the accumulation of iridophores. However, the specimen from Ucayali has a mostly transparent pericardium, with only a small and peripheral portion of the pericardium showing iridophores (Fig. 23). Nonetheless, we confirm its identification through phylogenetic analysis of DNA sequences (Fig. 2 a). Pericardium transparency is known to be an intraspecific polymorphic character in other species of Hyalinobatrachium (e.g., Guayasamin et al. 2006; Cisneros-Heredia & McDiarmid 2007; Castroviejo-Fisher et al. 2011; our Fig. 23). As suggested by previous authors, intraspecific variation should be examined to better determine if this character is diagnostic. In addition to the above locality, four additional localities are added: One adult male (voucher MNCN 45954) was collected from Puente Lahuarpia, 7 km NW from Pacayzapa, San Mart��n, Peru (6 �� 13 ' 35.56 "S, 76 �� 48 ' 58.03 "W, 787 m), on 8 February 2010 by ET, M. Pepper, and M. Sanchez-Rodriguez. Three individuals (CORBIDI-HE- 2010-8950, CORBIDI-HE- 2010-8954, and CORBIDI-HE- 2010-8959) were collected from 3 km E Pacayzapa (6 �� 16 ' 33.78 "S, 76 �� 44 ' 47.22 "W, 1021 m) on 28 July 2010 by ET, T. Kosch, and A. Delgado. One individual (CORBIDI 10474) was collected from 4.5 km E Pacayzapa (6 �� 16 ' 52.87 "S, 76 �� 43 ' 57.86 "W, 1111 m) on 6 June 2011 by JD and ET. This site corresponds to the type locality for H. lemur Duellman and Schulte (1993) (which is now in synonymy with H. pellucidum; see Castroviejo-Fisher et al. 2009). One individual (voucher ET- 12 -006) was collected from the upper R��o Shunte drainage, San Mart��n, Peru (8 �� 20 ' 45.24 "S, 76 �� 43 ' 29.10 "W, 966 m) on 12 March 2012 by JD and ET. These specimens fully concord with the original description, and identification of five of these specimens (from two localities; see Fig. 2 a and Appendix 3) was also confirmed through the phylogenetic analysis. These records from San Mart��n are important because they confirm the presence of the species in the area, which was previously based on a single, 16 year old specimen (the paratype of H. lemur, re-identified by Castroviejo-Fisher et al. 2009)., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on page 44, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33.","Guayasamin, J. M., Bustamante, M. R., Almeida-Reinoso, D. & Funk, W. C. (2006) Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnean Society, 147, 489 - 513. http: // dx. doi. org / 10.1111 / j. 1096 - 3642.2006.00223. x","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82.","Castroviejo-Fisher, S., Moravec, J., Aparicio, J., Guerrero-Reinhard, M. & Calderon, G. (2011) DNA taxonomy reveals two new species records of Hyalinobatrachium (Anura: Centrolenidae) for Bolivia. Zootaxa, 2798, 64 - 68."]}

Centrolene charapita new species Figures 4���6 Centrolene sp. 1 Castroviejo-Fisher, Guayasamin, Gonzalez-Voyer, and Vil�� 2014 Holotype. MHNCP 13933, adult male in good state of preservation (left hind limb was cut below the knee and preserved in 96 % ethanol for DNA analyses) from near the village of La Oliva past the village of Muyo (a larger village roughly 49 km N from Bagua), Amazonas, Peru (5 �� 18 ' 3.86 "S, 78 �� 23 ' 44.57 "W, 682 m), collected by M. Guerra Panaijo, E. Twomey, and S. Castroviejo-Fisher on 2 April 2010. Paratopotype. MNCN 45392, same data as holotype. Generic placement. The new species is placed in the genus Centrolene on the basis of our phylogenetic analyses that included complete or partial sequences of the mitochondrial genes 12 S and 16 S of the two known individuals of C. charapita (Fig. 2 b). Additionally, the following phenotypic characters frequently observed in species of Centrolene are present in the new species (sensu Guayasamin et al. 2009): trilobed liver, covered by a transparent hepatic peritoneum; ventral parietal peritoneum white anteriorly and transparent/translucent posteriorly; bones green in life; dorsum lavender in preservative; dorsum of males with spicules during breeding season; nuptial pads conspicuous in males; enameled pericloacal warts. However, the value of these characters as synapomorphies for Centrolene needs to be tested. Diagnosis. The following unique combination of characters distinguishes Centrolene charapita from other species of glassfrogs: (1) a dentigerous process on each vomer, each bearing a row of 4���6 small vomerine teeth; (2) snout truncate in dorsal view and truncate (slightly rounded) in profile; (3) tympanum covered by skin, not visible through skin; in preservative tympanic annulus not visible externally but appreciable under magnification in life; supratympanic fold absent; (4) dorsal skin texture complex, in life and preservative smooth with enameled microspicules and low enameled warts on ocelli, conspicuous enameled warts forming an irregular lateral band from snout to mid-part of body and extending into the dorsoposterior edge of the upper arm, skin of the venter and thighs coarsely areolate while all other ventral surfaces are smooth; (5) cloacal ornamentation conspicuous, formed by folded skin bearing four large enameled warts projected anteriorly (flaps,) surrounded by various lower warts with or without enameling; (6) parietal peritoneum anterior third white, posterior two-thirds transparent (state P 2); pericardium, hepatic, renal, and gonadal peritonea transparent, visceral (stomach and intestines) peritoneum white; (7) liver with three lobules (state H0); (8) humeral spines not present; (9) webbing formula of fingers III 2 - 1 + IV; (10) webbing formula of toes I 1 / 2 - 2 - II 1 - 2 - III 1 - 2 IV 2 - 1 V; (11) scalloped enameled ulnar and tarsal folds, extending from fringe on postaxial edge of Finger IV to elbow and from fringe on postaxial edge of Toe V to ankle, respectively; (12) nuptial pad Type I, not pigmented, enameled glands present on webbing and fringes of finger and toes, prepollex and prepollical spine concealed by skin; (13) when appressed, Finger I of equal length with Finger II; (14) eye diameter 2.5 times the width of disc on Finger III; (15) coloration in life: dorsal and dorsolateral surfaces with yellow or pale green spots of different size (ocelli) bearing enameled warts and microspicules set in a dark green reticulum (darker around the ocelli), upper lip and tip of fingers and Toes I, II, and III white (enameled), ventral surfaces, flanks, upper arms, Fingers I and II, and Toes I and II not pigmented, bones green; (16) coloration in preservative: dorsal and dorsolateral surfaces with cream spots of different size (ocelli) bearing enameled warts and microspicules set in a lavender-greyish reticulum formed by minute melanophores that are darker around the ocelli, upper lip and tips of fingers and toes I, II, and III white (enameled), ventral surfaces, flanks, upper arms, Fingers I and II, and Toes I and II not pigmented; (17) iris in life off-white to light grey with yellow tones and black dots and reticulation with a black ring delimiting the iris, iris in preservative white with black dots and reticulation with a black ring delimiting the iris; (18) in life and preservative tip (discs) of fingers and toes white (enameled), Finger IV and Toes IV and V pigmented, Finger III and Toe III not pigmented but for a small group of melanophores towards the tip, Fingers and Toes I and II not pigmented; (19) large size adult males, SVL = 34.7 ���37.0 mm (N = 2), unknown in females. Comparisons. Specimens of Centrolene charapita have a unique combination of characters that allows easy differentiation from all other known species of centrolenids. The presence of scalloped enameled ulnar and tarsal folds together with its large size (SVL> 34.0 mm) and a truncated snout in profile help diagnose this unique species. Specimens of Ce. daidaleum, Ce. paezorum, Ce. geckoideum, Ce. savagei, Ce. solitaria, Cochranella mache, Co. phryxa, Co. resplendens and V. uranoscopa have scalloped enameled ulnar and tarsal folds, weakly so in Ce. huilense, Ce. muelleri, and Nymphargus ocellatus; however all the aforementioned species are smaller (adult males SVL Ce. geckoideum, which has a truncated snout in profile but is much larger than the new species, up to 83.5 mm SVL in males and 72.0 mm SVL in females. Additionally, none of these species have the characteristic ocelli on dorsum and enameled warts on finger and toe webs and fringes of Ce. charapita (specimens of N. ocellatus have ocelli but no webs between Finger III���IV). Our analysis retrieved Ce. geckoideum as the sister species of Ce. charapita, but the following external characters are clearly diagnostic (character states of Ce. charapita in parenthesis): adult males with large and externally projecting conical humeral spines (humeral spines absent), forearms hyper-atrophied (normal), dorsal coloration dull green (dorsal coloration green with ocelli, Figs. 4 and 5), diameter of eye smaller than that of disc of Finger III (larger), adult males SVL> 50.0 mm (Description of holotype. Adult male with SVL 37.0 mm. Head about 1 / 3 wider than body; head width 36 % of SVL; head width 1.1 times head length. Snout truncate in dorsal view, in lateral view truncated (slightly rounded); eye-nostril distance/eye diameter 0.61 and eye-nostril distance/interorbital distance 0.88. Loreal region concave, nostrils round, flush with surrounding skin, slightly elevated; internarial region concave anterodorsally; canthus rostralis well defined. Eyes medium, directed anterolaterally, eyes angled 45.0�� relative to midline (where anteriorfacing eyes would be 90 �� relative to midline); eye diameter 2.5 times wider than width of disc on Finger III; eye diameter 52 % of head length and 1.44 times interorbital distance. Tympanum covered by skin, not visible through skin, tympanic annulus appreciable under magnification in life but not in preservative, supratympanic fold absent. A dentigerous process on each vomer present, each bearing a row of 4���6 vomerine small teeth (their size is so small that we could not determine the exact number); choanae large, drop-shaped, separated more widely than nostrils; tongue round, white in preservative, anterior 3 / 4 attached to floor of mouth; vocal slits present, extending from the sides of the base of tongue to the level of the mandibular joints. Forelimbs relatively robust, with forearm flattened and roughly 2 times as wide as arm; ulnar fold present, white (enameled) and scalloped; humeral spine absent. Relative length of fingers: I ~ II Coloration in life of both types (Fig. 4). Dorsal and dorsolateral surfaces with yellow or pale green spots of different size (ocelli), bearing enameled warts and microspicules, set in a dark green reticulum (darker around the ocelli). Upper lip and tips (discs) of fingers and toes I, II, and III white (enameled). Ventral surfaces, flanks, upper arms, Fingers I and II, and Toes I and II not pigmented. Iris delimited by a black ring, background coloration off white to light grey with yellow shades and black dots and reticulation. Parietal peritoneum anterior third white, posterior two-thirds transparent. Visceral peritoneum (stomach and intestines) white. Hepatic peritoneum transparent. Other peritonea not visible in life. Coloration in preservative of both types (Fig. 5). Dorsal and dorsolateral surfaces with cream spots of different size (ocelli), bearing enameled warts and microspicules, set in a lavender-greyish reticulum formed by minute melanophores that are darker around the ocelli. Upper lip and tips (discs) of fingers and toes I, II, and III white (enameled). Ventral surfaces, flanks, upper arms, Fingers I and II, and Toes I and II not pigmented. Finger III and Toe III not pigmented but for a small group of melanophores towards the tip. Finger IV and Toes IV and V pigmented as dorsum. Iris delimited by a black ring, background coloration white with black dots and reticulation. Parietal peritoneum anterior third white, posterior two thirds transparent, pericardium, hepatic, renal, and gonadal peritonea transparent, visceral (stomach and intestines) peritoneum white. Color of peritonea based on direct observations on the dissected holotype. Measurements. Holotype measurements (in mm) with paratype measurements given in parentheses: SVL 37.0 (34.7); HL 11.9 (10.5); HW 13.3 (12.5); IND 2.8 (2.6); IOD 4.3 (3.8); ED 6.2 (6.5); EA 44.8 �� (not measured in paratype); EW 3.7 (3.3); END 4.4 (4.3); HaL 11.0 (11.0); 3 DW 2.5 (2.3); TL 19.6 (19.0); SL 21.1 (20.1); FL 16.3 (15.5). Variation. Except for the size there is no evident variation in the two individuals collected. Even the number and distribution of the large dorsal ocelli are virtually the same in both specimens. Distribution and ecology. This species is known only from the type locality (Fig. 7). Both specimens were found perched on vegetation along a relatively large secondary stream east of the Mara����n River. This species likely occupies other streams in the mostly undisturbed forests of the surrounding mountains. The holotype was found perched on a branch approximately 2 m above the water. Its vocal sac was extended as typical of glassfrogs that have been recently calling. The paratype was found on the upper side of a Heliconia leaf (Heliconiaceae) approximately 5 m above the water. Other amphibians found in the same stream included Ameerega trivittata, Bolitoglossa altamazonica, Cochranella erminea, Hyloscirtus sp., Rulyrana mcdiarmidi, Pristimantis aff. acuminatus, Pristimantis sp., and Rhaebo glaberrimus. Advertisement call, fighting behavior, egg clutches, parental care, and tadpole are unknown for this species. Etymology. As applied here, the specific epithet ��� charapita ��� derives from a type of small yellow chili pepper (aji charapita) that is endemic to the Amazon lowlands and used in a variety of Amazonian dishes. These peppers resemble the yellowish ocelli characteristic of the dorsal coloration of this new species (Fig. 8). Remarks. Because we have only analyzed sequences of two genes (12 S and 16 S) and our results are not highly supported (Fig. 2 b; JK = 72; GB = 3), the phylogenetic position of Centrolene charapita might change as more molecular and/or phenotypic data are analyzed, or if more species are included, particularly species in the Centrolene geckoideum group (containing Ce. acanthidiocephalum, Ce. geckoideum, Ce. medemi, Ce. paezorum, and Ce. petrophilum). If the current phylogenetic position is corroborated, Ce. charapita would be the only species of Centrolene known to occur below 800 m., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 12-17, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Castroviejo-Fisher, S., Guayasamin, J. M., Gonzalez-Voyer, A. & Vila, C. (2014) Neotropical diversification seen through glassfrogs. Journal of Biogeography, 41, 66 - 80. http: // dx. doi. org / 10.1111 / jbi. 12208","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97."]}

Rulyrana mcdiarmidi (Cisneros-Heredia, Venegas, Rada, & Schulte 2008) Figure 27 Cochranella mcdiarmidi Cisneros-Heredia, Venegas, Rada, and Schulte 2008. Holotype DFCH-USFQ D 132, an adult female from ���a small rivulet tributary of the Jambue River, ca. 6 km S from Zamora (ca. 04��03���S, 78 �� 56 ���W, 1150 m), on the western slope of Contrafuerte de Tzunantza, Cordillera Oriental, eastern slopes of the Andes, Provincia de Zamora- Chinchipe, Rep��blica del Ecuador ��� Rulyrana mcdiarmidi��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil�� 2009 Background information. This species was described from a series of five specimens collected from two sites in southern Ecuador and one site in northern Peru at elevations between 1100���1210 m. The species was assigned to Cochranella on the basis of the humeral spine being absent in males (Cisneros-Heredia et al. 2008). Guayasamin et al. (2009) assigned this species to the genus Rulyrana considering morphological similarity to R. flavopunctata, a species that had been sequenced in their analysis. New data. Since 2010 we have encountered this species in a number of sites throughout northern Peru, extending the known range of this species roughly 100 km to the south and east of the Peruvian locality from the original description, and between elevations of 682���1811 m (Fig. 14). Here, we provide a list of new localities and associated voucher specimens (when collected): village of Cocachimba, Amazonas, Peru, near Gocta waterfall (CORBIDI-HE- 2010-6847 and CORBIDI-HE- 2010 -6848, 6�� 3 ' 44.28 "S, 77 �� 53 ' 36.47 "W, 1811 m); near the village of La Oliva past the village of Muyo (a larger village roughly 49 km N from Bagua), Amazonas, Peru (MNCN 45957 and MHNC 14015, 5�� 18 ' 3.86 "S, 78 �� 23 ' 44.57 "W, 682 m); south slope of Utcubamba canyon, 4.7 km NW from Pedro Ruiz, Amazonas, Peru (uncollected individuals, 5 �� 55 ' 5.80 "S, 78 ��0' 38.62 "W, 1238 m); north slope of Utcubamba canyon, 11.3 km WNW from Pedro Ruiz, Amazonas, Peru (CORBIDI 10470, 5�� 54 ' 50.93 "S, 78 �� 4 ' 35.20 "W, 1077 m); Quebrada Goca on Yambrasbamba road, about 1 km N from road entrance, Amazonas, Peru (CORBIDI 10473, 5�� 45 ' 50.78 "S, 77 �� 54 ' 46.52 "W, 1711 m). Variation. We have not noted variation in this species beyond what is reported in Cisneros-Heredia et al. (2008), except that dorsal coloration ranges from light to dark green and the intensity of dorsal spotting varies among individuals. One juvenile (CORBIDI 10470) had reduced spotting compared to adults photographed in a site nearby. Thus, intensity of spotting may exhibit ontogenetic variation. Vocalizations. We have heard R. mcdiarmidi calling in two sites (4.7 km NW from Pedro Ruiz and Quebrada Goca, mentioned above). Unfortunately, we have been unable to obtain a recording of this species. The call sounds similar to that of R. saxiscandens (described below), consisting of a single, piercing ���zip��� of short duration. In the site near Pedro Ruiz, several males were observed calling from a vertical rock face of a small waterfall. In Quebrada Goca, one male was observed calling (and several others were heard) on a small branch hanging roughly 1 m above the edge of the stream. These frogs were concentrated around fast-flowing sections of the stream, but despite background noise, the call could be heard up to roughly 50 m away. Distribution and ecology. Including the new localities mentioned above, R. mcdiarmidi is widely distributed in the Utcubamba and central Mara����n drainages in Amazonas, Peru and eastern foothills of southern Ecuador at elevations between 682���1811 m. This species has not been found on the Huallaga-side of Abra Patricia in northern Peru. R. mcdiarmidi occurs in a wide variety of habitats, ranging from pristine rainforest (e.g, near Muyo) to heavily disturbed, scrubby habitats (e.g., near Cocachimba). Adults and juveniles have all been found on rocks or vegetation typically within 2 m of the water surface. Eggs and tadpoles remain unknown. Rulyrana mcdiarmidi is sympatric with a number of species throughout its known range, including Nymphargus posadae in the southern Cordillera del Condor in Peru (Cisneros-Heredia et al. 2008), Cochranella erminea and Centrolene charapita in the Muyo site in Peru, and Espadarana cf. fernandoi in Quebrada Goca, Peru (Fig. 27 f). Taxonomic remarks. Some of the characters used to differentiate R. mcdiarmidi from R. saxiscandens (following our definition that considers R. tangarana as a junior synonym, see below) do not appear to be unambiguously diagnostic. For example, purple coloration in preservative is variable in R. saxiscandens, with some specimens appearing dull green or grayish lavender (see R. saxiscandens account below). On average, R. mcdiarmidi likely has a larger SVL, but the two species overlap with respect to this character (two R. mcdiarmidi males 25.4���26.9 mm, six R. saxiscandens males 22.3���26.6 mm). Ulnar folds are visible in life in some individuals of R. saxiscandens, but generally less-pronounced than in R. mcdiarmidi. Also, R. mcdiarmidi is described as having medium-sized eyes (ED/ 3 WD = 1.10���1.70), even though both male paratypes have ED/ 3 WD of 1.93, again, within the variation we observed in R. saxiscandens (ED/ 3 WD = 1.75���2.45, N = 6). However, presence of flat warts with low tubercles (which are typically lime green in life) in R. mcdiarmidi vs. skin uniformly spiculate and shagreened in R. saxiscandens may be a good diagnostic character between these two species. Nevertheless, the results from our phylogenetic analysis do not recover R. mcdiarmidi and R. saxiscandens as reciprocally monophyletic. Furthermore, nested within this clade is an individual referred to as R. flavopunctata (Guayasamin et al. 2008) from Provincia Morona-Santiago, Ecuador (Fig. 2 c). Also, the genetic distances among these terminals are low (p-distances = 0.0��� 0.6 %, Table 6). These results suggest the possibility of a single, widespread, morphologically variable species. However, our very limited sampling of R. flavopunctata (DNA sequences retrieved from GenBank of a specimen identified by others and that we have not examined) preclude us from taking further taxonomic decisions., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on page 48, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Cisneros-Heredia, D. F., Venegas, P. J., Rada, M. & Schulte, R. (2008) A new species of glassfrog (Anura: Centrolenidae) from the foothill Andean forests of Ecuador and Peru. Herpetologica, 64, 341 - 353. http: // dx. doi. org / 10.1655 / 06 - 078.1","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. & Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595. http: // dx. doi. org / 10.1016 / j. ympev. 2008.04.012"]}

Nymphargus chancas (Duellman & Schulte 1993) Figure 27 Cochranella chancas Duellman and Schulte 1993. Holotype KU 211778, an adult male from ���west slope of Abra Tangarana, 7 km by road northeast of San Juan de Pacaysapa (06�� 12 ��� S, 76 �� 44 ��� W, 1080 m), Provincia Lamas, Departamento San Mart��n, Peru ���. Nymphargus chancas��� Cisneros-Heredia and McDiarmid 2007. Background information. This species was described from a single adult male collected in Abra Tangarana, San Mart��n, Peru (Duellman & Schulte 1993). It was placed in the ocellata group due to the combination of white parietal peritoneum, clear visceral peritoneum, humeral spines in males absent, and vomerine teeth absent. This group was later described as a new genus Nymphargus (Cisneros-Heredia & McDiarmid 2007), and chancas was placed in this genus due to the combination of characters mentioned above, as well as its absence of hand webbing���a synapomorphy of Nymphargus. This species was later registered from the Cordillera del Condor in Ecuador (Guayasamin & Bonaccorso 2011). New data. Since 2005 we have encountered this species in two localities in northern San Mart��n (Fig. 14). In 2005, two individuals were found by day in a small stream roughly 25 km SW from Tarapoto (approx. 6 �� 35 ' 38.10 "S, 76 �� 34 ' 17.24 "W, 1131 m). On 6 June 2011 two adult males (CORBIDI 14148 and CORBIDI 10471) were collected from a small stream 7.5 km SE from the type locality (6 �� 19 ' 12.83 "S, 76 �� 41 ' 42.71 "W, 1003 m; Fig. 27). In addition, we have seen photographs of specimens from the village of Yurilamas (approximately 6 �� 14 'S, 76 �� 32 ' W, record courtesy M. Fash��) and a site roughly 14 km SE from the town of Shamboyacu, San Mart��n, Peru (7 �� 4 ' 8.9 ''S, 76 ��0' 55.2 ''W, 1122 m, voucher specimens CORBIDI-HE- 2010-8865 and CORBIDI-HE- 2010-8866, record courtesy P. Venegas). Variation. We noted variation in the prominence of yellow dorsal flecks, and rapid color-pattern changes. While all individuals we have seen possess distinct yellow flecks on the dorsum, the lone individual from Yurilamas has large, bright yellow flecks on the dorsum, appearing almost spotted. However, individuals seem to be capable of undergoing relatively rapid and dramatic color-pattern changes. For example, one of the individuals collected from Abra Tangarana had prominent occellations when we first found it in the field, but these quickly faded after capture and the frog took on the grayish-green dorsal coloration typical of this species. Iris color was described as pale yellowish-bronze in the original description (Duellman & Schulte 1993). In the specimens we have collected, iris color would be better described as silvery-white with a faint yellow suffusion throughout, and a horizontal black bar through the middle of the eye. Distribution and ecology. Nymphargus chancas is distributed throughout northern San Mart��n at elevations over 1000 m. Its distribution extends into southern Ecuador, where it was collected between 1256���1430 m (Guayasamin & Bonaccorso 2011). This species seems to be associated with small creeks with thick forest cover. Reproductive behaviors such as oviposition and calling have not yet been observed in this species. Taxonomic remarks. Our phylogenetic results revealed that Nymphargus chancas is paraphyletic. Of the three specimens included in the phylogenetic analyses, two of them cluster together as sister to N. puyoensis Flores and McDiarmid (1989) while the third one (QCAZ 41590) is sister to N. puyoensis (Fig. 2 b). Nonetheless, this third specimen is from Zamora-Chinchipe province in Ecuador and may represent a misidentification or an undescribed species. However, this specimen shows no differences with the holotype of N. chancas (J.M. Guayasamin, pers. comm.). Nymphargus chancas and N. puyoensis are easily diagnosed on the basis of vomerine teeth (present in N. puyoensis, absent in N. chancas) and dorsal coloration (dull yellowish-green with minute yellow flecks in N. chancas vs. dark green with low, flat warts corresponding to yellowish-green spots)., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on page 46, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33.","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82.","Guayasamin, J. M. & Bonaccorso, E. (2011) Evaluacion Ecologica Rapida de la Biodiversidad de los Tepuyes de la Cuenca Alta del Rio Nangaritza, Cordillera del Condor, Ecuador. Conservacion Internacional. Quito, Ecuador. [the total page number unkown]","Wiens, J. J., Fetzner, J. W., Parkinson, C. L. & Reeder, T. W. (2005) Hylid frog phylogeny and sampling strategies for speciose clades. Systematic Biology, 54, 778 - 807. http: // dx. doi. org / 10.1080 / 10635150500234625"]}

Teratohyla midas (Lynch & Duellman 1973) Centrolenella midas Lynch and Duellman 1973. Holotype KU 123219, an adult male from ���Santa Cecilia, 340 m, Provincia Napo, Ecuador. Cochranella midas��� Ruiz-Carranza and Lynch 1991 a. Teratohyla midas��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, Rada, and Vil�� 2009. Bioacoustics. This is a common, widespread species that occurs throughout much of the Amazon basin and into the northern lowlands of Brazil. For the purposes of this paper, we provide a description of the advertisement call, which has not been formally described (although see Kok & Castroviejo-Fisher 2008 for audiospectrograms and oscillograms). The following description is based on recordings from two males from two different sites in Peru. The first in an uncollected male recorded from San Jose, San Mart��n, Peru (6 �� 25 ' 14.67 "S, 76 �� 17 ' 28.42 "W, 518 m), on 4 July 2004 by J. L. Brown and ET at 22 ��C. The recording consists of a series of three notes given by the male over a 10 second period. Each note is 0.15��� 0.16 s in length, with a dominant frequency of 7030���7060 Hz, composed of four pulses each. The second recording (Fig. 30) is from Pongo de Rentema, Amazonas, Peru (5 �� 17 ' 43.68 "S, 78 �� 24 ' 22.17 "W, 557 m) recorded 2 April 2010 by SC-F and ET at 22 ��C (voucher MNCN 45963). It consists of two notes given by a single male over a 26 second period. Both notes were 0.10 s in length, each with a dominant frequency of 6760 Hz, composed of three pulses each., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on page 55, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Lynch, J. D. & Duellman, W. E. (1973) A review of the centrolenid frogs of Ecuador, with descriptions of new species. The University of Kansas Museum of Natural History Occasional Papers, 16, 1 - 66.","Ruiz-Carranza, P. & Lynch, J. D. (1991 a) Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Kok, P. J. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaieteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53."]}

Centrolene muelleri Duellman and Schulte 1993 Figure 9 Background information. Centrolene muelleri was described on the basis of a single adult male from ��� 14 km (by road) W of Venceremos, Provincia Rioja, Departamento San Mart��n, Per�� ��� (Duellman & Schulte 1993). Catenazzi et al. (2012) presented a phylogenetic tree including a sample of a frog identified as Ce. muelleri (collected by P. Venegas, but see ���Taxonomic remarks��� section for comments on this specimen) and found this species as sister of Ce. lynchi, a species from the western Cordillera of the Colombian and Ecuadorian Andes. Aside from this sequence data, no new information has become available regarding this species since the original description. New data. We add two new occurrence records (Fig. 7) and natural history observations. On two separate occasions this species was registered at the base of Gocta waterfall, Amazonas, Peru (6 �� 1 ' 37.80 "S, 77 �� 53 ' 15.15 "W, 1830 m,) by P. Venegas (six specimens, CORBIDI-HE- 2010 -0481 through 0487) in 2007 and M. Pepper in January 2011. On the first occasion (P. Venegas pers. comm.), these frogs were found at night after a heavy rainfall, with males calling 2���2.5 m above the water from the upper surfaces of leaves. Males were seen in combat while dangling from the edge of leaves. One clutch of eggs was found on the upper surface of a leaf that consisted of 46 white eggs surrounded in clear jelly. On the second occasion, several individuals were found in a small rivulet flowing into Gocta stream near the spray zone, where males were seen calling from the upper surfaces of leaves approximately 0.5 m above the water (M. Pepper pers. comm.). We (JD and ET) captured a single individual (CORBIDI 14667, Fig. 9) in a stream 7.4 km SE from Puente Vilcaniza along the Vilcaniza/Jumbilla road, Amazonas (5 �� 48 ' 52.25 " S, 77 �� 50 ' 6.19 "W, 1858 m) on 16 June 2011. This individual was calling from atop a leaf approximately 1.5 m above the water in a torrential cloud forest stream. This particular stream flowed through what appeared to be undisturbed highland forest, and was full of large rocks and deep pools with a strong current. On numerous occasions we have searched the type locality, but have been unable to find this species. However, the Vilcaniza locality is only 15 km airline from the type locality and the two sites are not separated by any barriers, thus the specimen from this locality may be a good representation of type material. Distribution and ecology. In Peru, this species has been registered from three sites spanning the divide between the Huallaga and Mara����n drainages. All three sites are located in the southern section of Departamento Amazonas between 1830 and 2050 m. Little is known regarding the natural history of this species. As mentioned above, male combat has been witnessed, and all males seen calling were on the upper surfaces of leaves. A single egg clutch, presumably belonging to this species, was found on the upper surface of a leaf. This species appears to be associated with undisturbed habitats with relatively dense canopy cover; however, the Gocta site is open, although this site lies deep in a ravine. Centrolene muelleri is sympatric with Ce. lemniscatum and Hyalinobatrachium anachoretus new species in the Alto Mayo drainage, and has been found in close proximity (Rulyrana mcdiarmidi populations in Gocta. Taxonomic remarks. In the original description, Centrolene muelleri was diagnosed primarily on the basis of large conical tubercles present on the dorsal surfaces of the head, body, and limbs (Duellman & Schulte 1993). Its body color in life was described as ���green with dark greenish-black spots and pale yellow tubercles���. Two years later, Ruiz-Carranza and Lynch (1995) described Ce. huilense, a species whose dorsal coloration was described as ���green with abundant dark green and cream flecks���. This species was not diagnosed against Ce. muelleri in the original description. Although these two species look similar (for example, both species are nearly identical with respect to snout shape in lateral profile, skin texture, dorsal coloration, absence of vomerine teeth, extent of hand webbing, and adult male SVL), Catenazzi et al. (2012) presented a molecular phylogeny supporting the status of both species. In their analysis Centrolene muelleri was sister to Ce. lynchi, while an individual of Ce. huilense from Yanayacu, Ecuador was sister to Ce. heloderma. However, our topology conflicts somewhat with the findings of Catenazzi et al. (2012). First, the Ce. muelleri from Gocta waterfall may represent an undescribed species as it is part of a larger clade containing several species (following our topology, Fig. 2 b). Second, our sequence of Ce. muelleri from Vilcaniza (15 km airline from the type locality) is sister to Ce. huilense. However, the latter sequence is from Yanayacu, Ecuador, and may not be representative of Ce. huilense sensu stricto. As we have not reviewed the specimens from Gocta, and molecular data from topotypic Ce. huilense remains unavailable, we refrain from taking any taxonomic actions at this time., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 17-18, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33."]}

Twomey, Evan, Delia, Jesse, Castroviejo-Fisher, Santiago (2014): A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species. Zootaxa 3851 (1): 1-87, DOI: http://dx.doi.org/10.11646/zootaxa.3851.1.1

Cochranella guayasamini new species Figures 19–21 Cochranella croceopodes Catenazzi and Venegas 2012 Cochranella sp. 2 Castroviejo-Fisher, Guayasamin, Gonzalez-Voyer, and Vilà 2014 Holotype. MHNC 13930, adult male in good state of preservation (right hind limb was cut below the knee and preserved for molecular analyses) collected from a stream near the village of San Jose, approximately 33 km (by road) northeast of Tarapoto, along the Yurimaguas road, San Martín, Peru (6 ° 25 ' 14.67 "S, 76 ° 17 ' 28.47 "W, 517 m), collected by S. Castroviejo-Fisher on March 31, 2010. Paratopotypes. MNCN 45950 and MHNC 13929, same data as holotype. Paratypes. MNCN 45951 and MHNC 13931, both adult males collected from a stream near the village of Santa Rosa, approximately 26 km (by road) northeast of Tarapoto along the Yurimaguas road, San Martín, Peru (6 ° 24 ' 44.79 "S, 76 ° 19 ' 2.09 "W, 686 m), collected by ET, M. Pepper, and M. Sanchez-Rodriguez on February 9, 2010; CORBIDI 8956, an adult male collected from a stream behind the village of Lejia near Abra Tangarana on the Tarapoto-Moyobamba road, San Martín, Peru (6 ° 18 ' 46.68 "S, 76 ° 43 ' 34.23 "W, 1120 m), collected by JD and ET on June 22, 2011. Referred specimens. CORBIDI-HE- 2010-6816, collected from a stream near “Puesto de Control 15 ”, near the town of Shamboyacu, San Martín, Peru (6 ° 56 ' 27.18 "S, 76 ° 3 ' 49.75 "W, 955 m), collected by P. Venegas on 2 June 2010; CORBIDI-HE- 2012-11392, collected from a stream in the northern Cordillera de Kampanquis, Amazonas, Peru (4 ° 2 ' 35.1 ''S, 77 ° 32 ' 28.3 ''W, 325 m), collected by A. Catenazzi on 12 August 2011. We identified these specimens on the basis of photographs taken by the collectors. Generic placement. The new species is placed in the genus Cochranella on the basis of our phylogenetic analyses that included complete or partial sequences of the mitochondrial genes 12 S and 16 S. Additionally, the following phenotypic characters typical of Cochranella (Guayasamin et al. 2009) are present in the new species: lobed liver covered by a transparent hepatic peritoneum; ventral parietal peritoneum white anteriorly and transparent posteriorly; moderate to extensive webbing between fingers III and IV; bones green in life; dentigerous process of the vomer and vomerine teeth present; males call from the upper surfaces of leaves, and females deposit eggs on the upper sides of leaves along streams. However, the value of these characters as synapomorphies for Cochranella needs to be tested. Diagnosis. The following unique combination of characters distinguishes Cochranella guayasamini from other species of centrolenids: (1) a dentigerous process on each vomer (in some specimens low and hardly visible), each bearing a row of two small vomerine teeth; (2) snout rounded in dorsal view, rounded and slightly inclined in profile; (3) tympanum visible, not covered by skin, angled laterally with slight dorsal inclination, tympanic annulus low, supratympanic fold absent; (4) skin of dorsum and flanks smooth, skin of venter and thighs coarsely areolate, skin of other ventral surfaces smooth; (5) cloacal ornamentation formed by folded skin bearing small and nonenameled warts and two paired round tubercles below vent; (6) parietal peritoneum anterior half white (both on the ventral and dorsal sides), posterior half transparent, pericardium white, hepatic peritoneum transparent, visceral (stomach and intestines), renal, and gonadal peritonea mostly transparent but with some small and isolated traces of iridophores; (7) liver with three lobules; (8) adult males with a small humeral spine; (9) webbing absent between fingers I and II, basal between fingers II and III, and III 2 - 1 + IV; (10) webbing formula of toes I 1 - 2 - II 1 - (2 – 2 -) III 1 - 2 IV (2 – 2 -) - 1 V; (11) folds on limbs absent, fringe on postaxial edge of finger IV and toe V; (12) nuptial pad Type I, not pigmented; (13) when appressed Finger I slightly longer than Finger II; (14) eye diameter 2.5 times the width of disc on Finger III; (15) coloration in life: dorsal and dorsolateral surfaces green with yellow tones and minute melanophores (absent in the upper lip), upper rim of lip white (enameled), eyelid rims bright yellow, ventral surfaces yellowish green, bones green; (16) coloration in preservative: dorsal and dorsolateral surfaces dull cream with minute melanophores and iridophores, the former are more conspicuous on the upper eyelid making it lavender, rim of eyelids conspicuously white due to iridophores, the thin layer of iridophores of the upper lip (enameled) is almost unappreciable, ventral surfaces and flanks dull cream; (17) iris in life reddish-bronze with minute black reticulation, black or grey spots, and a black ring delimiting the iris, yellow pupillary ring absent; in preservative, iris black with minute greyish spots and a black ring delimiting the iris; (18) in life hands and feet yellowish-green with fingers and toes yellowish, melanophores absent from hands and feet including all fingers and toes; (19) males call from upper surfaces of leaves, advertisement call consisting of one to two harsh buzzes lasting 0.166– 0.28 s (note length) and with 6080–6180 Hz dominant frequency; (20) eggs golden yellow with clear jelly, clutches deposited on upper surfaces of leaves overhanging streams (N = 10), clutch size 16– 38 eggs (N = 4); (21) medium size adult males, SVL 22.5–24.5 mm (N = 5); (23) tadpoles elongate, with sinistral spiracle, oral disc not emarginate, labial tooth row formula 1 / 3. Comparisons. No other species of glassfrog has a yellow rim around the eyelids in life, which are enameled white in preservative, except Nymphargus chancas and some specimens of " Centrolene " balionota. Specimens of the new species can be distinguished from N. chancas by (characters of the latter in parenthesis) webbing between external fingers developed, III 2 - 1 + IV (basal and hardly developed), snout rounded in dorsal and lateral views (truncate), dorsal coloration uniformly green (with yellow dots), iris reddish-bronze (white), humeral spine in adult males (absent). Specimens of the new species can be diagnosed from “ Centrolene ” balionota by (characters of the later in parenthesis) dorsal coloration uniformly green (green with large yellow patches and black dots), iris reddish-bronze (white), skin of flanks non-ornamented (ornamented with a longitudinal clusters of enameled glands). Within Cochranella, the only species with humeral spines is Co. litorale from the Pacific lowlands of the Choco in Ecuador and Colombia. However, this species has large black spots on its dorsum both in life and in preservative (absent in Co. guayasamini), and lacks dentigerous processes on the vomer (present in Co. guayasamini). Outside Cochranella, the most similar species is Centrolene durrellorum, known from the east versant of the Andes in Ecuador. However, this species has melanophores on its hands and feet including finger IV and toes IV and V (melanophores absent from hands, feet, fingers, and toes in Co. guayasamini); dorsum lavender in preservative (cream except for the lavender upper eyelids in Co. guayasamini), iris in life golden with a yellow pupillary ring (reddish-bronze without a yellow pupillary ring in Co. guayasamini). Description of holotype. Adult male with SVL of 23.4 mm. Head about 1 / 3 wider than body; head width 31.6 % of SVL; head width 1.1 times head length. Snout rounded in dorsal view, in lateral view rounded and slightly inclined; eye-nostril distance/eye diameter 0.6 and eye-nostril distance/interorbital distance 0.8. Loreal region slightly concaved, nostrils round, flush with surrounding skin, not elevated; internarial region concave anterodorsally; canthus rostralis not well defined (rounded). Eyes medium, directed anterolaterally, eyes angled 45.0° relative to midline (where anterior-facing eyes would be 90 ° relative to midline); eye diameter 2.6 times wider than width of disc on finger III; eye diameter 45.9 % of head length and 125.9 % of interorbital distance. Tympanum visible, not covered by skin, angled laterally with slight dorsal inclination, tympanic annulus low, supratympanic fold absent. A dentigerous process on each vomer present, each bearing a row of two vomerine teeth; choanae large, ovoid, separated more widely than nostrils; tongue round, white in preservative, anterior 3 / 4 attached to floor of mouth; vocal slits present, extending from the sides of the base of tongue to the level of the mandibular joints. Forelimbs relatively robust, with forearm flattened and roughly 2 times as wide as arm; ulnar fold absent; humeral spine in adult males present, small, evident as an elongated bump in external view. Relative length of fingers: II Coloration in life based on the type series. Dorsal and dorsolateral surfaces green with yellow tones and minute melanophores (Fig. 19). The melanophores are absent from the upper lip, hands, feet, and all fingers and toes. Upper lip rim white (enameled). Rim of eyelids bright yellow. Iris reddish-bronze with minute black reticulation and black or grey spots and a black ring delimiting the iris. Yellow pupillary ring absent. Hands and feet yellowish-green with fingers and toes yellowish. Ventral surfaces yellowish green. Bones green. Parietal peritoneum anterior half white, posterior half transparent. Hepatic and visceral (stomach and intestines) peritonea transparent. Other peritonea not visible in life. Coloration in preservative based on the type series. Dorsal and dorsolateral surfaces dull cream with minute melanophores and iridophores (Fig. 20). The melanophores are more conspicuous on the upper eyelid making it lavender and are absent from the upper lip, hands, feet, and all fingers and toes. The thin layer of iridophores of the upper lip (enameled) is almost unappreciable, most likely lost after more than two years in 70 % ethanol. Rim of eyelids conspicuously white due to iridophores. Iris black with minute greyish spots and a black ring delimiting the iris. Ventral surfaces and flanks dull cream. Parietal peritoneum anterior half white (both on the ventral and dorsal sides), posterior half transparent. Pericardium white. Hepatic peritoneum transparent. Visceral (stomach and intestines), renal, and gonadal peritonea mostly transparent but with some small and isolated traces of iridophores that seem to intrude from the dorsal portion of the parietal peritoneum. Observation made on the dissected paratype MHNC 13931. Measurements. Holotype measurements (in mm) as follows: SVL 23.4, HL 7.4, HW 8.1, TD 1.0, IND 1.3, IOD 2.7, ED 3.4, EA 44.9 °, EW 1.4, END 2.2, HaL 7.3, 3 DW 1.3, TL 12.4, SL 12.6, FL 10.6. Measurements of the complete paratype series are given in Table 2, except for MNCN 45951 (measurements not taken). Variation. This species displays remarkably little variation across its range. Two individuals from the Cordillera de Kampanquis in northern Peru (approximately 350 km NW from the type locality) exactly match the descriptions of the type specimens. The fingers and toes of this species vary from greenish-yellow to yellowishgreen. Tadpole. The following description is based on three preserved tadpoles of Gosner (1960) stages 26, 35 and 39 in order to observe developmental changes (Fig. 21). These tadpoles were from a collected egg clutch and reared in captivity. The first tadpole (stage 26) was 28.9 mm in total length; body length 9.5 mm; body width 4.6 mm; eyeeye distance 1.4 mm (30 % of body width); max. tail height 2.9 mm mid-way to tail end; tail musculature height 62 % of max. tail height; body color in life translucent white. The second tadpole (stage 35) had a total length of 35.5 mm; body length 11.3 mm; body width 5.3 mm; eye-eye distance 2.8 mm (52 % of body width); body color in life red, especially on ventral surfaces. The third tadpole (stage 39) reached a total length of 36.5 mm; body length 11.8 mm; body width 5.6 mm; eye-eye distance 3.8 mm (68 % of body width); limb length 12.1 mm; eyes directed anterodorsally; color in life grass green. General characteristics of the tadpoles are as follows: Body round, elongate (body length 2.1 times body width); eyes dorsal, becoming more lateral as development proceeds; spiracle sinistral and lateral, vent medial; tail musculature uniform white; upper and lower jaw sheaths narrow and serrated; lateral process on upper jaw medium length with distal tips slightly curled anteriorly; oral disc not emarginate; marginal papillae on ventral side complete while completely absent on dorsal side; submarginal papillae absent; labial tooth row formula 1 / 3 in all stages examined with no gaps present. Upon hatching, tadpoles of this species are typically reddish-gold and somewhat translucent. Over the course of development, they begin to turn bright pinkish-red, presumably due to increased vascularization near the skin surfaces (Fig. 21), which may be important for respiration if these tadpoles are fossorial. However, coloration of tadpoles likely depends on local environmental conditions, such as oxygen availability (Villa & Valerio 1982). As tadpoles approach metamorphosis, they begin to turn green although heavy vascularization is still apparent near the spine and buccal region. Vocalizations. The following call description comes from recordings of several different males from the Cainarachi valley at temperatures ranging from 20–24 °C ( Fig. 22). Two of these recordings were of two different choruses of males, one of which contained two males, the other of which contained at least four calling males. Cochranella guayasamini has a buzz-like advertisement call composed of one or two notes. The simpler one-note call consists of a single short buzz, 0.166– 0.208 s in length, composed of 9–10 pulses, pulse rate 48–54 pulses/s, dominant frequency 6080–6180 Hz. The two-note call includes a second buzz appended to the end of the first note, roughly 0.5 s after the end of the first note. This second note tends to be slightly longer in duration (roughly by 0.1 s) but is otherwise similar in structure to the first note. One of the most striking aspects of this species’ call is its complex calling behavior. Solitary males were sometimes encountered, but in these cases males called infrequently. For example, in one stream in Abra Tangarana (6 ° 16 ' 52.87 "S, 76 ° 43 ' 57.86 "W, 1003 m), a small number of solitary Co. guayasamini were heard, each typically calling once every 10–15 min. However, in a nearby site 3.6 km S (6 ° 18 ' 46.68 "S, 76 ° 43 ' 34.23 "W, 1120 m) where Co. guayasamini were more abundant, males called in dense aggregations and each male appeared to call far more frequently, corresponding to higher frequencies of chorusing bouts (individual males typically called every 1–5 minutes). Calling behavior also becomes quite complex in group situations, as males tend to call in distinct bouts. We have one recording of a calling bout between two males, each making a two-note call (Fig. 22). The second male starts his call immediately after (and slightly overlapping with) the first note of the first male’s call, while the second notes of each male more broadly overlap. This situation becomes far more complex in larger groups (commonly with 3–6 calling males), as individual males may be making either one- or two-note calls. It seems that one male serves as the chorus leader, and his call is rapidly followed by a cascade of calls from other nearby males (Fig. 22). In the field, this gives the impression of a continuous ‘wave’ throughout the chorus, making it extremely difficult to locate individual calling males. Similar behavior in Co. granulosa was studied in detail by Ibáñez (1991), who reported that males rapidly called in response (within 816 ms) to a neighbor’s call and overlaped their calls 92–100 % of the time (17–85 % of all notes). Chorusing behavior in Co. guayasamini, Co. erminea, and Co. granulosa seems similar in that males call once in discrete bouts (presumably responding to neighbor’s call), then remain silent until the next calling bout. In Co. granulosa, calling bouts can become calling waves traveling along the stream up to 600 m (Ibáñez 1991). Since high levels of call overlap can actually degrade the temporal information of the call and reduce males’ ability to attract mates in Co. granulosa (Ibáñez 1991), call overlap among these Cochranella may serve an anti-predator function interfering with detection (Ryan 1986). Distribution and ecology. Cochranella guayasamini occurs in the east-Andean versant of Peru from roughly 7 ° south latitude to 4 ° south latitude, at elevations between 250 and 1150 m (Fig. 18). In northern San Martín department (in the vicinity of Tarapoto), this species is commonly found in a wide variety of habitat types, from highly disturbed areas to primary forest. The highest density site we have found was at 1120 m near Abra Tangarana, in a stream flowing through a cattle pasture with low bushes growing along the edges. Prior to August 2011, this species was only known from northern San Martín. A record by Catenazzi and Venegas (2012) from the Cordillera de Kampanquis in northern Peru extends the known distribution of this species roughly 400 km NW from the type locality. Near San Jose we monitored 10 clutches of Cochranella guayasamini over several nights (typically involving 2–3 checks per night), and did not observe any parental behavio

Teratohyla amelie (Cisneros-Heredia & Meza-Ramos 2007) Figure 27 Cochranella amelie Cisneros-Heredia and Meza-Ramos 2007. Holotype DHMECN 3066, an adult male from ���Comunidad de Ogl��n, Cant��n Arajuno, Provincia de Pastaza, Rep��blica del Ecuador (01�� 18 ��� 65 ���S, 77 �� 42 ��� 41 ���W, 600 m elevation)��� Teratohyla amelie��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil�� 2009. Background information. This species was described on the basis of two adult males collected from a locality in the east-Andean foothills of central Ecuador. The combination of white visceral peritonea, green bones in life, transparent parietal peritoneum, humeral spines absent in adult males, bulbous liver, and dorsum lavender in preservative is unique among known centrolenid species. Guayasamin et al. (2008) provided molecular sequence data for a specimen identified as ��� Cochranella cf. amelie ��� from southern Peru, and Guayasamin et al. (2009) placed this species in the genus Teratohyla on the basis of genetics and morphology. New data. We have collected five specimens from two sites in northern Peru. Additionally, we have studied two specimens from southern Peru (one of them mentioned by Guayasamin et al. 2008). These seven specimens fit the description of Teratohyla amelie, which have a completely transparent parietal peritoneum, a character state only present in species of Chimerella, Hyalinobatrachium, and Teratohyla pulverata, Vitreorana antisthenesi, V. gorzulae, V. uranoscopa, V. eurygnatha, and V. pa r v u l a. Teratohyla amelie has green bones versus white in all species of Hyalinobatrachium, except for H. taylori from which it is distinguished by a rounded snout in profile (sloping in H. taylori). Males of Chimerella and V. gorzulae have humeral spines (absent in T. amelie). Vitreorana uranoscopa, V. eurygnatha, and V. parvula have a white urinary bladder peritoneum (transparent urinary bladder peritoneum in T. amelie). Teratohyla pulverata has a sloping snout in profile and scalloped, white ulnar and tarsal folds (snout rounded in profile and scalloped, white ulnar and tarsal folds absent). Four specimens (MNCN 45959 through MNCN 45962) were collected from a small stream running through an overgrown cacao plantation, roughly 92 km N of Bagua, 2.3 km south (airline) from the village of Huabico (4 �� 57 ' 16.59 "S, 78 �� 12 ' 39.92 "W, 349 m). An additional specimen (MNCN 45958) was found in southern San Mart��n department, Peru, in a stream on the Juanjui-Tocache road, near the village of Nuevo Jaen (7 �� 39 ' 16.20 "S, 76 �� 39 ' 24.60 "W, 591 m). The other two specimens (MHNC 5646 and MNCN 44212) were collected on February 1, 2007 by J. C. Chaparro, J. M. Padial, I. De la Riva, and SC-F at a stream crossing the road at 10 km from Quincemil towards Puerto Maldonado (13 �� 12 ���03.6������ S, 70 �� 40 ��� 28.9 ������ W; 572 m), Provincia Ouspicanchis, Departamento Cusco. These localities narrow the distributional gap between the specimens from Cusco, Peru and those from Ecuador. Vocalizations. We have not been able to obtain recordings of this species, although we observed males calling in Huabico and Cusco (ET and SC-F respectively). The call sounded similar to T. midas, consisting of a short ���chirp���, which sounded roughly similar (by ear) to the T. midas call described below. All individuals (including calling males) were found on the upper surfaces of leaves, roughly 2���3 m above the stream. Distribution and ecology. Considering the additional records provided above, Teratohyla amelie is distributed in the east-Andean foothills (349���600 m) from Central Ecuador to Central Peru, roughly between 1 �� and 7 �� south latitude, and with an isolated record in southern Peru. In Peru, T. amelie has been found alongside Cochranella erminea., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 53-54, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Cisneros-Heredia, D. F. & Meza-Ramos, P. (2007) An enigmatic new species of glassfrog (Amphibia: Anura: Centrolenidae) from the Amazonian Andean slopes of Ecuador. Zootaxa, 1485, 33 - 41.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. & Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595. http: // dx. doi. org / 10.1016 / j. ympev. 2008.04.012"]}

Cochranella erminea Torres-Gastello, Su��rez-Segovia, and Cisneros Heredia 2007 Figure 17 Cochranella erminea Torres-Gastello, Su��rez-Segovia, and Cisneros-Heredia 2007. Holotype MUSM 24056, collected from ���Sabetari stream (11 �� 14 ' 31 " S, 73 �� 31 ' 33 " W, 370 m), on the Tambo River Basin, 9 km NNE of the indigenous community of Quitepampani 2, province of Satipo, department of Jun��n, Peru.��� Rulyrana erminea��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil�� 2009. Rulyrana erminae��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil�� 2009. Incorrect subsequent spelling. 2. The original description states that the type locality is " [...] 9 km NNE of the indigenous community of Quitepampani, province of Satipo, department of Jun��n". Kitepampani (correct spelling) is a Machiguenga community located in Cusco, not Jun��n. The type locality is situated in the Mayapo community, which is an Ash��ninka community. Cochranella erminea��� Castroviejo-Fisher, Guayasamin, Gonzalez-Voyer, and Vil�� 2014. Background information. This species was described on the basis of a single adult female collected from Departamento Jun��n in south-central Peru (Torres-Gastello et al. 2007), and was distinguished from other centrolenid species by its combination of extensive webbing between fingers III and IV, transparent visceral peritonea, and dorsum covered in abundant white enameled spots with scattered dark spots. The authors considered the placement of this species in the genus Cochranella tentative, as no males were available and thus the condition of the humeral spine was unknown at the time of description. Guayasamin et al. (2009) moved this species to the newly-erected genus Rulyrana, as more recent observations indicated that males of this species lack humeral spines. Mercado (2012) added four specimens from three new localities from the department of Cusco, Peru, extending the range of this species 111.5 km to the south and up to 870 m elevation. New data. In February 2010, during field work in northern Peru, ET, M. Pepper, M. Sanchez-Rodriguez, and J. Tumulty encountered two individuals of Cochranella erminea, confirmed by comparisons of morphology, bioacoustics, and phylogenetics (Figs. 2 c and 17). These specimens were collected from two sites north of Bagua, Amazonas, Peru. The first site was a small stream running through an overgrown cacao plantation, roughly 92 km N of Bagua, 2.3 km south (airline) from the village of Huabico, Amazonas (4 �� 57 ' 16.59 "S, 78 �� 12 ' 39.92 "W, 349 m). The second site was roughly 49 km N of Bagua, 14 km north (airline) from the Pongo de Rentema near the town of Muyo (5 �� 17 ' 53.87 "S, 78 �� 24 ' 1.99 "W, 629 m), in a small stream with a relatively open forest cover. Three months later, we (SC-F and ET) returned to this area and encountered another two specimens in a relatively large stream flowing through undisturbed forest, close to the second site mentioned above (5 �� 18 ' 3.86 "S, 78 �� 23 ' 44.57 "W, 682 m). These records extend the known distribution of Co. erminea 875 km northwest from the type locality. In 2011, we filled in this 875 km sampling gap with a single specimen of Co. erminea (CORBIDI 10477) taken from a site 28 km S (airline) from the city of Juanjui, San Mart��n, Peru (7 �� 25 ' 38.62 "S, 76 �� 39 ' 53.28 "W, 366 m), which is located 545 km NW from the type locality. This was an adult male observed calling from a leaf 4 m above the edge of a waterfall. An additional record (uncollected specimen) of Co. erminea was obtained from the R��o Shunt�� drainage, 26.2 km SW from the city of Tocache, San Mart��n, Peru (8 �� 19 ' 25.68 "S, 76 �� 42 ' 35.82 "W, 923 m) on 12 March 2012 by JD and ET. Variation. Cochranella erminea exhibits moderate morphological variation over its large range. While our records from near Juanjui and Tocache appear nearly identical to material from the type locality, individuals from northern Peru are notable in that they have a distinctly pinkish-red iris, and they show variation in the number and distribution of dorsal spots (Fig. 17). Furthermore, the white enameled spots as mentioned in the holotype are less notable in these northern populations. Vocalizations. The following call description is based on a single call from a male recorded by SC-F and ET on 3 April 2010 from Pongo de Rentema (5 �� 18 ' 3.86 "S, 78 �� 23 ' 44.57 "W, 682 m), temperature 21.4 ��C. The call consists of two harsh buzz-notes given in quick succession. The first of these notes starts quietly and increases in amplitude throughout, with a duration 0.313 s, 14 pulses, dominant frequency 4870 Hz. This is followed by a single ���chirp��� given 0.173 s after the first note. This chirp is 0.054 s in duration, with 3 pulses, dominant frequency 4866 Hz. In the field, this sounds like a ���zzzZZZT ZT���. We have heard calls from other populations (i.e., Juanjui and the type locality), and the calls all sound similar to the ear, with the only noticeable difference is that the number of trailing chirps can vary from zero to two. This call is similar to the call of Cochranella granulosa, which has a ���pulsed primary note followed by 0���5 shorter pulsed secondary notes��� (Ib����ez 1991). The call of Co. guayasamini new species (see below) is superficially similar to Co. erminea, except that Co. guayasamini tends to use a two-note call only in groups (often just one note when alone), whereas Co. erminea typically uses a two-note call even when alone. In addition, in Co. guayasamini the first and second notes are equal in duration, and not amplitude modulated (vs. first note amplitude modulated and longer than second note in Co. erminea). Distribution and ecology. This species is widely distributed throughout the east-Andean versant in Peru, occurring from roughly 11 �� south latitude to 5 �� south latitude, at elevations between 349 and 923 m (Fig. 18). However, Cochranella erminea appears to be patchily distributed within this region. For example, this species has not been registered from the mountains surrounding Tarapoto, Peru, despite frequent searches in this area. Some of this patchiness may be attributable to the presence of the closely related Co. guayasamini. These two species are sister taxa, have similar calls, occur at similar elevations, and appear to overlap in the montane regions extending from northern San Mart��n to central Amazonas. However, these two species have never been found together in the same site (N = 7 sites for Co. erminea and N = 8 sites for Co. guayasamini). It is possible that competitive exclusion or microhabitat preferences could be important regulators of community structure at small spatial scales. Cochranella erminea occurs in a wide variety of habitats ranging from primary premontane forest to relatively disturbed, lowland foothill habitats. The site where we have witnessed the highest density of this species was an overgrown cacao plantation at 349 m. The only notable behavior we have observed in this species has been male calling activity. Males call from the tops of leaves, typically in groups of 2���5 individuals, although solitary males have also been observed calling. When calling in groups, males seem to stagger their calls sequentially and calls of different males do not typically overlap. At one site, males called sequentially downstream, with the chorus starting with an upstream male and finishing with a downstream male. See details below on Co. guayasamini for noted similarities among Cochranella. We observed two egg clutches near a chorus of Co. erminea males that correspond to Cochranella eggs; a globular cluster of pigmented eggs dangling from leaf tips. At the type locality as well as our record from near Juanjui, Cochranella erminea was the only centrolenid species registered (Torres-Gastello et al. 2007). In the R��o Shunt�� site, Co. erminea was found alongside Hyalinobatrachium carlesvilai and Rulyrana cf. flavopunctata. In northern Peru, Co. erminea has been found alongside Centrolene charapita, Teratohyla amelie, T. midas, and Rulyrana mcdiarmidi. Taxonomic remarks. Based on results from our phylogenetic analysis (Fig. 2 c), Cochranella erminea is wellsupported as a member of the genus Cochranella (JK = 100; GB = 16) and is similar in overall morphology to Co. granulosa. Furthermore, this species shows substantial similarity in advertisement call to other members of Cochranella, most notably Co. granulosa and Co. guayasamini., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 27-29, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Torres-Gastello, C. P., Suarez-Segovia, J. & Cisneros-Heredia, D. F. (2007) Cochranella erminea, a new species of Centrolenidae (Amphibia: Anura: Athesphatanura) from Amazonian Peru. Journal of the National Museum (Prague), Natural History Series, 176, 1 - 12.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Castroviejo-Fisher, S., Guayasamin, J. M., Gonzalez-Voyer, A. & Vila, C. (2014) Neotropical diversification seen through glassfrogs. Journal of Biogeography, 41, 66 - 80. http: // dx. doi. org / 10.1111 / jbi. 12208","Mercado, D. A. R. (2012) Rulyrana erminea (Torres-Gastello, Suarez-Segovia and Cisneros-Heredia, 2007) (Anura: Centrolenidae): Distribution and range extension, Peru. Check List, 8, 894 - 895.","Ibanez, R. (1991) Synchronized calling in Centrolenella granulosa and Smilisca sila (Amphibia, Anura). Dissertation. University of Connecticut. [total page number unkown]"]}

Hyalinobatrachium pellucidum (Lynch & Duellman 1973) Figure 26 Centrolenella pellucida Lynch and Duellman 1973. Holotype KU 143298, an adult female collected from ���Rio Azuela, 1740 m, Quito-Lago Agrio road, Provincia Napo, Ecuador ���. Hyalinobatrachium pellucidum��� Ruiz-Carranza and Lynch 1991 Hyalinobatrachium lemur Duellman and Schulte 1993 Background information. This species was described from a single adult female (Lynch & Duellman 1973) based on the following characters: (1) heart visible in life (at the time, a state known only in two other species of Ecuadorian centrolenids), and (2) ulnar, anal, and tarsal folds present. Twenty years later, Duellman and Schulte (1993) described Hyalinobatrachium lemur from the east-Andean versant of San Mart��n, Peru. They diagnosed H. lemur from H. pellucidum on the basis of skin texture (smooth in H. lemur vs. shagreened in H. pellucidum) and the extent of webbing on Finger IV (webbing more extensive in H. lemur). Castroviejo-Fisher et al. (2009) reviewed the type series of both H. pellucidum and H. lemur, as well as additional fresh specimens of the former species both from Ecuador and southern Peru. They concluded that the characters originally used to diagnose H. lemur fell within the range of variability exhibited by H. pellucidum. Thus, they designated H. lemur as a junior synonym of H. pellucidum and identify the paratype as a H. carlesvilai. New data. Specimens have now been found in the Cainarachi valley near Tarapoto, San Mart��n, Peru (6 �� 25 ' 16.7 ''S, 76 �� 17 '' 28.5 W, 523 m). Wen et al. (2012) referred these specimens to H. pellucidum and also provided a description of the advertisement call, which is described as a single, monotone note without amplitude modulation. Also, a gravid female MNCN 45953 was collected on 28 March 2010 by M. Guerra Panaijo and Santiago Castroviejo-Fisher at Puente Chino on the Tocache-Pucallpa road, approximately 29 km NE from Tingo Maria (9 �� 8 ' 9.03 "S, 75 �� 47 ' 15.78 "W, 1147 m), Ucayali, Peru, where it is sympatric with H. carlesvilai. The specimen was found on the upper side of a tree leaf ~ 4 m above the water level. This is the first report of this species for Ucayali. The identity of this specimen was confirmed by morphology and phylogenetic analysis (Fig. 2 a). Natural history. Spider predation was observed in several clutches from a single male���s territory at the San Jose locality mentioned above. During a brief visit to the study stream on 15 May 2011 a male was encountered brooding a newly deposited clutch at 22: 57. We returned again on May 24 th, finding that the male had sired an additional 2 clutches all positioned next to each other on the lower surfaces of a leaf. At 22:00 a small spider was observed on the same leaf as the male and his egg clutches (see Fig. 26 for the following sequence). We lightly tapped the spider with a blade of grass on the abdomen. The spider then moved towards the oldest clutch, immediately removed an embryo from the clutch with its chelicerae and began consuming the embryo. The male, who was positioned on the opposite side of the clutches, moved over the eggs towards the spider with roughly half of his body positioned over two clutches. The male remained in this position for ca. 2 minutes, even when the spider came in contact with the male���s hind limb. After ca. 2 min the male moved back on top of a clutch and began brooding while the spider continued consuming the embryo. We returned to this site again on 30 May, finding that the male had abandoned his territory (or was predated), and that the spider had built a web over the clutches (Fig. 26). The spider was observed to consume embryos during 4 nights until 2 June, when all embryos had either hatched or were predated. Over this period the spider was found at the nest site during both day and night, but was only observed consuming embryos at night., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 44-45, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Lynch, J. D. & Duellman, W. E. (1973) A review of the centrolenid frogs of Ecuador, with descriptions of new species. The University of Kansas Museum of Natural History Occasional Papers, 16, 1 - 66.","Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33.","Wen, A., Vasquez, N. & Castroviejo-Fisher, S. (2012) Description of the previously unknown advertisement calls of Hyalinobatrachium fragile, H. pellucidum, and Vitreorana antisthenesi (Amphibia: Centrolenidae). Zootaxa, 3480, 80 - 87."]}

Rulyrana saxiscandens (Duellman & Schulte 1993) Figures 28, 29 Cochranella saxiscandens Duellman and Schulte, 1993. Holotype KU 211779, an adult male from ���Catarata Ahuashiyacu, (06�� 30 ��� S, 76 �� 20 ��� W, 730 m), 14 km (by road) northeast of Tarapoto, Provincia San Mart��n, Departamento San Mart��n, Per�� ��� Cochranella tangarana Duellman and Schulte, 1993. New synonym. Cochranella croceopodes Duellman and Schulte, 1993. New synonym. ��� Cochranella ��� croceopodes��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil�� 2009. Rulyrana saxiscandens��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil��, 2009. Rulyrana tangarana��� Guayasamin, Castroviejo-Fisher, Trueb, Ayarzag��ena, and Vil��, 2009. Background information. Cochranella saxiscandens was described from a series of 30 adult males collected from Ahuashiyacu waterfalls (730 m), 14 km NE from Tarapoto, San Mart��n, Peru (Duellman & Schulte 1993). Cochranella tangarana was described on the basis of two adult males and one adult female, all collected from Abra Tangarana (1080 m), San Mart��n, Peru, a site roughly 50.8 km W from Ahuashiyacu. Both species were allocated to the Cochranella ocellata group due to the combination of white parietal peritonea with clear visceral peritonea. Additional characters of these two species include vomerine teeth present, bones green, dorsal skin finely spiculate, humeral spine absent, and color in life dark green to black. Duellman & Schulte (1993) distinguished these two species from each other based on color in preservative (dark grey to black in R. saxiscandens vs. lavender in R. tangarana), presence/absence of melanophores on ventral surface of shanks and tarsi (melanophores present in R. saxiscandens vs. absent in R. tangarana), and presence/absence of inner tarsal fold (fold present in R. saxiscandens vs. absent in R. tangarana). Both species were removed from Cochranella and placed in the genus Rulyrana by Guayasamin et al. (2009) on the basis of overall similarity with other species of the genus. Cochranella croceopodes was described on the basis of two adult females, the holotype collected from km. 23.2 on the Tarapoto-Yurimaguas road and the paratype collected from Ahuashiyacu waterfalls, both San Mart��n, Peru (Duellman & Schulte 1993). This species was assigned to the Co. granulosa group due to the combination of white parietal and visceral peritonea. Cochranella croceopodes was described as having a ���diffuse yellow line on flanks from axilla to groin��� (Duellman & Schulte 1993) and diagnosed only against species having dorsolateral stripes; i.e., Centrolene hesperium and Ce. lemniscatum, both of which lack vomerine teeth and have first finger shorter than the second. Cochranella croceopodes was moved from the granulosa group to the spinosa group (Ruiz-Carranza & Lynch 1995), then back to the granulosa group (Guayasamin & Bonnacorso 2004). Cisneros- Heredia and McDiarmid (2007) removed it from the granulosa group but did not assign it to any group. Guayasamin et al. (2009) left Co. croceopodes as incertae sedis within the subfamily Centroleninae. New data. In 2010 and 2011 we collected a series of 10 R. saxiscandens from two sites: three males (CORBIDI-HE- 2012-14150, CORBIDI-HE- 2012-14151, and CORBIDI-HE- 2012-14153) and three females (CORBIDI-HE- 2012-14149, CORBIDI-HE- 2012-14152, and ET- 10-124) from the type locality of Ahuashiyacu (6 �� 27 ' 19.69 "S, 76 �� 18 ' 32.33 "W, 797 m); and four males (ET- 10-126, ET- 10-127, CORBIDI 14668, CORBIDI 10464) from 2.3 km E of the type locality (6 �� 27 ' 30.28 "S, 76 �� 17 ' 14.35 "W, 988 m, Figs. 7, 28). In addition, we collected four R. tangarana males, three of which were from the type locality (6 �� 16 ' 52.86 "S, 76 �� 43 ' 57.86 "W, 1047 m), and one from 6 km SE from the type locality (6 �� 19 ' 22.40 "S, 76 �� 41 ' 43.55 "W, 1003 m, Figs. 7, 28). Morphological, bioacoustic, and genetic data all strongly suggest the existence of a single species. With respect to color in preservative, of the 10 R. saxiscandens, two (CORBIDI 10464 and CORBIDI 14668) were lavender in preservative; seven (CORBIDI-HE- 2012-14149 through CORBIDI-HE- 2012-14153, ET- 10-124, ET- 10-126, and ET- 10-127) ranged from bright green or yellowish green to dark or dull green in preservative, and one individual (CORBIDI-HE- 2012-14151) was completely black in preservative. In contrast with Duellman and Schulte (1993), none of the R. tangarana were lavender in preservative but rather dull green. With respect to melanophore presence/absence on the ventral surface of the tarsi and shanks, of the 10 R. saxiscandens, two (ET- 10-126 and ET- 10-127) entirely lacked melanophores in this area, six (CORBIDI-HE- 2012-14149 through CORBIDI-HE- 2012- 14151, ET- 10-124, MNCN 45962, and CORBIDI 14668) had faint melanophores restricted to the edges of the leg (barely extending to ventral surface), and two (CORBIDI-HE- 2012-14152 and CORBIDI-HE- 2012-14153) had distinct melanophores present. Of the four R. tangarana, two individuals (CORBIDI 14149 and CORBIDI 10476) completely lacked melanophores in this area. The remaining two individuals (CORBIDI 10478 and CORBIDI 14151) had melanophores present only on the leg edges, similar to six of the R. saxiscandens mentioned above. Finally, with respect to presence/absence of the inner tarsal fold, of the 10 R. saxiscandens, six individuals (CORBIDI-HE- 2012-14151, CORBIDI-HE- 2012-14153, ET- 10-124, ET- 10-127, CORBIDI 10464, and CORBIDI 14668) had a distinct inner tarsal fold, while one individual (ET- 10-126) lacked a fold completely. The remaining three individuals (CORBIDI-HE- 2012-14149 through CORBIDI-HE- 2012-14151) have very faint folds, mainly present as a row of yellowish discoloration extending from the base of the foot along the inner surface of the tarsus. In the series of four R. tangarana, one individual from 6 km SE from the type locality (CORBIDI 10478) clearly has an inner tarsal fold. Two individuals (CORBIDI 10476 and CORBIDI 14151) from the type locality do not. One individual (CORBIDI 14149) from the type locality has a discoloration along the inner tarsus, similar to three of the R. saxiscandens, but lacks a fold. Specimens from the type locality of R. saxiscandens display considerable variation in dorsal coloration in life. In a series of five individuals collected in a single night, one individual (CORBIDI-HE- 2012-14153) was black (Fig. 28 a), one was bluish-gray (ET- 10-124), one dull green (CORBIDI-HE- 2012-14152), and two lime-green (CORBIDI-HE- 2012-14149 and CORBIDI-HE- 2012-14151) (e.g., Fig. 28 c). Four individuals (ET- 10-126, ET- 10- 127, CORBIDI 10464, and CORBIDI 14668) found 2.3 km E of the type locality ranged from black to bright green; furthermore, two of these individuals (CORBIDI 10464 and CORBIDI 14668) had faint lime-green flecks on the dorsum. The four specimens collected in or near the type locality of R. tangarana, dorsal coloration ranged from dull to bright green. Two (CORBIDI 14149 and CORBIDI 10476) had prominent lime-green flecks on the dorsum, which appear to vary in number from just a few to consistently scattered throughout the dorsum. Much of this variation may be due to the apparent ability for individuals to change color. Two individuals (CORBIDI 10464 and CORBIDI 14668) from 2.3 km E of Ahuashiyacu were lime green upon collection but turned nearly black by the following morning. Iris color also varies from dark bronze to light bronze (Fig. 28), and appears to be related to dorsal color (with darker individuals tending to have darker irises). Skin texture of R. saxiscandens from the type locality varies from spiculate to shagreened to nearly smooth. The two specimens from the R. tangarana locality both had spiculate skin. There is variation in the extent to which iridophores cover the parietal peritoneum among R. saxiscandens specimens and those collected from the type locality of R. tangarana. In some individuals the iridophores cover nearly the entire venter (e.g., Fig. 28 b) whereas in others they are restricted to only the anterior ~ 1 / 2 of the venter (e.g., Fig. 28 f). Additionally, there is some variation in the prominence of the inner tarsal fold (see above) and extent of melanophores on ventral surfaces of limbs (see above). Calls obtained from both the type locality of R. tangarana and R. saxiscandens are nearly identical. The call consists of a short, high-pitched ���zip��� given every few seconds. We obtained a series of six calls from a single male at the type locality of R. tangarana, recorded 6 June 2011 at 21.5 ��C. Note length was 0.038��� 0.05 s (mean = 0.046 s), each note was composed of 7 or 8 pulses (mean = 7.8 pulses), pulse rate 160���184 pulses/s (mean = 170.3 pulses/ s), dominant frequency 6103���7106 Hz (mean = 6599 Hz). We also recorded calls of three R. saxiscandens (two from Ahuashiyacu and one from 2.3 km E Ahuashiyacu) at temperatures ranging from 20���23 ��C. Calls were similar to R. tangarana calls; note length 0.028��� 0.056 s (mean 0.041 s), notes composed of 7���12 pulses (mean = 8.2 pulses), pulse rate 179���250 pulses/s (mean = 205.6 pulses/s), dominant frequency 6568���6915 Hz (mean = 6833 Hz). While there are slight differences between the two nominal taxa (e.g., when comparing means), the amount of interspecific variation is in most cases is equal to or lower than intraspecific or even intra-individual variation (i.e., different notes given by a single male). Thus, there are no fixed, discernible differences in the calls of these two taxa. Our new molecular data reveals that Rulyrana mcdiarmidi, R. tangarana, R. saxiscandens, and an individual referred to as R. flavopunctata (Guayasamin et al. 2008) from Provincia Morona-Santiago, Ecuador are not reciprocally monophyletic (Fig. 2 c). The specimen R. tangarana CORBIDI 14149 is retrieved as sister to all other specimens, which are grouped in a polytomy. Within this polytomy, there are eight single terminals corresponding to specimens of R. saxiscandens and R. tangarana, and three clades. The first clade groups two specimens of R. saxiscandens (CORBIDI-HE- 2012-14150 and CORBIDI-HE- 2012-14151), the second clade groups another three individuals of R. saxiscandens (ET- 10-124, CORBIDI 14668, and CORBIDI-HE- 2012-14152) in a polytomy, and the third clade groups three specimens of R. mcdiarmidi (CORBIDI-HE- 2010-6848, CORBIDI 10470, and CORBIDI 10473) with the specimen of R. flavopunctata QCAZ 32265. Intra- and interspecific genetic distances among these terminals are low (p-distances = 0.0��� 0.6 %) and overlapping (Table 6). Comparisons. Based on the above data, none of the original diagnostic characters between these two species are supported. Color in preservative is variable both within and among populations. Furthermore, none of the Rulyrana tangarana specimens turned lavender in preservative (as mentioned in the original diagnosis, although our specimens were fixed in ethanol and not in formalin, but this did not prevent other specimens from turning lavender). Presence/absence of melanophores on the ventral surfaces of the tarsi and shanks is variable within populations and is not fixed in either type locality of the nominal taxa. Finally, the inner tarsal fold appears to vary within populations and is also likely influenced by preservation. For example, CORBIDI 14668, (a R. saxiscandens from the type locality), has the most prominent inner tarsal fold of any of the specimens mentioned above. This individual also appears to be the most desiccated and also has prominent dorsolateral ridges, which were not visible in life. Thus, the prominence of the inner tarsal fold was likely exaggerated in this specimen due to desiccation. The modification on ulnar and tarsal folds due to preservation artifacts in centrolenids was already mentioned by Kok and Castroviejo-Fisher (2008). Uncorrected pairwise genetic distances among DNA sequences of a fragment of 16 S also indicate that the two populations are genetically similar (p-distances 0.0��� 0.4 %, see Table 6). Furthermore, as they were not recovered as reciprocally monophyletic in the phylogenetic analysis (Fig. 2 c), we consider them to be conspecific. In the original description, Duellman and Schulte (1993) did not compare Cochranella croceopodes with either Rulyrana saxiscandens or R. tangarana. We studied in detail the types of Co. croceopodes (Fig. 29), as well as all available photos of the live types deposited at KU (see Duellman & Schulte 1993). We found that: (1) there are problems in the original description of C. croceopodes and (2) the types are extremely similar to the syntopic R. saxiscandens, which is similar to R. tangarana (see above). Below, we list and discuss the ways in which the original description of Co. croceopodes is inconsistent with our own observations on the type specimens and how they fall within the variability observed in R. saxiscandens and R. tangarana. The visceral peritoneum of Cochranella croceopodes was described as white, but our examination of the holotype revealed that the visceral peritoneum is transparent (Fig. 29), which is also the character state in all specimens of R. saxiscandens and R. tangarana examined. Cochranella croceopodes was described as having a "diffuse yellow line present on flanks from axila to groin" Duellman and Schulte (1993: 21). The diffuse yellow line on flanks is not appreciable in the preserved types as described in Duellman and Schulte (1993: 21). Based on the examination of the photos of the live types, we interpret that this diffuse yellow line is formed by the absence of melanophores on the pigments of the flanks (which are a continuation of the dorsal pigments). A similar diffuse yellow line is also present in two (CORBIDI-HE- 2012-14149 and CORBIDI-HE- 2012-14151) of the newly collected specimens of R. saxiscandens (Fig. 28 c). We interpret this character as variability in coloration. Cochranella croceopodes had shagreened dorsal skin in the description (Duellman & Schulte 1993, Fig. 2, upper left); however, it appears smooth after 23 years in ethanol (Fig. 29). Most specimens of Rulyrana saxiscandens have spicules in dorsal surfaces but their distribution and density varies substantially to the point that females of our newly collected specimens have shagreened skin with no spicules. We interpret differences in this character as variability in dorsal skin texture, sexual dimorphism, and effects of preservation. Cochranella croceopodes was described as having a snout ���nearly truncate in profile��� but examination of the holotype reveals that it is round (Fig. 29) as in R. saxiscandens. Lastly, examination of photos of the live types of Co. croceopodes taken in situ by the authors revealed that those specimens show the same overall coloration pattern, skin texture, and habitus as the newly collected specimens of R. saxiscandens and R. tangarana. Taxonomic remarks. Based on available morphological, bioacoustic, and phylogenetic evidence, we hereby consider Cochranella croceopodes, Rulyrana saxiscandens, and Rulyrana tangarana synonymous. Under the Principle of the First Reviser of the ICZN (article 24.2.2) we give precedence to saxiscandens, rendering Rulyrana tangarana and Cochranella croceopodes junior synonyms of Rulyrana saxiscandens. We also recognize that Cisneros-Heredia & Guayasamin (in press) arrived at a similar conclusion regarding the synonymy of R. saxiscandens and R. tangarana. Distribution and ecology. In addition to the four localities mentioned above, we have also obtained photographic records of this species from two sites in the Cainarachi valley north of Tarapoto (Fig. 7): near the village of Santa Rosa (6 �� 24 ' 44.79 "S, 76 �� 19 ' 2.09 "W, 685 m) and San Jose (6 �� 25 ' 14.67 "S, 76 �� 17 ' 28.47 "W, 517 m). Considering these nominal taxa as a single species (see ���Taxonomic remarks���), this species is distributed in the Cordillera Escalera, roughly between the cities of Tarapoto and Moyobamba, at elevations of 517���1047 m. This species occurs in habitats ranging from small creeks to torrential streams (e.g., Ahuashiyacu) to spray zones of waterfalls. This species has never been registered from heavily disturbed habitat, but does occur in secondary forest. At the type locality of Ahuashiyacu, we have noticed that individuals encountered in the streamside vegetation tend to be bright green whereas those collected from the rocks (particularly near the spray zone of the waterfall) tend to be dark green or black. Members of the genus Rulyrana are thought to be primarily rock-breeding species (Cisneros-Heredia et al. 2008), but we found that they often call from vegetation along rocky streams (i.e., our observations in both R. mcdiarmidi and R. saxiscandens). Observed variation in coloration among individuals reflected microhabitat use when collected (vegetation vs. rocks), suggesting that the ability to change color may be a form of crypsis to aid in dynamic microhabitat use. Neither eggs nor tadpoles have been found from this species. Adults are active in a variety of microhabitats such as small trees (calling males have been heard as high as ~ 4 m), streamside vegetation, and rocks along streams., Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 49-53, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Guayasamin, J. M. & Bonaccorso, E. (2004) A new species of glass frog (Centrolenidae: Cochranella) from the lowlands of northwestern Ecuador, with comments on the Cochranella granulosa group. Herpetologica, 60, 485 - 494. http: // dx. doi. org / 10.1655 / 03 - 74","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82.","Cisneros-Heredia, D. F., Delia, J., Yanez-Munoz, M. H. & Ortega-Andrade, H. M. (2010) Endemic Ecuadorian glassfrog Cochranella mache is Critically Endangered because of habitat loss. Oryx, 44, 114 - 117. http: // dx. doi. org / 10.1017 / s 0030605309990640","Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. & Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595. http: // dx. doi. org / 10.1016 / j. ympev. 2008.04.012","Kok, P. J. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaieteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53."]}

Hyalinobatrachium anachoretus new species Figures 24, 25 Holotype. CORBIDI 10472, adult male collected from a stream on the east slope of Abra Patricia (the divide between the R��o Mara����n and R��o Huallaga drainages), 2.2 km by road NE from Puente Nieva, Provincia Rioja, Departamento San Mart��n, Peru, 5 �� 40 ' 39.06 "S, 77 �� 46 ' 23.99 "W, 2001 m, collected by E. Twomey, T. Kosch, and M. Panaijo on 5 May 2011. Paratopotype. CORBIDI 10462, adult male, same locality and collection data as holotype. Generic placement. The new species is placed in the genus Hyalinobatrachium on the basis of our phylogenetic analyses that included complete or partial sequences of the mitochondrial genes 12 S and 16 S. Additionally, the following phenotypic characters typical of Hyalinobatrachium (Guayasamin et al. 2009) are present in the new species: (1) humeral spine absent in adult males; (2) digestive tract and bulbous liver covered by iridophores (white); (3) ventral parietal peritoneum and pericardium completely transparent; (4) white bones in life; (5) when present, nuptial pads small and restricted to the medial edge of Finger I in males; (6) dentigerous process of the vomer and vomerine teeth absent; (7) males seen vocalizing from the undersides of leaves; and (8) eggs deposited on the undersides of leaves. However, the value of these characters as synapomorphies for Hyalinobatrachium needs to be tested. Diagnosis. The following combination of characters can distinguish Hyalinobatrachium anachoretus from other species in the family: (1) dentigerous process of the vomer and vomerine teeth absent; (2) snout truncate in dorsal and lateral view; (3) tympanum barely visible with equal coloration and granulation as surrounding skin; (4) dorsal skin weakly granular on body, slightly more granular on lateral regions of the head; (5) ventral skin granular, cloacal ornamentation absent, paired round tubercles below vent absent; (6) parietal peritoneum transparent; heart and urinary bladder lacking iridophores (= uncovered), kidney with iridophores on medial third, lateral two-thirds of kidney uncovered; liver and viscera mostly covered by iridophores except for inferior surface of liver and the adjacent surface of the intestine which are uncovered; testes completely covered by iridophores; (7) liver bulbous; (8) humeral spine absent in adult males; (9) finger webbing absent between fingers I and II, basal webbing between Fingers II and III, III 2 - 2 - IV; (10) toe webbing I 1 + - 2 - II 1 + - 1 + III 1 - - 1 + IV 1 + - 1 - V; (11) fringe on postaxial edge of finger IV present, metacarpal fold and ulnar fold present and enameled, metatarsal and tarsal fold present and enameled; (12) nuptial excrescence present as a small pad on medial edge of Finger I (Type V), prepollex not enlarged; prepollical spine not projecting (spine not exposed); (13) when appressed, finger I longer than II; (14) diameter of eye roughly 2.2 times wider than disc on Finger III; (15) coloration in life: dorsal surfaces apple green with small yellow spots and minute melanophores, bones white; (16) coloration in preservative pale yellow, dorsum scattered with dark melanophores; (17) iris coloration in life: creamy white with irregular dark flecks; flecks concentrated around pupil giving impression of diffuse grey ring; (18) distribution of melanophores on digits constant in both specimens, absent from fingers, present on toes IV and V; in life hands and feet are light green, tips of fingers and toes pale orange; (19) males were seen calling from the undersides of leaves; advertisement call consisting of a single note, distinctly pulsed (5���6 pulses); call duration note 0.32��� 0.37 s, dominant frequency 4670���4800 Hz with no apparent frequency modulation; (20) clutches were located on the lower surfaces overhanging streams, clutch size 19��� 20 eggs (N = 3); (21) SVL in males 20.6���21.4 mm (N = 2); (22) enameled glands absent from lower part of upper lip and skin covering jaw. Comparisons. Many species of Hyalinobatrachium are difficult to diagnose using only morphological characters, especially when using preserved specimens (Castroviejo-Fisher et al. 2009; 2011). In our diagnosis we combine typical morphological characters (in most cases they show only subtle differences that might need a trained eye and direct comparisons of specimens to detect), with bioacoustic evidence and phylogenetic relationships inferred from DNA sequences (Table 3, Fig. 2 a). As the new species occurs in the east-Andean versant, we limit our comparisons to species which occur in this ecoregion, except for two closely related species occurring west of the Andes: H. colymbiphyllum and H. chirripoi, both of which occur in Central America and northeastern South America (H. colymbiphyllum ranges into the R��o Cauca and R��o Magdalena valleys of Colombia). Character states of H. anachoretus are in parentheses. Hyalinobatrachium bergeri has a hand webbing formula III 3 - 2 + IV (III 2 - 2 ��� IV); advertisement call consisting of a single monotone note, call duration 0.129��� 0.198 s (single trilled note with 5���6 pulses, 0.32��� 0.37 s); pericardium with at least some iridophores (pericardium without iridophores); they are not sister species in phylogenetic analysis. Hyalinobatrachium carlesvilai has a hand webbing formula III 2 ��� - 1 + IV (III 2 - 2 ��� IV); advertisement call consisting of a single note, the first third being pulsed and the second third tonal, call duration 0.102��� 0.152 s (single trilled note with 5���6 pulses, 0.32��� 0.37 s); pericardium with at least some iridophores (pericardium without iridophores); they are not sister species in phylogenetic analysis. Hyalinobatrachium chirripoi has ���extensive��� hand webbing (as described by Kubicki 2004, 2007) (hand webbing moderate, i.e., III 2 - 2 ��� IV); advertisement call with pulse rate 55���57 pulses/s (13.8���19.2 pulses/s); SVL 23.7���26.4 mm (20.6���21.4 mm). Hyalinobatrachium colymbiphyllum has a single trilled note advertisement call with 7���9 pulses and lower dominant frequency 3375���3658 Hz (single trilled note with 5���6 pulses and higher dominant frequency 4670���4800 Hz); SVL 23.0��� 30.6 mm (20.6���21.4 mm); yellow or gold iris in life (white or cream iris). Hyalinobatrachium esmeralda has a hand webbing formula III 2 + - 2 + IV (III 2 - 2 ��� IV); pericardium white or partially pigmented (transparent, but see Fig. 23), single tonal note advertisement call (single trilled note with 5���6 pulses) (M. A. Rada, pers. comm.); snout rounded in dorsal and lateral views (truncate). Hyalinobatrachium ibama has a white pericardium (transparent); dorsal surfaces with minute and small melanophores (only minute melanophores); they are not sister species in phylogenetic analysis. Hyalinobatrachium mondolfii has a hand webbing formula III 2 - (1 + ��� 1 1 / 2) IV (III 2 - 2 ��� IV); single tonal note advertisement call (single trilled note with 5���6 pulses); pericardium white (transparent); snout rounded in dorsal and lateral views (truncate); they are not sister species in phylogenetic analysis. Hyalinobatrachium munozorum has a hand webbing formula III 2 ��� - 1 + IV (III 2 - 2 ��� IV); pericardium white (transparent); snout rounded in dorsal and lateral views (truncate); they are not sister species in phylogenetic analysis. Hyalinobatrachium pellucidum has a single tonal note advertisement call, each note 0.12��� 0.15 s in duration, dominant frequency 4863���5408 Hz (vs. single trilled note with 5���6 distinct pulses, notes 0.32��� 0.37 s in duration, dominant frequency 4670���4800 Hz); they are not sister species in phylogenetic analysis. Hyalinobatrachium ruedai has a hand webbing formula III 2 - 1 3 / 4 IV (III 2 - 2 ��� IV); pericardium white (transparent); snout rounded in dorsal and lateral views (truncate). Description of the holotype. Adult male with SVL 21.4 mm. Head just wider than body; head width 38 % of SVL; head width 1.32 times head length. Snout truncate in dorsal and lateral view; eye-nostril distance/eye diameter = 1.12 and eye-nostril distance/interorbital distance = 0.66. Loreal region flat and nearly vertical, nostrils slightly protuberant, elliptical; internarial region concave anterodorsally; canthus rostralis well defined. Eyes small, directed anterolaterally, eyes 47.5 �� relative to midline (where anterior-facing eyes would be 90 �� relative to midline); eye diameter 2.2 times wider than width of disc on finger III; eye diameter 36 % of head length and 59 % of interorbital distance. Tympanum hardly visible, with no annulus nor supratympanic fold visible externally, center colored as the dorsum; tympanum with slight dorsal inclination. Dentigerous processes on vomers absent, choanae large, circular, separated more widely than nostrils; tongue round, white in preservative, anterior 2 / 3 attached to mouth; vocal slits present, extending along floor of mouth lateral to tongue; enameled glands absent from lower part of upper jaw. Forelimbs slim, with forearm roughly 1.5 times as wide as arm; ulnar fold present and enameled; humeral spine absent. Relative length of fingers: I Coloration in life. Overall impression is apple-green with irregular, diffuse yellow blotches. Dorsum peppered with small melanophores that appear dark green (Fig. 24). Legs and arms pale green, melanophores absent in circular patches giving the impression of faint pale-yellow spots. Finger and toe discs pale orange. Flanks white. Iris creamy white with irregular dark flecks. Flecks concentrated around pupil giving impression of diffuse grey ring surrounding pupil. Ventrally, parietal peritoneum transparent; pericardium also transparent with heart showing red in life; visceral peritoneum of gall bladder and urinary bladder also transparent. Hepatic and visceral peritonea white. Ventral vein red, prominent. Coloration in preservative. General appearance pale yellow (Fig. 25). Dorsal surfaces dotted with sparse melanophores; faint yellow spots present on the legs. Venter appearing uniform white. Iris and peritonea as in life. Measurements. Holotype measurements (in mm) with paratype measurements given in parentheses: SVL 21.4 (20.6), HL 6.2 (6.1), HW 8.2 (8.0), TD 0.8, IND 1.3 (1.5), IOD 3.4 (3.4), ED 2.0 (2.0), EW 3.0 (2.9), END 2.2 (2.3), HaL 6.0 (5.3), 3 DW 0.9 (0.9), TL 11.2 (10.5), SL 11.6 (11.6), FL 9.1 (8.9). Note that in the paratype, the tympanum was not visible, thus tympanum diameter is not reported. Variation. In life, the holotype has slightly more prominent spots on the legs than the paratype. One uncollected male, which was photographed near an egg clutch (Fig. 24) had more prominent spotting on the dorsum than either of the types. In the paratype, iridophores cover just the medial-most edge of the kidneys, whereas the inner third of the kidneys have iridophores in the holotype. Vocalizations. Numerous males were calling on the night the type series was collected. The analysis here is based on a recording of a single male from the type locality, taken at 15.5 ��C. Although the recorded call belongs to an uncollected male, we (ET) observed a male calling from the underside of a leaf next to eggs, so the call reported here is definitely attributable to H. anachoretus. The entire recording is 58 s long and contains 4 notes all given by the same male. The call consists of a single note that is distinctly pulsed, with notes given 13���16 s apart. Each note is 0.32��� 0.37 s in duration, containing 5���6 distinct pulses (pulse rate 13.8���19.2 pulses/s). Dominant frequency 4670���4800 Hz with no apparent frequency modulation. Unfortunately, the recording has too much background noise for illustration purposes; however, the recording is of high enough quality to extract relevant call parameters. This pulsed call is unlike any call known from Hyalinobatrachium in the east-Andean versant, and is most similar to H. colymbiphyllum, which occurs in southeastern Central America and the Pacific lowlands and R��o Cauca and Magdalena drainages of Colombia (AmphibiaWeb, 2014). However, there appear to be some differences in calls between these species. We (JD) have recorded 5 individuals of H. colymbiphyllum from R��o Frijoles ca. 300 m, Parque Nacional Soberan��a, Panama, and the calls have more pulses (7���9 pulses in H. colymbiphyllum vs. 5���6 in H. anachoretus), and lower dominant frequency (3375���3658 Hz in H. colymbiphyllum vs. 4670���4800 Hz in H. anachoretus). Hyalinobatrachium chirripoi and H. pellucidum (two other closely related species to H. anachoretus) have distinct calls. The call of H. chirripoi (based on a recording from a single male found along a tributary of R��o Banano at 80 m, Lim��n, Costa Rica; Kubicki 2004, call courtesy B. Kubicki) is much more highly pulsed (pulse rate 55���57 pulses/s in H. chirripoi vs. 13.8���19.2 pulses/s in H. anachoretus), whereas the call of H. pellucidum is a single, monotone, non-pulsed note (Wen et al. 2012). The advertisement calls of Hyalinobatrachium esmeralda and H. aff. pellucidum have not been described; however, we have analyzed recordings of each taxon (from the type locality and from Chivor, Boyac��, respectively) graciously provided by M. A. Rada. A full description of the calls will be published elsewhere (M. A. Rada in prep.), but both are clearly different from that of H. anachoretus in that they are composed of a single, unpulsed, tonal note. Distribution and ecology. Hyalinobatrachium anachoretus is known only from the vicinity of the type locality (Fig. 18). Males were also heard calling in a stream near Puente Nieva, which is roughly 2 km by road from the type locality and located at 2050 m. As far as we can tell, the latter site is also the type locality of Centrolene lemniscatum and Ce. muelleri (Duellman & Schulte 1993), although we have not encountered either species at that site. Thus, H. anachoretus appears to be sympatric with Ce. lemniscatum and Ce. muelleri at least in this area. Calling behavior and general activity in this species is perplexing. We have visited the type locality at night on three occasions (24 May 2010, 5 May 2011, and 14 June 2011) and were only able to hear and find frogs on the night of 5 May 2011. In addition, R. Schulte visited this site at least once while collecting glassfrogs (the types of Ce. lemniscatum and Ce. muelleri were collected 9 July 1989 from a stream less than 2 km from the type locality of H. anachoretus, a site where we also heard H. anachoretus calling on 5 May 2011), yet H. anachoretus was not recorded. It is worth mentioning that on the one night these frogs were active they were abundant and conspicuous, with high densities of males calling along every drainage in the area (even R��o Nieva, the largest drainage in the area). Weather conditions are known to affect calling rates in some Central American Hyalinobatrachium, with a decline in vocal activity during drier evenings (Vockenhuber et al. 2008; Kubicki 2007). During the three nights when we visited the area there was significant rainfall and high humidity, typical for this high cloud forest region. A possible explanation is that the breeding season of H. anachoretus ends sometime in early to mid-May, which coincides with the end of the wet season. Although we did not directly observe oviposition, three clutches were found near calling males (Fig. 24). These were typical of Hyalinobatrachium, consisting of a monolayer array of unpigmented eggs attached to the lower surfaces of broadleaf vegetation. Clutch sized ranged from 21��� 25 eggs (N = 3), which is notably smaller than that of H. chirripoi and H. colymbiphyllum. Hyalinobatrachium chirripoi clutch size ranges from 65��� 80 eggs (Kubicki 2004). In H. colymbiphyllum from near Gamboa, Panama clutch sized ranged from 51��� 102 eggs (JD, unpublished), and near Rinc��n de Osa, Costa Rica, clutch size ranged from 33��� 70 eggs (McDiarmid 1978). Kubicki (2007) reported smaller clutches (25��� 35 eggs) in a high-elevation population near Monteverde, Costa Rica. In the closely related H. pellucidum clutch size ranged from 24��� 41 eggs near San Jose (JD, unpublished.). Combat behavior, tadpoles, and parental care are unknown for this species. Etymology. The name is derived from the Latin noun anachoretus, which means ���hermit���. This name seems appropriate for two reasons. First, this species appears to be isolated from other members of the genus in that it occurs in the cool, rainy highlands of the Alto Mayo drainage. Second, the species seems to be reclusive, as we have only been able to locate it on a single night of collecting despite several visits to the locality. Remarks. Our phylogeny shows H. anachoretus as part of a clade containing H. aff. pellucidum, H. colymbiphyllum, and H. chirripoi. The topology we recovered did not support a monophyletic H. colymbiphyllum. One interpretation of this topology would be that H. chirripoi is a synonym of H. colymbiphyllum. However, H. chirripoi has a distinct advertisement call (see above) and is diagnosable from H. colymbiphyllum on the basis of hand webbing (extensive in H. chirripoi vs. moderate in H. colymbiphyllum; Kubicki 2004). Anot, Published as part of Twomey, Evan, Delia, Jesse & Castroviejo-Fisher, Santiago, 2014, A review of Northern Peruvian glassfrogs (Centrolenidae), with the description of four new remarkable species, pp. 1-87 in Zootaxa 3851 (1) on pages 39-43, DOI: 10.11646/zootaxa.3851.1.1, http://zenodo.org/record/286921, {"references":["Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Kubicki, B. (2004) Rediscovery of Hyalinobatrachium chirripoi (Anura: Centrolenidae) in southeastern Costa Rica. Revista de Biologia Tropical, 52, 214 - 218.","Kubicki, B. (2007) Glass frogs of Costa Rica. Instituto Nacional de Biodiversidad (INBio), Heredoa, Costa Rica. [total page number unkown]","AmphibiaWeb: Information on amphibian biology and conservation (2014) Web application. 2014. Berkeley, California: AmphibiaWeb. Available from: http: // amphibiaweb. org / (accessed 27 May 2014)","Wen, A., Vasquez, N. & Castroviejo-Fisher, S. (2012) Description of the previously unknown advertisement calls of Hyalinobatrachium fragile, H. pellucidum, and Vitreorana antisthenesi (Amphibia: Centrolenidae). Zootaxa, 3480, 80 - 87.","Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. The University of Kansas Museum of Natural History Occasional Papers, 155, 1 - 33.","McDiarmid, R. W. (1978) Evolution of parental care in frogs. In: The Development of Behavior. Garland STPM Press, New York, New York, USA, pp. 127 - 148.","Crawford, A. J., Lips, K. R. & Bermingham, E. (2010) Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama. Proceedings of the National Academy of Sciences, 107, 13777 - 13782. http: // dx. doi. org / 10.1073 / pnas. 0914115107"]}

Ranitomeya vanzolinii species group Figs. 3, 4, 9, 23 – 29 Tables 1, 4 – 6 A monophyletic assemblage of six species: Ranitomeya vanzolinii Myers 1982; R. sirensis Aichinger 1991 with its junior synonyms R. biolat Morales 1992 and R. lamasi Morales 1992; R. imitator Schulte 1986 with its junior synonyms R. imitator intermedius Schulte 1999 and R. imitator yurimaguensis Schulte 1999; R. flavovittata Schulte 1999; R. yavaricola Perez-Peña, Chavez, Twomey & Brown 2010 and R. cyanovittata Perez-Peña, Chavez, Twomey & Brown 2010. Definition and diagnosis. Medium to large adult SVL (17 – 21 mm); black dorsum; highly variable morphology; LTRF 2(2)/3(1), oral disc emarginated; large intestine unpigmented; larvae whitish to gray; eggs cream; territoriality present in males; promiscuous (known in R. sirensis) or monogamous mating system (known in R. vanzolinii and R. imitator), male parental care (observed in R. sirensis) or biparental (observed in R. vanzolinii and R. imitator); females produce between 2 – 4 eggs per mating. The advertisement call of species in this group consists of a loud trill, highly tonal, with notes 0.4 – 1.1 sec in length, repeated at 2 – 14 notes per minute (Fig. 28)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 61, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Myers, C., W. (1982) Spotted poison frogs: Description of Three new Dendrobates from Western Amazonia, and resurrection of a lost species from \" Chiriqui \". American Museum Novitates, 2721, 23.","Aichinger, M. (1991) A new species of poison-dart frog (Anura: Dendrobatidae) from the Serrania de Sira, Peru. Herpetologica, 47, 1 - 5.","Morales, V. (1992) Dos especies nuevas de Dendrobates (Anura: Dendrobatidae) para Peru. Caribbean Journal of Science, 28, 191 - 199.","Schulte, R. (1986) Eine neue Dendrobates - Art aus Ostperu (Amphibia: Salentia: Dendrobatidae). Sauria, 8, 11 - 20.","Schulte, R. (1999) Pfeilgiftfrosche \" Artenteil - Peru \". INBICO, Wailblingen, Germany, 294 pp.","Perez-Pena, P., Chavez, G., Twomey, E. & Brown, J. L. (2010) Two new species of Ranitomeya (Anura: Dendrobatidae) from eastern Amazonian Peru. Zootaxa, 2439, 1 - 23."]}

Ranitomeya variabilis species group Figs. 3, 4, 9, 14, 32 – 37 Tables 1, 4 – 6 A monophyletic assemblage of two species: Ranitomeya variabilis Zimmermann & Zimmermann 1988 and R. amazonica Schulte 1999. Definition and diagnosis. Medium to large adult size (SVL 14 – 21 mm); two prevalent dorsal patterns: large ovoid black spots on greenish dorsum or complete yellow to red dorsolateral stripes, middorsal stripe and oblique lateral stripes (note that occasionally these species possess ventrolateral stripes, or an intermediate between oblique lateral and ventrolateral stripes, suggesting this trait is a continuous character); large intestine of larvae entirely pigmented; LTRF 2 (2)/3(1), oral disc emarginated; larvae gray, tadpoles cannibalistic on embryos and other tadpoles; eggs gray; neither males nor females territorial; promiscuous mating system with male parental care; females produce between 2 – 6 eggs per mating. Advertisement calls consist of short, regularly spaced buzz calls, notes 0.16 – 0.44 sec in length, repeated at 24 – 70 notes per minute (Fig. 14)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 78, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Zimmermann, H., & Zimmermann, E. (1988) Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfroschen (Anura: Dendrobatidae). Salamandra, 24, 125 - 160.","Schulte, R. (1999) Pfeilgiftfrosche \" Artenteil - Peru \". INBICO, Wailblingen, Germany, 294 pp."]}

Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel, Summers, Kyle (2011): A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083. Zootaxa 3083 (1): 1-120, DOI: 10.11646/zootaxa.3083.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3083.1.1

Ranitomeya reticulata species group Figs. 3, 4, 9, 15 – 18 Tables 1, 4 – 6 A monophyletic assemblage of six species: Ranitomeya reticulata Boulenger 1884 “1883” with its junior synonym R. ignea Melin 1941; R. fantastica Boulenger, 1884 “1883”; R. ventrimaculata Shreve 1935; R. uakarii Brown, Schulte & Summers 2006; R. summersi Brown, Twomey, Pepper & Sanchez-Rodriguez 2008 and R. benedicta Brown, Twomey, Pepper, & Sanchez-Rodriguez 2008. Definition and diagnosis. Medium to large adult SVL (17 – 21 mm); black dorsum; typically orange to red dorsal patterning; LTRF 2 (2)/3, oral disc emarginate; large intestine entirely pigmented; eggs dark; promiscuous mating system, male parental care; grayish tadpoles and embryos, females produce between 2 – 5 eggs per mating. Vocalizations consist of a series of very short buzz-like notes (0.1 – 0.5 sec in length) given in rapid succession (104 – 200 notes per minute) (Fig. 18)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 48, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Boulenger, G. A. (1884 \" 1883 \") On a collection of frogs from Yurimaguas, Huallaga River, Northern Peru. Proceedings of the Zoological Society of London, 1883, 635 - 638, 1 colour pl, 1 B & W pl.","Melin, D. E. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungl. Vetenskaps-och Vitterhetssamhalles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1 - 71.","Shreve, B. (1935) On a new Teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional Papers of the Boston Society of Natural History, 8, 209 - 218.","Brown, J. L., Schulte, R. & Summers, K. (2006) A new species of Dendrobates (Anura: Dendrobatidae) from the Amazonian lowlands in Peru. Zootaxa, 45 - 58."]}

Andinobates minutus species group Figs. 3 – 5 (a – m), 8 Tables 4 –6 A monophyletic assemblage of two divergent species: Andinobates minutus (Shreve 1935) comb. nov. and A. claudiae (Jungfer, Lötters & Jörgens 2000) comb. nov. Definition and diagnosis. SVL 13–16 mm; dark brown or black dorsum; largely complete light colored dorsolateral stripes not extending to thigh and oblique lateral stripes present, typically incomplete and not extending to eye; large spots near body on upper surface of legs and forearms; limbs and venter black with pale marbling on venter; LTRF either 2(2)/3(1) (A. claudiae) or 2(2)/3 (A. minutus); larvae with complete papillae on posterior labium and wide gap in papillae on anterior labium; oral disc emarginated (Table 4); larvae gray; eggs dark (Table 6). Vocalizations short, tonal buzz-notes, notes less than 1 sec in length, repeated at 3 – 17 notes per minute (Fig. 8, Table 5).

Ranitomeya yavaricola sp. nov. Figures 2 –5, 11 Holotype. MZUNAP-01- 520 (Fig 3.), an adult male collected by Pedro Perez-Peña nearby Lago Preto, 17 km W of Estiron de Ecuador, Provincia Ramon Castilla, Departamento Loreto, Peru; 4 ° 27 ' 35.0" S, 71 ° 45 ’ 3.5 "W, 120 m elevation; August 2009; found foraging in leaf litter within terra firme forest. Paratypes. All from same locality as holotype (MZUNAP-01- 518, 519) an adult female and male (respectively) collected by P. Pérez-Peña in August 2009. Etymology. The specific epithet is noun in apposition that means “inhabitant of the Yavarí” and is formed from the Latin suffix “ -icola ” for “dweller” or “inhabitant” and Río Yavarí, the watershed where this species occurs. Definition and diagnosis. Assigned to the genus Ranitomeya due to the combination of the following characteristics: small size ( Ranitomeya yavaricola can be distinguished from other species of Ranitomeya by the combination of irregular, pale turquoise spots and stripes on the dorsum, solid-bronze limbs, and irregular sky-blue spots on the ventral surface of the upper thighs. Ranitomeya flavovittata possesses irregular, bright yellow spots and stripes (often broken) on the dorsum (vs. pale turquoise in R. yavaricola). Additionally, the limbs of R. flavovittata are black with light blue reticulation (vs. limbs solid bronze in R. yavaricola), and R. flavovittata typically possesses a complete (to nearly complete) yellow median dorsal stripe and lacks conspicuous spots on the ventral surface of the upper thighs (vs. median dorsal stripe absent and ventral thigh spots present in R. yavaricola). Ranitomeya vanzolinii possesses bright yellow dorsal spotting, light blue reticulation on a ground color of black on limbs, and lacks conspicuous spots on the ventral surface of the upper thighs. Ranitomeya yavaricola is similar in appearance to the nominal morph of Excidobates captivus and Adelphobates castaneoticus. The nominal morph of Excidobates captivus has dull brown limbs (vs. bronze in R. yavaricola), bears yellow spots on the flanks (vs. flank spots absent in R. yavaricola), and has paired reddishorange dorsolateral dashes (vs. dorsolateral spots and stripes pale turquoise to spring green in R. yavaricola). Adelphobates castaneoticus is larger in size (SVL up 23 mm), lacks the inner metacarpal tubercle, and lacks distinct bronze limbs. Furthermore, it has conspicuous red flash-marks on the upper surface of the forearms, thighs, and calves. Measurements (in mm) of holotype. The male holotype (Fig. 3) has SVL 15.2; FL 7.1; TL 7.2; KK 13.1; FoL 4.9; HaL 3.4; HL 5.5; HW 5.0; BW 4.3; UEW 2.3; IOD 2.2; IND 1.8; TD 0.9; ED 1.9; DET 0.5; L 1 F 1.9; L 2 F 2.2; W 3 D 0.9; W 3 F 0.3. For paratype measurements see Table 1. Description of holotype. Widest part of head at jaw articulations. Head width slightly wider than body. Tongue ovoid; teeth absent. Snout sloping and rounded in lateral profile, slightly blunted in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but not from dorsal view. Canthus rostralis rounded, loreal region flat and nearly vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to horizontal eye diameter. Tympanum round, partially concealed posterodorsally. In life, skin texture nearly smooth on dorsal surfaces of body and head; limbs and rump weakly granular. Venter weakly granular on limbs and body, ventral surface of head nearly smooth. Hands (Fig. 4) relatively large, length 22 % of SVL. Relative length of appressed fingers III> IV> II> I; first finger 90 % length of second; finger discs greatly expanded, width of disc on finger III 2.6 times width of adjacent phalanx. Unpigmented median metacarpal tubercle present on base of palm; inner metacarpal tubercle present near base of finger I; unpigmented proximal subarticular tubercles present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on finger III. All tubercles raised above level of hands; scutes present on dorsal surface of fingers. Hind limbs moderate length, with heel of appressed hind limbs reaching level of eye. Femur and tibia roughly equal in length, femur 99 % length of tibia; knee–knee distance 86 % of SVL. Relative lengths of appressed toes IV> III> V> II> I (Fig. 4); first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes III–V moderately expanded. Two unpigmented metatarsal tubercles on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I, II, III due to lack of pigmentation. Toes III and V with two large subarticular tubercles, toe IV with three subarticular tubercles. Tarsal keel extends from below knee to medial metatarsal tubercle at foot. Tarsal tubercle absent; feet and hands lack webbing and lateral fringing. Color in life. In life, body black with metallic sage-green dashes anterior and posterior to each eye, a single spot between eyes; metallic-sage dorsolateral stripes extend from upper thighs to shoulders where they each form a single dash. Broad sage oblique-lateral lines extend from the axillae to groin, where they fuse with a large spot on upper surfaces of each thigh. Broad labial stripe present, continues posteriorly to upper surfaces of arms. Limbs and digits solid bronze. Paired sage dashes on underside of thighs. Underside of head sage with two pairs of gular spots, creating appearance of an hourglass. Venter black with coarse, irregular sage marbling. Iris black. Color in preservative. In preservative, turquoise/sky-blue coloration turns grey and bronze limb coloration turns brown. Variation (based on 15 adults). Adults 15.2–17.7 mm SVL (mean 16.4 mm). Head about as wide as body except in single gravid female whose body was wider than the head. Head width 98 % of body width (range 81–109 %). Head width 30–36 % of SVL in adults. No apparent sexual dimorphism in external morphology except that males possess faint vocal slits on the floor of the mouth and have a slightly expanded subgular pouch. The completeness of dorsolateral and medial lateral stripes varies considerably between individuals (Fig. 5). Line/dash/spot coloration varies subtly between a sky-blue to a very light bluish-yellow to pale sage-green, with a majority of individuals being light turquoise. Venter coloration is typically darker than dorsal coloration and is less variable, being predominantly skyblue. Ventral ground coloration varies from dark brown to black. Venter reticulation is largely symmetrical and often forms a broad, irregular, black medial-stripe in half of the specimens (Fig. 5). Most individuals possess a large gular spot in the center of the throat (80 %). In all individuals, sky-blue dashes are present on the ventral surface of the upper thighs, though their size varies considerably between individuals. Hands relatively large, length 22–26 % of SVL. First finger 65–95 % length of second; finger discs moderately expanded in both males and females, width of disc on finger III 2 –3.6 times width of adjacent phalanx. Tibia 86–107 % length of femur (mean 99 %); knee–knee distance 82–92 % of SVL (mean 87 %). Tadpole measurements (in mm). A stage 25 tadpole (MZUNAP-01- 521) was used for the description (Fig. 6). Total length 12.5; body length 4.7; internarial distance 0.9; eye to nares distance 0.8; eye diameter 0.5; interorbital distance 0.7; tail length 7.8. Tadpole description. Snout rounded when viewed from above; body ovoid in dorsal view. Eyes dorsal, angled laterally, pupils white in preservative. Nares not forming tube, situated half-way between eye and tip of snout, directed dorsolaterally. Spiracle sinistral; vent dextral. Ventral tail fin begins at tail base, dorsal tail fin begins just posterior to plane of vent opening, ventral and dorsal fins relatively uniform in thickness throughout tail, tapering towards tip. Musculature depth uniform throughout, tapering toward tip. Mouth directed anteroventrally. Oral disc emarginate, anterior and posterior labia forming flaps free from body wall, 1.4 mm in width. Marginal papillae absent on anterior labium, present in one complete row on posterior labium. Papillae rounded; submarginal papillae absent. Jaw sheaths deep in longitudinal width, serrate, lacking indentations. Lateral processes short, extending barely past lower jaw. Labial tooth row formula is 2 (2)/ 3 [1]. A- 1 complete (62 teeth), A- 2 with medial gap (34 teeth), same width as A- 1. P- 1 with medial gap (46 teeth), P- 2 (44 teeth), and P- 3 complete (44 teeth); P- 1 and P- 2 equal width, P- 3 slightly shorter. Color in Life. In life, the head appears light grey. Pigmentation on dorsum mottled brown, ground color weakly transparent. Eyes black, papillae white. Ventral coloration is transparent (most internal organs are visible) with irregular faint red flecking that is dense around mouth and nares. Tail musculature white with abundant brown mottling, fins almost transparent. Color in Preservative. In preservative, coloration is identical, though red pigmentation turns to brown. Vocalizations. The following values are presented as: min-max (average ± SD, number of individuals). The advertisement call is a short trilled note (Fig. 7) with duration between 630–880 ms (760 ± 140 ms, 7) and is repeated at irregular intervals of 2–7 notes per minute (3.14 ± 1.80 notes per minute, 6). Each note consists of 20–27 pulses (mean = 24). Calling activity is sporadic and continues throughout daylight, but peaks in the early morning and late afternoon. Dominant frequency is 5400–6000 Hz (5600 ± 2000 Hz, 5) at temperatures between 24.5–26 °C. The call of R. yavaricola sounds similar to the calls of other species in the vanzolinii group, although there are some slight differences within this group. Ranitomeya flavovittata has slightly longer notes (1 sec vs. 760 ms in R. yavaricola). Ranitomeya imitator (n = 25) has a call that is slightly shorter in duration (686 ms vs. 760 ms in R. yavaricola) and has a slightly lower dominant frequency (5200 Hz vs 5600 Hz in R. yavaricola). Distribution and natural history. Ranitomeya yavaricola occurs in primary forests in a small area of northeastern Peru. This species likely occurs more widely within the broad interfluvium bordered by the Ucayali, Amazon, Yavarí, and Blanco rivers (Fig. 1). Much of the area bordering these rivers is seasonally or permanently flooded and it is possible that these large expanses impose limits to this species’ distribution (Fig. 8). Ranitomeya yavaricola is only known from forests near Lago Preto (but see discussion). Lago Preto is a large oxbow lake at the confluence of Yavarí and Yavari-Mirin rivers. Near Lago Preto, several forest types are present: seasonally flooded forests, swamp forests that are saturated year-round, and upland terra firme forests. At Lago Preto, R. yavaricola occurs in upland forests and very low lying forests, just above the flood zone of seasonally flooded forests (i.e. várzea forests) and swamps. These sites contained many large trees and a relatively sparse understory. The upland forests in this region are some of the most diverse in the Amazon basin, with estimates of tree diversity exceeding 300 species per hectare in some areas (Pitman et al. 2003, Fine et al. 2006) These forests are dominated by trees of the families Fabaceae (legumes), Bombacaceae (“mallows” such as kapoks), and Moraceae (figs) (Fine et al. 2006). This species is sympatric with three other dendrobatids: Ranitomeya uakarii, Ameerega hahneli, and A. trivittata. This species potentially also co-occurs with R. flavovittata and R. ventrimaculata, both of which have been observed less than 90 km from Lago Preto (Fig. 8). Ranitomeya yavaricola was typically observed foraging throughout the leaf litter or calling from the leaves of terrestrial palms (Genoma spp.), epiphytes (primarily bromeliads), and on the branches of fallen trees (between 0.2–3 m above the ground). Ranitomeya yavaricola is an extremely shy species and when encountered dives to the ground and hides within the leaf litter or within the roots of plants. A single tadpole was observed in the phytotelm of a small bromeliad that was ca. 1.5 m above the ground. Prior to the discovery of the tadpole, a male was observed calling near the bromeliad (Ranitomeya yavaricola is the most abundant species of Ranitomeya at Lago Preto. Using visual encounters, we observed 3 individuals in 30 man-hours of observation (0.1 encounters per man-hour) and using acoustic estimates, we recorded 20 different individuals in 3.7 man-hours of observation (5.6 encounters per man-hour). Lastly, as the result of casual observations, over a two week period in June 2009, 20 people encountered 6 individuals. During this study 12 individuals of R. uakarii were observed at Lago Preto (as the result of both casual observation and surveys). Conservation status. Following the IUCN Red List criteria (IUCN 2001), this species should be listed as Data Deficient (DD). It is currently known from only a single locality but probably occurs more widely.

Ranitomeya cyanovittata sp. nov. Figures 8, 10 Holotype. CORBIDI 0 2266 (Fig. 10 A), an adult female collected by Diego Vasquez in the Rio Blanco Basin near the Zona Reservada Sierra del Divisor, Departamento Loreto, Peru; 6 �� 55 ��� 12 ���S, 73 �� 50 ��� 45 W, 206 m elevation; September 2008. Paratype. CORBIDI 0 2991 (Fig. 10 B), an adult male collected by Diego Vasquez in the Rio Blanco Basin near the Zona Reservada Sierra del Divisor, Departamento Loreto, Peru; 6 �� 54 ��� 57 ���S, 73 �� 50 ��� 45 ���W, 257 m elevation; December 2008. Etymology. The specific epithet, an adjective, means ���blue striped��� and is formed from the combination of the ancient Greek noun �����υ����������� (or ��� cyanos ���) for ���a blue substance used to adorn armor in the Heroic age��� and the Latin adjective ��� vittatus ��� for ���banded.��� Definition and diagnosis. Assigned to the genus Ranitomeya due to the combination of the following characteristics: small size (Ranitomeya cyanovittata is a relatively small species of Ranitomeya, adult SVL of approximately 13.8���17.3 mm. Bicolored pattern bright blue and black; teeth absent; finger I shorter than finger II; disc of third finger 1.9���2.2 times wider than finger width. Ranitomeya cyanovittata can be distinguished from all other dendrobatids by its distinctive bicolored pattern of a black background with turquoise-blue lines or reticulations over the entire body (dorsal and ventral side). Six other species of Ranitomeya have a bicolored pattern. Ranitomeya fulgurita (Silverstone, 1975) ranges from Panama to Northwestern Colombia and is bicolored dorsally and ventrally, but this pattern consists of a greenish-yellow body with black stripes or spots (L��tters et al., 2007). Ranitomeya abdita from southern Ecuador (Myers & Daly, 1976) has a solid dark bronze to black body with orange spots on the posterior areas of the upper arm and thigh (L��tters et al., 2007). Ranitomeya altobueyensis from the Serrania de Baud�� in Colombia (Silverstone, 1975) has a uniform yellow or golden metallic body (sometimes with a greenish hue), occasionally with small black spots on the dorsum and often on the venter (L��tters et al., 2007). Ranitomeya dorisswansonae from the Cordillera Central in Tolima, Colombia (Rueda-Almonacid et al., 2006) has a dark brown or reddish brown background with irregular red to yellowish-orange spots that may be absent on the limbs (L��tters et al., 2007). Ranitomeya sirensis from the Cordillera El Sira in Central Amazonian Peru (Aichinger, 1991) has a red dorsum with turquoise-green legs and arms, a turquoise-green venter with a red patch on the belly, and lacks dorsal reticulations or spots (L��tters et al., 2007). Finally, Ranitomeya summersi from Central Amazonian Peru has a black ground color with orange stripes on the dorsum and venter (Brown et al., 2008). Ranitomeya cyanovittata is most similar in appearance to the following species: Ranitomeya biolat, R. flavovittata, R. imitator, R. lamasi, and R. ventrimaculata. Ranitomeya biolat and most lowland populations of R. imitator possess yellow dorsal stripes (vs. blue stripes in R. cyanovittata); R. flavovittata has broken, irregular, yellow dorsal stripes (vs. unbroken blue stripes in R. cyanovittata); R. lamasi has dorsal stripes which can be green, yellow, orange, or red (but never blue as in R. cyanovittata) and typically has a yellow patch on the belly (vs. patch absent in R. cyanovittata). Ranitomeya ventrimaculata has yellow dorsal stripes that are not parallel (vs. blue stripes parallel in R. cyanovittata). Measurements (in mm) of holotype. The female holotype (Fig. 10 A) has SVL 17.3; FL 6.5; TL 6.8; KK 13.3; FoL 6.4; HaL 4.2; HL 5.1; HW 5.6; BW 6.4; UEW 1.2; IOD 2.6; IND 2.0; TD 0.46; ED 1.7; DET 0.48; L 1 F 1.77; L 2 F 2.28; W 3 D 1.09; W 3 F 0.53. Paratype measurements (Fig. 10 B): SVL 13.8; FL 5.5; TL 6.1; KK 11.7; FoL 5.09; HaL 3.6; HL 3.6; HW 3.9; BW 3.9; UEW 0.94; IOD 2.0; IND 1.7; TD 0.35; ED 1.3; DET 0.35; L 1 F 1.28; L 2 F 1.85; W 3 D 0.56; W 3 F 0.26. Description of holotype. Widest part of head at jaw articulations. Head slightly narrower than body. Tongue ovoid; teeth absent. In life, head black with blue spot in pit of snout, transverse blue stripe between eyes extending through upper eyelid anterior to upper insertion point of thigh; labial stripe extends to upper insertion of arm. Dorsum black with a vertebral line from posterior region of head to vent. Weakly-defined spot present on dorsal surface of axilla. Arms, legs, and toes are covered in a pale reticulations on a black background color, creating round black spots. Underside of head with lateral gular spots, creating the appearance of an hourglass. Venter black with blue irregular reticulation, creating black spots, much like the limbs. In life, skin texture nearly smooth on the dorsal surfaces of the body and head; limbs and rump weakly granular. Venter weakly granular on limbs and body, ventral surface of head nearly smooth. Snout sloping and rounded in lateral profile, round or slightly blunted in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but not from dorsal view. Canthus rostralis rounded, loreal region flat and nearly vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to eye width. Tympanum round, partially concealed posterodorsally. Hands relatively large, length 24.5 % of SVL. Relative length of appressed fingers III> IV> II> I; first finger 77.6 % length of second; finger discs moderately expanded, width of disc on finger III 2.05 times width of adjacent phalanx. An unpigmented median metacarpal tubercle is present on base of palm; inner metacarpal tubercle present near base of finger I; unpigmented proximal subarticular tubercle present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on fingers III and IV. All tubercles raised above level of hands; scutes present on dorsal surface of fingers. Hind limbs moderate length, with heel of appressed hind limbs reaching level of eye. Femur and tibia roughly equal in length, tibia 99.1 % length of femur; knee���knee distance 77 % of SVL. Relative lengths of appressed toes IV> III> V> II> I; first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes III���V moderately expanded. Two unpigmented metatarsal tubercles present on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I and II due to their lack of pigmentation. Toes III and V with two subarticular tubercles, toe IV with three subarticular tubercles. A tarsal keel is present starting below the knee and turning into the medial metatarsal tubercle at the foot. Tarsal tubercle absent; feet and hands lacking webbing and lateral fringing. Color in life: Dorsal body, legs, and head are black with blue longitudinal irregular stripes; a longitudinal, bright blue stripe is present from the tip of the labial region to the insertion point of the arms, a blue lateral stripe is present from the armpit to the groin. There is a weakly-defined pale blue spot on the dorsal surface of the axilla. The arms, legs, and toes are covered in pale blue reticulations on a black background color, creating round black spots; toes are blue-black. Underside of head blue with lateral black gular spots, creating appearance of a blue hourglass. Lower lip blue. Venter black with blue irregular spots, creating black spots, much like the limbs. Limbs are brownish. Iris black. Color in preservative. In preservative, coloration is identical to color in life, with the exception of blue coloration being grey. Distribution and natural history. Ranitomeya cyanovittata is known only from two sites near Nueva Capanahua community, in the Rio Blanco basin near the Sierra del Divisor, between 200 and 300 m elevation (Figs. 1 and 11). This species inhabits undisturbed upland forests near small streams and rivulets (Fig. 12). The vegetation consists principally of the following trees: Moronobea spp., Cedrella spp., and Cedrelinga cateniformis. Lianas are common but were not noted to occur in dense tangles. At ground level, vegetation was sparse, with some ferns and other large, herbaceous plants (such as Heliconia spp.). Bromeliads were not recorded. The leaf litter was sparse and appeared dry after several rainless days. The two individuals observed were foraging in leaf litter (Pers. Comm. Diego Vasquez): this is notable because other members of Ranitomeya vanzolini group are almost always found within or upon some sort of plant (e.g. Heliconia, Xanthosoma, or Dieffenbachia). Both specimens were encountered during the day and were sympatric with Ameerega ignipedis and A. cf. hahneli: of these species, Ameerega ignipedis was abundant and was frequently observed during the day in the leaf litter (Pers. Comm. Diego Vasquez). Neither courtship nor other behavioral characteristics were observed in R. cyanovittata. Conservation status: Following the IUCN Red List criteria (IUCN 2001), this species should be listed as Data Deficient (DD). Being known from only from two localities that are less than 1 km apart, its extent of occurrence is unknown., Published as part of Perez-Pe��a, Pedro E., Chavez, German, Twomey, Evan & Brown, Jason L., 2010, Two new species of Ranitomeya (Anura: Dendrobatidae) from eastern Amazonian Peru, pp. 1-23 in Zootaxa 2439 on pages 12-16, DOI: 10.5281/zenodo.194909, {"references":["Silverstone, P. A. (1975) A revision of the poison-arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County, 21, 1 - 55.","Lotters, S., Jungfer, K. - H., Schmidt, W. & Henkel, F. W. (2007) Poison Frogs: Biology, Species and Captive Husbandry. Edition Chimaira, Frankfurt am Main, 667 pg.","Myers, C. W. & Daly, J. W. (1976) A new species of poison frog (Dendrobates) from Andean Ecuador, including an analysis of its skin toxins. Occasional papers of the Museum of Natural History, the University of Kansas, 59, 1 - 12.","Rueda-Almonacid, J. V., Rada, M., Sanchez-Pacheco, S. J., Velasquez-Alvarez, A. A. & Quevedo, A. (2006) Two new and exceptional poison dart frogs of the genus Dendrobates (Anura: Dendrobatidae) from the Northeastern Flank of the cordillera Central of Colombia. Zootaxa, 1259, 39 - 54.","Aichinger, M. (1991) A new species of poison-dart frog (Anura: Dendrobatidae) from the Serrania de Sira, Peru. Herpetologica, 47, 1 - 5.","IUCN (2001) IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission, IUCN, Gland, Switzerland and Cambridge, UK."]}

Ranitomeya yavaricola sp. nov. Figures 2 ���5, 11 Holotype. MZUNAP-01- 520 (Fig 3.), an adult male collected by Pedro Perez-Pe��a nearby Lago Preto, 17 km W of Estiron de Ecuador, Provincia Ramon Castilla, Departamento Loreto, Peru; 4 �� 27 ' 35.0" S, 71 �� 45 ��� 3.5 "W, 120 m elevation; August 2009; found foraging in leaf litter within terra firme forest. Paratypes. All from same locality as holotype (MZUNAP-01- 518, 519) an adult female and male (respectively) collected by P. P��rez-Pe��a in August 2009. Etymology. The specific epithet is noun in apposition that means ���inhabitant of the Yavar����� and is formed from the Latin suffix ��� -icola ��� for ���dweller��� or ���inhabitant��� and R��o Yavar��, the watershed where this species occurs. Definition and diagnosis. Assigned to the genus Ranitomeya due to the combination of the following characteristics: small size (Ranitomeya yavaricola can be distinguished from other species of Ranitomeya by the combination of irregular, pale turquoise spots and stripes on the dorsum, solid-bronze limbs, and irregular sky-blue spots on the ventral surface of the upper thighs. Ranitomeya flavovittata possesses irregular, bright yellow spots and stripes (often broken) on the dorsum (vs. pale turquoise in R. yavaricola). Additionally, the limbs of R. flavovittata are black with light blue reticulation (vs. limbs solid bronze in R. yavaricola), and R. flavovittata typically possesses a complete (to nearly complete) yellow median dorsal stripe and lacks conspicuous spots on the ventral surface of the upper thighs (vs. median dorsal stripe absent and ventral thigh spots present in R. yavaricola). Ranitomeya vanzolinii possesses bright yellow dorsal spotting, light blue reticulation on a ground color of black on limbs, and lacks conspicuous spots on the ventral surface of the upper thighs. Ranitomeya yavaricola is similar in appearance to the nominal morph of Excidobates captivus and Adelphobates castaneoticus. The nominal morph of Excidobates captivus has dull brown limbs (vs. bronze in R. yavaricola), bears yellow spots on the flanks (vs. flank spots absent in R. yavaricola), and has paired reddishorange dorsolateral dashes (vs. dorsolateral spots and stripes pale turquoise to spring green in R. yavaricola). Adelphobates castaneoticus is larger in size (SVL up 23 mm), lacks the inner metacarpal tubercle, and lacks distinct bronze limbs. Furthermore, it has conspicuous red flash-marks on the upper surface of the forearms, thighs, and calves. Measurements (in mm) of holotype. The male holotype (Fig. 3) has SVL 15.2; FL 7.1; TL 7.2; KK 13.1; FoL 4.9; HaL 3.4; HL 5.5; HW 5.0; BW 4.3; UEW 2.3; IOD 2.2; IND 1.8; TD 0.9; ED 1.9; DET 0.5; L 1 F 1.9; L 2 F 2.2; W 3 D 0.9; W 3 F 0.3. For paratype measurements see Table 1. Description of holotype. Widest part of head at jaw articulations. Head width slightly wider than body. Tongue ovoid; teeth absent. Snout sloping and rounded in lateral profile, slightly blunted in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but not from dorsal view. Canthus rostralis rounded, loreal region flat and nearly vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to horizontal eye diameter. Tympanum round, partially concealed posterodorsally. In life, skin texture nearly smooth on dorsal surfaces of body and head; limbs and rump weakly granular. Venter weakly granular on limbs and body, ventral surface of head nearly smooth. Hands (Fig. 4) relatively large, length 22 % of SVL. Relative length of appressed fingers III> IV> II> I; first finger 90 % length of second; finger discs greatly expanded, width of disc on finger III 2.6 times width of adjacent phalanx. Unpigmented median metacarpal tubercle present on base of palm; inner metacarpal tubercle present near base of finger I; unpigmented proximal subarticular tubercles present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on finger III. All tubercles raised above level of hands; scutes present on dorsal surface of fingers. Hind limbs moderate length, with heel of appressed hind limbs reaching level of eye. Femur and tibia roughly equal in length, femur 99 % length of tibia; knee���knee distance 86 % of SVL. Relative lengths of appressed toes IV> III> V> II> I (Fig. 4); first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes III���V moderately expanded. Two unpigmented metatarsal tubercles on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I, II, III due to lack of pigmentation. Toes III and V with two large subarticular tubercles, toe IV with three subarticular tubercles. Tarsal keel extends from below knee to medial metatarsal tubercle at foot. Tarsal tubercle absent; feet and hands lack webbing and lateral fringing. Color in life. In life, body black with metallic sage-green dashes anterior and posterior to each eye, a single spot between eyes; metallic-sage dorsolateral stripes extend from upper thighs to shoulders where they each form a single dash. Broad sage oblique-lateral lines extend from the axillae to groin, where they fuse with a large spot on upper surfaces of each thigh. Broad labial stripe present, continues posteriorly to upper surfaces of arms. Limbs and digits solid bronze. Paired sage dashes on underside of thighs. Underside of head sage with two pairs of gular spots, creating appearance of an hourglass. Venter black with coarse, irregular sage marbling. Iris black. Color in preservative. In preservative, turquoise/sky-blue coloration turns grey and bronze limb coloration turns brown. Variation (based on 15 adults). Adults 15.2���17.7 mm SVL (mean 16.4 mm). Head about as wide as body except in single gravid female whose body was wider than the head. Head width 98 % of body width (range 81���109 %). Head width 30���36 % of SVL in adults. No apparent sexual dimorphism in external morphology except that males possess faint vocal slits on the floor of the mouth and have a slightly expanded subgular pouch. The completeness of dorsolateral and medial lateral stripes varies considerably between individuals (Fig. 5). Line/dash/spot coloration varies subtly between a sky-blue to a very light bluish-yellow to pale sage-green, with a majority of individuals being light turquoise. Venter coloration is typically darker than dorsal coloration and is less variable, being predominantly skyblue. Ventral ground coloration varies from dark brown to black. Venter reticulation is largely symmetrical and often forms a broad, irregular, black medial-stripe in half of the specimens (Fig. 5). Most individuals possess a large gular spot in the center of the throat (80 %). In all individuals, sky-blue dashes are present on the ventral surface of the upper thighs, though their size varies considerably between individuals. Hands relatively large, length 22���26 % of SVL. First finger 65���95 % length of second; finger discs moderately expanded in both males and females, width of disc on finger III 2 ���3.6 times width of adjacent phalanx. Tibia 86���107 % length of femur (mean 99 %); knee���knee distance 82���92 % of SVL (mean 87 %). Tadpole measurements (in mm). A stage 25 tadpole (MZUNAP-01- 521) was used for the description (Fig. 6). Total length 12.5; body length 4.7; internarial distance 0.9; eye to nares distance 0.8; eye diameter 0.5; interorbital distance 0.7; tail length 7.8. Tadpole description. Snout rounded when viewed from above; body ovoid in dorsal view. Eyes dorsal, angled laterally, pupils white in preservative. Nares not forming tube, situated half-way between eye and tip of snout, directed dorsolaterally. Spiracle sinistral; vent dextral. Ventral tail fin begins at tail base, dorsal tail fin begins just posterior to plane of vent opening, ventral and dorsal fins relatively uniform in thickness throughout tail, tapering towards tip. Musculature depth uniform throughout, tapering toward tip. Mouth directed anteroventrally. Oral disc emarginate, anterior and posterior labia forming flaps free from body wall, 1.4 mm in width. Marginal papillae absent on anterior labium, present in one complete row on posterior labium. Papillae rounded; submarginal papillae absent. Jaw sheaths deep in longitudinal width, serrate, lacking indentations. Lateral processes short, extending barely past lower jaw. Labial tooth row formula is 2 (2)/ 3 [1]. A- 1 complete (62 teeth), A- 2 with medial gap (34 teeth), same width as A- 1. P- 1 with medial gap (46 teeth), P- 2 (44 teeth), and P- 3 complete (44 teeth); P- 1 and P- 2 equal width, P- 3 slightly shorter. Color in Life. In life, the head appears light grey. Pigmentation on dorsum mottled brown, ground color weakly transparent. Eyes black, papillae white. Ventral coloration is transparent (most internal organs are visible) with irregular faint red flecking that is dense around mouth and nares. Tail musculature white with abundant brown mottling, fins almost transparent. Color in Preservative. In preservative, coloration is identical, though red pigmentation turns to brown. Vocalizations. The following values are presented as: min-max (average �� SD, number of individuals). The advertisement call is a short trilled note (Fig. 7) with duration between 630���880 ms (760 �� 140 ms, 7) and is repeated at irregular intervals of 2���7 notes per minute (3.14 �� 1.80 notes per minute, 6). Each note consists of 20���27 pulses (mean = 24). Calling activity is sporadic and continues throughout daylight, but peaks in the early morning and late afternoon. Dominant frequency is 5400���6000 Hz (5600 �� 2000 Hz, 5) at temperatures between 24.5���26 ��C. The call of R. yavaricola sounds similar to the calls of other species in the vanzolinii group, although there are some slight differences within this group. Ranitomeya flavovittata has slightly longer notes (1 sec vs. 760 ms in R. yavaricola). Ranitomeya imitator (n = 25) has a call that is slightly shorter in duration (686 ms vs. 760 ms in R. yavaricola) and has a slightly lower dominant frequency (5200 Hz vs 5600 Hz in R. yavaricola). Distribution and natural history. Ranitomeya yavaricola occurs in primary forests in a small area of northeastern Peru. This species likely occurs more widely within the broad interfluvium bordered by the Ucayali, Amazon, Yavar��, and Blanco rivers (Fig. 1). Much of the area bordering these rivers is seasonally or permanently flooded and it is possible that these large expanses impose limits to this species��� distribution (Fig. 8). Ranitomeya yavaricola is only known from forests near Lago Preto (but see discussion). Lago Preto is a large oxbow lake at the confluence of Yavar�� and Yavari-Mirin rivers. Near Lago Preto, several forest types are present: seasonally flooded forests, swamp forests that are saturated year-round, and upland terra firme forests. At Lago Preto, R. yavaricola occurs in upland forests and very low lying forests, just above the flood zone of seasonally flooded forests (i.e. v��rzea forests) and swamps. These sites contained many large trees and a relatively sparse understory. The upland forests in this region are some of the most diverse in the Amazon basin, with estimates of tree diversity exceeding 300 species per hectare in some areas (Pitman et al. 2003, Fine et al. 2006) These forests are dominated by trees of the families Fabaceae (legumes), Bombacaceae (���mallows��� such as kapoks), and Moraceae (figs) (Fine et al. 2006). This species is sympatric with three other dendrobatids: Ranitomeya uakarii, Ameerega hahneli, and A. trivittata. This species potentially also co-occurs with R. flavovittata and R. ventrimaculata, both of which have been observed less than 90 km from Lago Preto (Fig. 8). Ranitomeya yavaricola was typically observed foraging throughout the leaf litter or calling from the leaves of terrestrial palms (Genoma spp.), epiphytes (primarily bromeliads), and on the branches of fallen trees (between 0.2���3 m above the ground). Ranitomeya yavaricola is an extremely shy species and when encountered dives to the ground and hides within the leaf litter or within the roots of plants. A single tadpole was observed in the phytotelm of a small bromeliad that was ca. 1.5 m above the ground. Prior to the discovery of the tadpole, a male was observed calling near the bromeliad (Ranitomeya yavaricola is the most abundant species of Ranitomeya at Lago Preto. Using visual encounters, we observed 3 individuals in 30 man-hours of observation (0.1 encounters per man-hour) and using acoustic estimates, we recorded 20 different individuals in 3.7 man-hours of observation (5.6 encounters per man-hour). Lastly, as the result of casual observations, over a two week period in June 2009, 20 people encountered 6 individuals. During this study 12 individuals of R. uakarii were observed at Lago Preto (as the result of both casual observation and surveys). Conservation status. Following the IUCN Red List criteria (IUCN 2001), this species should be listed as Data Deficient (DD). It is currently known from only a single locality but probably occurs more widely., Published as part of Perez-Pe��a, Pedro E., Chavez, German, Twomey, Evan & Brown, Jason L., 2010, Two new species of Ranitomeya (Anura: Dendrobatidae) from eastern Amazonian Peru, pp. 1-23 in Zootaxa 2439 on pages 4-12, DOI: 10.5281/zenodo.194909, {"references":["Pitman, N., Vriesendorp, C. & Moskovits, D. (2003) Rapid biological inventory 11: Peru: Yavari. The Field Museum, Chicago, U. S. A., pp. 1 - 195.","Fine, P., Davila, N., Foster, R., Mesones, I. & Vriesendorp, C. (2006) Flora and vegetation. In: Vriesendorp, C., Pitman, N., Rojas-Moscoso, J. I., Pawlak B. A., Rivera-Chavez, L., Calixto-Mendez, L., Vela-Collantes, M., Fasabi-Rimachi, P. F. (Eds), Rapid biological inventory 16: Peru: Matses. The Field Museum, Chicago, U. S. A., pp. 174 - 183.","IUCN (2001) IUCN Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission, IUCN, Gland, Switzerland and Cambridge, UK."]}

Ameerega ignipedis sp. nov. Figures 1, 11, 16 Phyllobates petersi: Silverstone 1976 p. 37 (FMNH 56248 collected in 1947 by J. M. Schunke at ���Cerro Azul, 24 km E Contaman�����). Holotype. MUSM 24948, an adult female (Fig. 1) collected on 19 July 2006 by M. Pepper, E. Twomey, and J. Brown in Departamento Loreto, Peru, 17.5 km NE Contamana at the western foot of the Serran��a de Contamana, 240 m elevation, 7 �� 11 ��� 55.46 ��� S, 74 �� 57 ��� 35.28 ��� W. Type locality near ���El Uni��n���, a campsite located at the confluence of a hot-water and cold-water stream. Paratypes. Five adults (MUSM 24947, 24949��� 24952) collected on 19 July 2006, same locality as holotype. Etymology. The species name ignipedis is a Latin adjective meaning ���fiery-footed���, referring to the fact that the type locality is located alongside a geothermal stream. Our campsite in the Serran��a de Contamana was located at the confluence of two streams, one of which was fed by hot springs and reached temperatures exceeding 90 �� C in some places. We found A. ignipedis along the cool-water stream in low abundances, but along the hot-water stream they were much more common. Definition and diagnosis. Assigned to the genus Ameerega on the basis of the following: first finger longer than second, webbing absent between the toes, dorsal skin granular (Myers 1982, Grant et al. 2006). This is a small species of Ameerega with an adult SVL of approximately 20���24 mm. Dorsum granular and brown medially, black laterally; pale yellowish-green dorsolateral stripes extending from loreal region to groin. Pale yellowish-green labial stripe present starting behind nares and terminating above forelimb as a yellow patch. Yellow spots present and distinct above groin, most individuals (~ 60 %) also possess a yellow spot on the medial face of the tibia. Venter sky-blue with black marbling. Teeth present. Appressed first finger longer than second; finger discs weakly expanded; hands and feet lacking webbing between digits. Vocalization consists of a series of regularly-spaced notes occurring at a rate of ~ 2 notes per second; note length short (97 ms), dominant frequency of 4700 Hz. Ameerega ignipedis is the sister taxon to a larger clade containing Ameerega bassleri and the two additional new species described in this paper. However, A. ignipedis bears little resemblance to any of these species, as they are larger and usually have a yellow, orange, or red dorsum. Ameerega ignipedis bears a pattern most similar to A. pongoensis, A. petersi, A. simulans, and A. smaragdina, all of which have a brown or black dorsum with green or yellow dorsolateral stripes. For A. ignipedis individuals that possess a spot on the medial face of the tibia, diagnosis can be made on this character alone, as none of the four similar species possess such spots. Individuals that lack this shank spot can be diagnosed on the basis of body size, advertisement call, and ventral coloration. Both A. petersi and A. smaragdina have a larger SVL than A. ignipedis (Table 1). Furthermore, the dorsolateral stripes of A. smaragdina (and many populations of A. petersi) are emerald-green as opposed to pale yellowish-green in A. ignipedis. Both A. petersi and A. smaragdina have ventral coloration that is green and blue (Figures 11 & 12) (vs. uniform sky-blue in A. ignipedis), and A. smaragdina lacks black marbling on the venter (vs. black marbling present in A. ignipedis). Diagnosis against A. simulans can be made since A. simulans is smaller than A. ignipedis with respect to SVL (Table 1). Ameerega simulans also lacks spots above the axillae and groin (as opposed spots above axillae and groin present in A. ignipedis). Ameerega ignipedis can be distinguished from A. pongoensis in that the latter species lacks distinct yellow spots above the groin (vs. large, distinct spots in A. ignipedis), and by differences in advertisement call (Fig. 2). Ameerega pongoensis has a call consisting of a single- or double-peep which is repeated irregularly at a rate of one note (or note couplet) every ~ 2 seconds (vs. notes repeated regularly at a rate of two notes per second in A. ignipedis). Ameerega ignipedis can furthermore be diagnosed on the basis of 20 unambiguous nucleotide substitutions in the 16 s and CytB regions of the mitochondrial genome. Measurements (in mm) of holotype. The female holotype (Fig. 1) has SVL 23.8; FL 10.5; TL 11.1; KK 20.9; FoL 9.9; HaL 6.0; HL 6.0 HW 7.1; BW 7.0; UEW 4.6; IOD 3.4; IND 2.8; TD 2.6; ED 4.1; DET 1.4; L 1 F 4.7; L 2 F 2.9; W 3 D 0.9; W 3 F 0.6. Measurements of paratypes are given in Table 3. Description of holotype. Size medium, SVL 23.8 mm. Dorsal skin of head, body, and hind limbs granular; skin smooth or nearly smooth on forelimbs and smooth on sides of head and body and ventral surfaces. In life, the dorsum is brown medially with weak black marbling; laterally the dorsum is black. A pale yellow dorsolateral stripe present on either side beginning at nares, passing over the eyelids, and continuing posteriorly to terminate above the groin as a bright yellow spot. Stripe starts as pale yellow and blends to bright yellow towards the groin. Dorsolateral lines are ~ 50 % wider at groin than head. Pale yellow labial stripe present beginning near nares, extending posteriorly, terminating above axillae as a weakly defined yellow patch. Flanks black from groin to snout, ventral margin is pale yellow and fades to sky-blue ventrally. Upper forelimbs are yellow-bronze dorsally and conspicuously yellow ventrally; lower part of forelimbs brown dorsally and sky-blue with black reticulation ventrally. Hind limbs brown with irregular black markings dorsally; sky-blue with black reticulation ventrally. A conspicuous yellow spot is present on the medial surface of the tibia near the knee. Ventral surface of limbs, belly, and head sky-blue with a coarse reticulum of irregular black lines. Hands and feet brown dorsally. Iris black. Widest part of head between jaw articulations, head narrower than body; greatest head-width 29.9 % of SVL. Tongue medium sized, oval. Premaxillary and maxillary teeth present. Vocal slits absent. Snout narrow, sloping from lateral view; bluntly rounded from dorsal view; truncate from ventral view. Nares situated and directed posterolaterally to the tip of snout; nares visible from front and below but not from above. Canthus rostralis sloped, slightly rounded; loreal region nearly vertical and slightly concave. Upper eyelid 1.4 times wider than interorbital distance. Eye large and prominent with a maximum diameter of 17.2 % of the snout vent length, pupil rounded and horizontally elliptical. Tympanum circular, partially concealed posterodorsally, lacking tympanic annulus; its diameter less than 63.4 % of ED; supratympanic fold absent. Hands relatively small, length being 25.2 % of SVL. Relative length of appressed fingers: I ��� II ��� IV Variation in type series. The most notable variation in the type series is the yellow shank-spot present in four individuals and absent in two. The width of the oblique dorsolateral stripe and degree of reticulation varies between individuals (Figures 1 & 11); in some individuals this stripe blends completely with ventral reticulation. Within populations, the anterior terminus of dorsolateral lines varies between exterior edges of the eyelids to the superior part of the snout. Vocalizations. The advertisement call for A. ignipedis (Fig. 2) can be characterized as a ���retarded trill��� following L��tters et al. (2003). The call consists of regularly-spaced ���peeps��� which are repeated at a rate of about 1.7 notes per second. Note duration is 97 ms on average and notes are spaced 287���752 ms apart. Dominant frequency is slightly modulated, starting at 4398 Hz at the beginning of the note and ending at 4730 Hz. We witnessed a single male calling in the type locality after a heavy rain. Calling lasted for several minutes. The call sounds very similar to A. altamazonica (in fact their calls are nearly indistinguishable), although we have never encountered A. altamazonica in Contamana, or in any other site east of R��o Ucayali. In the type locality we also heard an unidentified species of Allobates calling whose call resembled that of A. ignipedis, but differed by having notes about 1000 Hz higher in pitch and half the duration of A. ignipedis. The call differs from that of A. pongoensis (Fig. 2), which consists of irregularly-spaced ���peeps��� repeated at a rate of 1 note every 2���3 seconds. These notes are sometimes given in couplets. Note duration 9���11 ms, dominant frequency 4233 Hz. Distribution and natural history. Ameerega ignipedis is known from two localities in the Serran��a de Contamana but probably occurs more widely (Fig. 3). Because these localities lie at the foot of low mountains, we suspect that this species may occur widely throughout the foothills of the Serran��a de Contamana, as well as other parts of the Sierra del Divisor. Few dendrobatid species are known to occur on both sides of Rio Ucayali, so it is probable that A. ignipedis occurs only to the east of this river. This species occurs in lower montane rainforest. Although the elevation is relatively low (240 m), this habitat is more reminiscent of higher elevation forests in the east Andean versant than the surrounding lowlands. For example, at the foot of the Serran��a de Contamana there is an abrupt change from typical lowland forest to a lush habitat which is characterized by abundant epiphytic plants, large tree ferns, and a comparatively moist leaf litter. This may be in part attributable to a slight Massenerhebung effect of these isolated hills, trapping moisture as clouds pass across the lowland plain. These hills also have a surprisingly diverse poison frog community. In addition to A. ignipedis, we found A. hahneli, A. trivittata, A. yoshina sp. nov., Ranitomeya lamasi, and R. ventrimaculata. Within this habitat, we found A. ignipedis near a small stream (Fig. 15 h). Most individuals were observed and collected in the forest near the stream, although we also saw a few individuals around the vegetation growing in the sandy banks, one of which was a male transporting tadpoles (which were preserved but later damaged). Since we found no other bodies of water nearby, we presume that he was in the process of depositing tadpoles in this stream. The use of flowing water for tadpole deposition is rare in Ameerega species, although has been observed in a few species such as A. cainarachi (pers. obsv.), A. macero (Rodriguez and Myers 1993), and A. pongoensis (M. Pepper, pers. comm.). Additionally, we found tadpoles of A. trivittata and what appeared to be a relatively large species of Leptodactylus. Fish were observed in the deeper pools of this stream. Males were heard calling throughout the day, particularly after rain. Conservation status. Following the IUCN Red List criteria (IUCN 2001), this species should be listed as Data Deficient (DD). Being known from only two localities that are less than 3 km apart, its extent of occurrence is unknown but probably occurs more widely. It is fairly common in the type locality and its habitat appears to be remote enough that deforestation risk is low., Published as part of Brown, Jason L. & Twomey, Evan, 2009, Complicated histories: three new species of poison frogs of the genus Ameerega (Anura: Dendrobatidae) from north-central Peru, pp. 1-38 in Zootaxa 2049 on pages 5-10, DOI: 10.5281/zenodo.186518, {"references":["Silverstone, P. A. (1976) A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra (Family Dendrobatidae). Bulletin of the Natural History Museum of Los Angeles County, 27, 1 - 53.","Myers, C. W. (1982) Spotted poison frogs: Descriptions of three new Dendrobates from western Amazonia, and resurrection of a lost species from \" Chiriqui \". American Museum Novitates, 272, 1 - 23.","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, B. D., Noonan, B. P., Schargel, W. C. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Anura: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262.","Rodriguez, L. & Myers, C. W. (1993) A new poison frog from Manu National Park, southeastern Peru (Dendrobatidae, Epipedobates). American Museum Novitates, 3068, 1 - 15."]}

Ameerega pepperi sp. nov. Figures 1, 14��� 16 Phyllobates bassleri: Silverstone 1976 p. 45 (AMNH 42327, 42867, 43402 collected in 1926 and 1928 by H. Bassler at ���Pachiza���). Holotype. MUSM 26940 (Fig. 16), an adult male collected on 14 July 2006 by M. Pepper, Provincia Tocache, Departamento San Mart��n, Peru, 2 km NE of San Francisco, 980 m elevation, 8 �� 18 ' 30.3 " S, 76 �� 40 ' 37.6 " W. Found on the ground near a small waterfall. Paratypes. All from San Mart��n, Peru. One adult female (MUSM 26942), collected 21 July 2006 by M. Pepper and E. Twomey, and two adult females (MUSM 26975, 26976), collected 25 June 2007 by J. Brown, E. Twomey, and K. Fieselman, 390 m elevation, 7.0 km SW of Huicungo at 7 �� 21 ' 58.6 " S, 76 �� 48 ' 44.1 " W: Two adult females (MUSM 26979, 26980), collected on 25 June by J. Brown, E. Twomey, and K. Fieselman, 384 elevation, 6.6 km N of Campanilla at 7 �� 25 ' 38.6 " S, 76 �� 39 ' 53.3 " W: One adult female (MUSM 26968), collected on 15 July 2006 by M. Pepper, 570 m elevation, SW of Chumanza (alternate spelling: Shumanza) at 7 �� 34 ' 7.8 " S, 76 �� 41 ' 34.4 " W: One adult male (MUSM 26941), collected at same locality of holotype, 980 m elevation, 2 km NE of San Francisco at 8 �� 18 ' 30.3 " S, 76 �� 40 ' 37.6 " W. Etymology. The specific epithet is a patronym for the discoverer of this species, Mark Pepper, an enthusiastic Canadian explorer and conservationist whose discoveries have greatly contributed to the authors��� understanding of poison frog biogeography and taxonomy. Definition and diagnosis. Assigned to the genus Ameerega on the basis of the following: first finger longer than second, webbing absent between the toes, dorsal skin granular (Myers 1982, Grant et al. 2006). This is a medium-large species of Ameerega with an adult SVL of approximately 28���34 mm. Dorsum has a black ground color, finely stippled brick-red to bright red in southern populations (e.g. San Francisco) to coarsely stippled orange or yellow northern populations (e.g. Campanilla), stippling concentrated near head and is less dense near sacral region. Bright yellow dorsolateral stripes extend from the eyelid to the groin; a yellow labial stripe is also present starting near nares and extending posterior to the axillary region. Faded yellow-green to white spot present above groin in some populations, shank spots and axillary spots absent. Limbs greenish-bronze or black; venter blue with strong black marbling. Teeth present. Appressed first finger longer than second; finger discs moderately expanded, hands and feet lacking webbing between digits. Vocalization consists of a loud retarded trill of whistle-like notes given at a rate of 0.9���1.3 notes per second; dominant frequency 2800���3160 Hz (Fig. 4, Table 2). Ameerega pepperi can be distinguished from most other species of Ameerega by its red, orange, or yellow dorsum. It is similar in appearance to the following species: A. bassleri, A. bilinguis, A. cainarachi, A. macero, A. parvula, and A. yoshina, all of which have (or can have) a solid red, orange, or yellow dorsum. Ameerega parvula and A. bilinguis both lack distinct dorsolateral stripes (vs. dorsolateral stripes present and conspicuous in A. pepperi). Ameerega cainarachi is smaller, females up to 31.3 mm SVL (vs. 34 mm SVL in A. pepperi), and has an advertisement call consisting of a regular chain of notes which are not frequency modulated, repeated at a rate of 9 notes per second (vs. ���bursts��� of frequency-modulated notes, repeated 0.9���1.3 notes per second in A. pepperi). Ameerega macero is smaller than A. pepperi, females up to 29.5 mm SVL, and has a call consisting of a series of harsh ���peeps���, repeated regularly at a rate of 10 notes per second. Ameerega bassleri has an advertisement call consisting of a series of regularly-spaced whistle-like notes given continuously for several minutes at a rate of 1.6���2.1 notes per second (vs. notes given at a rate of 0.9���1.3 notes per second in A. pepperi). Ameerega yoshina has an advertisement call given in bursts of 4���31 notes, given at a rate of 3.7���4.1 notes per second, each burst less than 8 seconds in total duration (vs. notes given at a rate of 0.9���1.3 notes per second in A. pepperi). Measurements (in mm) of holotype. The male holotype (Fig. 16) has SVL 28.6; FL 13.4; TL 16.1; KK 28.2; FoL 13.3; HaL 7.5; HL 7.7; HW 7.8; BW 8.3; UEW 5.5; IOD 3.9; IND 4.0; TD 3.0; ED 4.2; DET 1.6; L 1 F 6.9; L 2 F 6.1; W 3 D 0.8; W 3 F 0.6. Measurements of paratypes are given in Table 5. Description of holotype. Size medium, SVL 28.6 mm (Table 5). Dorsal skin is coarsely granular and is pigmented black with brick red mottling densest near the snout. Thin yellow dorsolateral stripes extend from groin, extending dorsally around snout; posterior to eyes the stripes blend into the red dorsum. Dorsolateral stripe is sightly wider at groin. Flanks are black extending from groin to nares. Forelimbs lack granulation and are colored tan and flecked in olive. Yellow coloration extends up upper arm surface, anteriorly around lip. Legs are coarsely granulated; contain yellow femoral spots, dorsal surface colored olive green. The venter is yellow-gold with olive marbling; marbling extends primarily on throat and margins of legs. Iris black. Widest part of head between jaw articulations and tympanum, head narrower than body. Greatest head width 27.0 % of SVL. Eyes very protuberant. Tongue medium sized, oval. Premaxillary and maxillary teeth present. Vocal slits present. Dorsal skin of head and body is finely granular; skin smooth or nearly smooth on limbs and smooth on sides of head and body and ventral surfaces. Snout sloping laterally; bluntly rounded dorsally; truncate ventrally. Nares situated and directed posterolaterally to the tip of snout; nares visible from front and below but not from above. Canthus rostralis sloped, slightly rounded; loreal region nearly vertical and slightly concave. Upper eyelid 1.4 times wider than interorbital distance. Eye large and prominent with a maximum diameter of 14.6 % of SVL, pupil rounded and horizontally elliptical. Tympanum circular, without tympanic annulus its diameter less than 73.0 % of ED, supratympanic fold absent. Hands relatively small, length being 26.2 % of SVL. Relative length of appressed fingers I ��� II ��� IV Variation. The dorsal coloration varies over a cline from the north to south. In southern populations, the dorsum is finely stippled brick-red to bright red and gradually changes to coarsely stippled orange or yellow in northern populations (Figures 14 & 15). Vocalizations. The advertisement call for A. pepperi (Fig. 4) can be classified as a ���retarded trill��� following L��tters et al. (2003). The call consists of a series of musical notes resembling short ���whistles��� at a rate of 0.9���1.3 notes per second. Notes are 130���220 ms long, spaced 560���1030 ms apart, overall dominant frequency 2970 Hz. The call of A. pepperi can be distinguished from the advertisement call of A. bassleri by the rate the notes are repeated, with A. bassleri repeating notes at 1.6���2.1 notes per second (vs. 0.9���1.3 in A. pepperi). Distribution and natural history. Ameerega pepperi is known from throughout the upper Huallaga Valley, south of R��o Huayabama (near Huicungo) to the southern border of San Mart��n at elevations from 380 m to approximately 1000 m elevation. Most of the known localities for this species run along the main road to Tingo Maria, which is primarily on the eastern side of R��o Huallaga (i.e., the western slope of the Cordillera Azul). This species also occurs on the western side of the Huallaga (i.e., the eastern slope of the Andes) in San Francisco and Huicungo, and likely occurs throughout the area between these two sites based on the results of our niche model (Fig. 17). This model also predicts suitable habitat throughout much of the R��o Biabo valley in the Cordillera Azul, although no surveys have taken place in this area. There appears to be very little predicted overlap in the distributions of A. pepperi and A. bassleri in the niche model (Fig. 17), with the only overlapping areas occurring near the town of Juanjui. In 2007 we spent a day in the mountains just west of Juanjui and did not detect either species. The area near Juanjui appears to be somewhat drier than surrounding areas, and this may function as a weak barrier that separates the distributions of these two species. Ameerega pepperi occurs in undisturbed premontane forests and secondary premontane forest, particularly in habitats which are adjacent to streams (Fig. 15 i). Unlike A. bassleri, which we have often observed far from streams (> 300 m), we have never observed A. pepperi greater than ~ 30 m from a stream. This species can occur in moderately disturbed areas provided the streamside habitats are relatively shaded. During the day, adults tend to hide amongst streamside boulders. Calling activity seems to peak during late evening, just prior to dusk. Males call from elevated positions on boulders, although courtship and oviposition appears to take place on the ground in the leaf litter. Clutches contain 22- 44 eggs and are guarded by males (M. Pepper pers. comm.). These eggs typically hatch in approximately 18 days. Tadpoles have been found in shallow eddies or streamside pools which are sometimes left by receding water levels. Clay-bottomed streams appear to be most commonly used, and tadpoles will often submerge themselves under the sediment, particularly during the hotter hours in the afternoon. These pools also contain tadpoles of the sympatric A. altamazonica and A. trivittata. Tadpoles of Ameerega pepperi complete development in as little as 5-6 weeks. Conservation status. Following the IUCN Red List criteria (IUCN 2001), we tentatively suggest that A. pepperi should be listed as Least Concern (LC) based on the following: (1) we estimate the extent of occurrence to be at least 5,500 km 2, (2) populations appear to be continuous and (3) this species occurs in largely undisturbed forests, though there has been a small amount of potential habitat loss in the Cordillera Azul and in the upper Huallaga Valley., Published as part of Brown, Jason L. & Twomey, Evan, 2009, Complicated histories: three new species of poison frogs of the genus Ameerega (Anura: Dendrobatidae) from north-central Peru, pp. 1-38 in Zootaxa 2049 on pages 16-18, DOI: 10.5281/zenodo.186518, {"references":["Silverstone, P. A. (1976) A revision of the poison-arrow frogs of the genus Phyllobates Bibron in Sagra (Family Dendrobatidae). Bulletin of the Natural History Museum of Los Angeles County, 27, 1 - 53.","Myers, C. W. (1982) Spotted poison frogs: Descriptions of three new Dendrobates from western Amazonia, and resurrection of a lost species from \" Chiriqui \". American Museum Novitates, 272, 1 - 23.","Grant, T., Frost, D. R., Caldwell, J. P., Gagliardo, R., Haddad, C. F. B., Kok, P. J. R., Means, B. D., Noonan, B. P., Schargel, W. C. & Wheeler, W. C. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Anura: Athesphatanura: Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1 - 262."]}