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We present the results of herpetological surveys in two adjacent mountains where the EcoMinga Foundation protects the cloud forest in the Upper Rio Pastaza watershed, in the Llanganates Sangay Ecological Corridor in Ecuador. A rapid assessment of the amphibian communities of the study sites reveals a diverse and heterogeneous composition, dominated by terrestrial frogs from the genus Pristimantis. We also identify a cryptic diversity with a significant number of candidate new species. We describe two new species of terrestrial frogs of the genus Pristimantis. Pristimantis maryanneaesp. nov. is characterised by not having tympanum externally visible and having 2–3 subconical tubercles in the upper eyelid; and Pristimantis burtoniorumsp. nov. is characterised by the presence of red colouration in hidden surfaces of the hind-limbs, tubercles on the upper eyelid, interorbital tubercle and a row of rounded tubercles along the snout to the tip and a pale red venter with dark brown mottled pattern. Our samples from the two Reserves do not share species between them, so the proportion of shared species seems to be relatively low. In addition, we highlight the importance of updating the knowledge of amphibians that are restricted to this important conservation region and comment about the threats and composition of the amphibian communities on the eastern slopes of the Upper Rio Pastaza watershed.

The Neotropical frog genus Cryptobatrachus includes six currently recognized species distributed throughout the northern Andes in Colombia and Venezuela. Cryptobatrachus conditus, C. pedroruizi, and C. remotus were described from the Colombian and Venezuelan slopes of the Serranía del Perijá. Due to the great morphological similarity among these species, we re-assess their taxonomic status based on morphological and molecular data from types, topotypes, and specimens from localities referred to in the original descriptions of these species. Morphometric comparisons showed that all these species are virtually indistinguishable, although some subtle qualitative differences in morphological traits distinguish C. conditus from the other species. Phylogenetic analyses of DNA sequences found that C. remotus and C. pedroruizi are not reciprocally monophyletic and exhibit low genetic divergence (< 1 %). Therefore, C. remotus should be considered a junior synonym of C. pedroruizi. This work stresses the importance of comprehensive taxon sampling in poorly explored areas, especially between neighbouring countries.

[EN] The genus Gastrotheca (Anura: Hemiphractidae) is a group of marsupial frogs particularly diverse in Andean regions. Several taxonomic studies of this genus have been conducted in the humid cloud forests—or Yungas—of the Andean eastern slopes of central Bolivia (departments of Cochabamba and Santa Cruz). Yet, the distinction among three species that occur sympatrically in these forests, G. lauzuricae (proposed as a junior synonym of G. coeruleomaculatus in 2015), G. piperata, and G. splendens, remains unclear since the morphological characters that purportedly support their differentiation are variable and partly shared among them. We have carried out external morphological studies, including multivariate morphometric analyses, to assess how they support the taxonomic status of these three species. We also evaluated characters of the cranial osteology of a sample of six individuals using micro CT-scanning. Principal component and linear discriminant analyses resulted in a great overlap among the putative species. Cranial osteological comparisons did not reveal highly significant differences among them, but suggested that different degrees of hyperossification could be related to the developmental state of individuals. Our results indicate that most morphological and osteological reported differences between the three species likely represent intraspecific variation. Thus, we propose that the three nominal species belong to a single biological entity, for which the name Gastrotheca splendens (Schmidt, 1857) has priority. We also restrict the name Gastrotheca coeruleomaculatus (Werner, 1899) to externally similar congeneric populations from the Yungas forests of department of La Paz, but highlighting the need of a detailed evaluation of their taxonomic identity., [ES] El género Gastrotheca (Anura: Hemiphractidae) es un grupo de ranas marsupiales altamente diverso, especialmente en las regiones andinas. Varios estudios taxonómicos de este género se han llevado a cabo en los bosques húmedos montañosos—o Yungas—de la vertiente oriental de los Andes del centro de Bolivia (departamentos de Cochabamba y Santa Cruz). La distinción entre tres especies que se encuentran en simpatría en estos bosques,G. lauzuricae (propuesta como sinónimo posterior de G. coeruleomaculatus), G. piperata y G. splendens, no está clara, ya que los caracteres morfológicos que supuestamente apoyan su diferenciación son variables y están parcialmente compartidos entre ellas. Mediante estudios de morfología externa—incluyendo análisis morfométricos multivariantes—evaluamos la validez taxonómica de estas tres especies. Además, examinamos la osteología craneal de una muestra de seis individuos con microtomografía computarizada. Los análisis de componentes principales y discriminante lineal mostraron un gran solapamiento entre las supuestas especies, y las comparaciones osteológicas no revelaron diferencias sustanciales entre ellas, pero sí sugieren que los diferentes grados de hiperosificación podrían estar relacionados con el estado de desarrollo de los individuos. Nuestros resultados indican que la mayoría de las diferencias morfológicas y osteológicas descritas para las tres especies probablemente representan variación intraespecífica, por lo que proponemos la existencia de una única entidad biológica, para la que el nombre Gastrotheca splendens (Schmidt, 1857) tiene prioridad. Se establece que el nombre aplicable a las poblaciones similares de Gastrotheca de los Yungas del departamento de La Paz sería Gastrotheca coeruleomaculatus (Werner, 1899), pendiente de posteriores estudios taxonómicos., This paper was financed by projects CGL2014-53523-P and PGC2018-097421-B-I00 (PI, Ignacio De la Riva) of the Spanish Ministry of Science and Innovation, partially with funds from the European Union.

We describe a new species of marsupial frog, genus Gastrotheca, using morphological characters and molecular data as lines of evidence. The new species was discovered in the páramo and the ecotone between páramo and humid montane forest of Cordillera de Colán, at elevations between 3136 and 3179 m a.s.l., in northeastern Peru. The new species is distinguished from all its congeners by the combination of the following characters: coarsely granular skin on dorsum, a green dorsal coloration without pattern, finger I shorter than finger II, turquoise iris, and a venter without blotches, flecks or dots. Furthermore, we include a detailed osteological description of the new Gastrotheca species based on Micro-CT scanning. Based on our phylogenetic analyses, the new species belongs to the Gastrotheca marsupiata species group, is sister to G. oresbios and closely related to G. psychrophila, G. spectabilis, G. stictopleura and one undescribed species. Additionally, we test for the presence of the fungal pathogen Batrachochytrium dendrobatidis (Bd). No Bd infection was detected for G. gemma sp. nov. specimens but Bd prevalence was detected among syntopic frogs.

This article provides a brief account of the career of Eugenia M. del Pino. Casual events and serendipity played important roles in modeling her career as a developmental biologist. In collaboration with colleagues and students, she analyzed the biology and development of the marsupial frog Gastrotheca riobambae (family: Hemiphractidae) in comparison with Xenopus laevis and tropical frogs. The emphasis was placed on oogenesis and the early stages of development. Topics include the mono- and multi-nucleated modes of oogenesis. She described two modes of gastrulation in frogs, gastrulation modes one and two, according to the timing of notochord elongation. She was able to establish a pioneer laboratory for the comparative analysis of frog development in Ibero America at the Pontifical Catholic University of Ecuador, in Quito. Her contributions to society include her influence in the establishment of the National Academy of Sciences of Ecuador, and efforts toward the conservation of the Galápagos Archipelago. She is part of a pioneer group of professors that placed Biology as an academic discipline in Ecuador. The experiences of her career reveal that we all face difficulties in our jobs. However, nothing is impossible when we follow a passion. Her work reveals that the key to success is to turn obstacles into opportunities.

Dollo's law of irreversibility states that once a complex structure is lost, it cannot be regained in the same form. Several putative exceptions to Dollo's law have been identified using phylogenetic comparative methods, but the anatomy and development of these traits are often poorly understood. Gastrotheca guentheri is renowned as the only frog with teeth on the lower jaw. Mandibular teeth were lost in the ancestor of frogs more than 200 million years ago and subsequently regained in G. guentheri. Little is known about the teeth in this species despite being a frequent example of trait "re-evolution," leaving open the possibility that it may have mandibular pseudoteeth. We assessed the dental anatomy of G. guentheri using micro-computed tomography and histology and confirmed the longstanding assumption that true mandibular teeth are present. Remarkably, the mandibular teeth of G. guentheri are nearly identical in gross morphology and development to upper jaw teeth in closely related species. The developmental genetics of tooth formation are unknown in this possibly extinct species. Our results suggest that an ancestral odontogenic pathway has been conserved but suppressed in the lower jaw since the origin of frogs, providing a possible mechanism underlying the re-evolution of lost mandibular teeth.

Made available in DSpace on 2021-06-25T10:49:37Z (GMT). No. of bitstreams: 0 Previous issue date: 2021-01-15 Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Aplectana longa n. sp. (Ascaridida: Cosmocercidae) from the small intestine of Gastrotheca microdiscus (Amphibia: Hemiphractidae) is described and illustrated. The new species is characterized by the combination of a unique set of morphological characters: 1) Large body size in both sexes; 2) Lateral alae absent; 3) Gubernaculum present, small, thin, weakly sclerotized, pointed at the distal end and curved proximally at the end, with a small and punctiform papilla-like dilation, and 4) Caudal papillae arrangement (9+1:0:6). The distribution pattern of caudal papillae is similar only to Aplectana chamaeleonsis. Nevertheless, Aplectana longa n. sp. is easily differentiated from this species by the arrangement of precloacal papillae. This is the 57th species of the Aplectana and the 16th species reported from Brazil. Universidade Estadual Paulista (UNESP) Campus de Botucatu Instituto de Biociências Setor de Parasitologia, R. Prof. Dr. Antônio Celso Wagner Zanin, 250-Distrito de Rubião Junior Universidade Federal da Bahia Instituto de Biologia Campus de Ondina Universidade Federal de Uberlândia (UFU) Instituto de Ciências Agrárias, LMG-746, Km 1, Monte Carmelo Universidade Estadual Paulista (UNESP) Campus de Botucatu Instituto de Biociências Setor de Parasitologia, R. Prof. Dr. Antônio Celso Wagner Zanin, 250-Distrito de Rubião Junior CNPq: 140871/2017-9 CNPq: 141322/2018-7 CNPq: 150041/2017-9 CNPq: 309125/2017-0 CNPq: 313241/2018-0

We study the phylogenetic relationships of egg-brooding frogs, a group of 118 neotropical species, unique among anurans by having embryos with large bell-shaped gills and females carrying their eggs on the dorsum, exposed or inside a pouch. We assembled a total evidence dataset of published and newly generated data containing 51 phenotypic characters and DNA sequences of 20 loci for 143 hemiphractids and 127 outgroup terminals. We performed six analytical strategies combining different optimality criteria (parsimony and maximum likelihood), alignment methods (tree- and similarity-alignment), and three different indel coding schemes (fifth character state, unknown nucleotide, and presence/absence characters matrix). Furthermore, we analyzed a subset of the total evidence dataset to evaluate the impact of phenotypic characters on hemiphractid phylogenetic relationships. Our main results include: (i) monophyly of Hemiphractidae and its six genera for all our analyses, novel relationships among hemiphractid genera, and non-monophyly of Hemiphractinae according to our preferred phylogenetic hypothesis; (ii) non-monophyly of current supraspecific taxonomies of Gastrotheca, an updated taxonomy is provided; (iii) previous differences among studies were mainly caused by differences in analytical factors, not by differences in character/taxon sampling; (iv) optimality criteria, alignment method, and indel coding caused differences among optimal topologies, in that order of degree; (v) in most cases, parsimony analyses are more sensitive to the addition of phenotypic data than maximum likelihood analyses; (vi) adding phenotypic data resulted in an increase of shared clades for most analyses., Funding for this study was provided by Conselho Nacional de Desenvolvimento Cientıfico e Tecnologico (CNPq), Brazil to LYE and SC-F (scholarship numbers 830432/1999-0 and 312744/2017-0), Coordenacao de Aperfeicoamento de Pessoal de Nıvel Superior (CAPES), Brazil to LYE (scholarship number 88887.179352/2018-00) and projects CGL2014-53523-P and PGC2018-097421-B-100 of the Spanish Government (PI, Ignacio De la Riva).

We studied the amphibian community of the Parque Nacional da Serra dos Órgãos (PARNASO) for over thirty years. The area of 20,024 hectares has a steep altitudinal gradient (200-2,263 m a.s.l.), and it is located in the municipalities of Guapimirim, Magé, Petrópolis and Teresópolis, middle of the state of Rio de Janeiro, Brazil. Most data were obtained from sampling sites in the municipality of Teresópolis, with additional data from zoological collections and bibliography. We recorded 83 amphibian species distributed in two orders, Anura, 13 families: Aromobatidae (1), Brachycephalidae (11), Bufonidae (5), Centrolenidae (2), Craugastoridae (2), Cycloramphidae (8), Hemiphractidae (7), Hylidae (28), Hylodidae (6), Leptodactylidae (5), Microhylidae (1), Odontophrynidae (3), Phyllomedusidae (3) and Gymnophiona, one family: Siphonopidae (1). In addition, we present six species that occurs in the buffer zone. Ten of these species are endemic of the park, 18 have PARNASO as its type locality, and five the type locality is at the buffer zone. Resumo: Estudamos a comunidade de anfíbios do Parque Nacional da Serra dos Órgãos (PARNASO) por mais de 30 anos. A área de 20.024 hectares possui um gradiente altitudinal de 200 a 2.263m acima do nível do mar, e está localizada nos municípios de Guapimirim, Magé, Petrópolis e Teresópolis, que ficam no meio do estado do Rio de Janeiro, Brasil. A maioria das coletas foram feitas no município de Teresópolis, com dados adicionais de outras coleções zoológicas e de bibliografia. Foram registradas ao todo 83 espécies de anfíbios distribuídos em duas ordens, Anura, 13 famílias: Aromobatidae (1), Brachycephalidae (11), Bufonidae (5), Centrolenidae (2), Craugastoridae (2), Cycloramphidae (8), Hemiphractidae (7), Hylidae (28), Hylodidae (6), Leptodactylidae (5), Microhylidae (1), Odontophrynidae (3), Phyllomedusidae (3) e Gymnophiona, uma família: Siphonopidae (1). Foram registradas, também, seis espécies que ocorrem na zona de amortecimento. Destas espécies 10 são endêmicas, 18 têm o PARNASO como localidade tipo e cinco têm a localidade tipo na zona de amortecimento.

An inventory of the amphibians of the Reserva Ecológica Michelin – REM in southern Bahia, Brazil is presented. Sixty-nine species were recorded during a ten-year sampling period. Amphibians were distributed in two orders (Gymnophiona and Anura), belonging to twelve families [Aromobatidae (1), Bufonidae (3), Centrolenidae (1), Craugastoridae (5), Eleutherodactylidae (3), Hemiphractidae (2), Hylidae (34), Phyllomedusidae (5) Leptodactylidae (7), Microhylidae (4), Odontophrynidae (3) and Caeciliidae (1)]. Fifty per cent of the reproductive modes known for Atlantic forest anurans were recorded. While no threatened species were found at REM, six species are classified as data deficient (DD) by the Brazilian Red List of threatened species and deserve additional attention.PhasmahylatimboandVitreoranaeurygnataare listed as endangered in Bahia according to the list of threatened species of the state. Despite a higher diversity of amphibians in the Atlantic forest having been reported for mountainous regions, our results revealed that amphibian richness for lowland forests is also high.

Report the first records of Gastrotheca longipes, Cochranella resplendens and Teratohyla midas in the Allpahuayo-Mishana National Reserve, Loreto region, Peru. We update the distribution maps of these three species within the Amazon basin, provide information about their natural history, and describe puffing-up body and mouth gapping behaviours in Gastrotheca longipes. Revisado por pares

Gastrotheca yacuri sp. nov. urn:lsid:zoobank.org:act: 6261932E-02AB-4C9E-B2BD-BD213BBDC771 Holotype. CJ 7822 (Figs. 21–22), an adult male, from El Salado de Jimbura, at about 2.5 km (by road) west from the entrance in the Parque Nacional Yacuri, 2914 m (04° 42' 08.37" S, 79° 27' 00.09" W), Loja Province, Ecuador, collected on 9 April 2016 by Paul Székely and Diana Székely. Paratype. Ecuador: Loja: QCAZ 21105, adult male, from El Salado de Jimbura, 2712 m, on 26 June 2002 by Fernando Nogales Sornoza. Referred specimen. Ecuador: Loja: Not collected (Fig. 11O), subadult, 43 mm SVL, from nearby of Lagunas Negras, Parque Nacional Yacuri, 3492 m (04° 42' 31.81" S, 79° 25' 49.61" W). Diagnosis. Included in the genus Gastrotheca by molecular evidence and general morphological similarity with species of Gastrotheca. Although females are unknown, we infer presence of dorsal brood pouch because all its closely related species have it. A moderately large species (57.4–58.9 mm SVL in males, n = 2) with tibia length 47¯49% SVL, larger than foot; (2) interorbital distance greater than width of upper eyelid; (3) skin on dorsum weakly granular to smooth, co-ossified with skull, having a transverse ridge at occipital region; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I shorter than Finger II, width of discs much wider than digits; (8) fingers unwebbed; (9) toes one-half webbed, webbing extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum uniform green or brown with brown paravertebral marks; (11) head markings consisting of brown labial stripe with cream glandular areas at posterior end, canthal and supraciliary stripe of black, brown, and bronze; (12) dorsolateral stripe absent or present, when present above consisting in a series of bronze and cream warts, bordered below with a black and brown line, (13) flanks brown and green with a blue and black reticulum towards the lower flanks, axillae, groin, anterior and posterior surfaces of thighs, shanks, and tarsus; brown pelvic patch; (14) venter cream with nearly uniformly distributed, small, dark brown marks or brown with cream marks; gular region dark brown in males. Gastrotheca yacuri differs from all other species of Gastrotheca in southern Ecuador by having a blue and black reticulated pattern on flanks, axilla, groin, thighs, and shanks. It most closely resembles two other species, G. pseustes and G. psychrophila by having the blue colors, but they lack a blue and black reticulum (compare these species in Fig. 10). Gastrotheca yacuri differs from its sister species, G. aguaruna, from the Cordillera Central in northern Peru, by lacking blue colors, having the skin co-ossified with skull, and being smaller (SVL of males of G. yacuri 57.4–58.9 mm vs 41.6–46.8 mm in G. aguaruna). The genetic distance between them is of at least 2.8 % (in a DNA dataset of 438 bp, 16S gene). Description of the holotype. An adult male (Figs. 21–22); body moderately robust; SVL 57.4 mm; head wider that long; snout slightly acuminate in dorsal view, bluntly rounded in profile; canthus rostralis angular in section; loreal region concave; lips rounded; top of head flat; interorbital distance 92% of width of upper eyelid; internarial area flat; nostrils not protuberant, directed anterolaterally, at about level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye by distance about equal to diameter of tympanum; tympanic evident; supratympanic fold elevated, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing four teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 28% of SVL); fingers unwebbed bearing fringes; discs large and rounded, width of disc of Finger III lesser that diameter of tympanum; relative lengths of fingers I Skin on dorsum smooth to shagreen; smooth occipital ridge present; U-shaped ridges (vestigial) in sacrum region; skin on flanks rugose and coarsely areolate; skin on throat loose bearing folds of smooth vocal sac; skin on ventral surfaces of thighs, arms, and belly heavely areolate; skin on venter surface of shanks smooth; rugose surfaces around cloacal opening. Tongue broad, suboval, not notched posteriorly, fully attached to mouth floor. Coloration in life. The dorsal surfaces of the head, body, and limbs are green; the hidden surfaces of the forearm, thighs, and shanks are reticulated with black and blue markings (Figs. 21–22). Head markings are brown canthal and labial stripes. The canthal stripe is diffuse; the labial stripe varies from brown anteriorly to white posteriorly until the insertion of the forelimb. The dorsolateral stripe is thin and consists of a series of bronze and cream warts, bordered below with a black and brown line. The flanks are brown and green with a blue and black reticulum towards the lower flanks, axillae, groin, anterior and posterior surfaces of thighs, shanks, and tarsus; the pelvic patch is brown. The venter is cream with nearly uniformly distributed, small, dark brown marks; gular sac is nearly uniform grayish-brown with small dark brown marks. Palms and soles are pinkish gray; ventral fingers and toes are fleshy. The iris is copper with abundant black reticulations. Coloration in preservative. Similar to coloration in life, but green and blue surfaces have begun to fade. Measurements (in mm). SVL: 57.4, TIBL: 27.2, FL: 24.8, HL: 17.1, HW: 19.2, IOD: 6.9, EW: 5.2, IND: 2.8, ED: 6.0, EN: 3.0, TD: 3.3, FFL: 8.2, TFL: 16.2, TFD: 2.8. Variation. Morphometric variation of the male paratype is as follows: SVL: 58.9, TIBL: 28.6, FL: 28.3, HL: 17.9, HW: 20.1, IOD: 6.7, EW: 4.3, IND: 2.8, ED: 5.6, EN: 5.0, TD: 3.0, FFL: 10.8, TFL: 19.0, TFD: 2.7. The skin on the dorsum varies in the subadult by lacking a smooth tranverse occipital ridge. Color variation in life. The referred subadult specimen varies from the holotype in that it is mostly brown; it has a light reddish-brown dorsum of body and limbs with darker, well-defined stripes, two paravertebral and one at the coccyx position; scattered black spots are present on dorsum towards the flanks (Fig. 10O). The loreal region is brown, and a thin dark brown canthal line is present; the labial stripe is dark brown and bordered along its upper margin by a thin pale stripe that varies from brown anteriorly to cream posteriorly until the insertion of the forelimb. There is no dorsolateral stripe but the reddish-brown dorsal color contrasts with a broad dark brown band on the flanks. The flanks are brown with black and cream marks (without blue colors). The venter is mostly brown with cream marks that are larger towards the chest; the gular region is slightly darker that the venter. The palms and soles are gray. The iris is cooper with black reticulations and has a shade of red in the lower half. Vocalization. Four individuals of Gastrotheca yacuri were recorded at El Salado, Loja Province (Appendix III). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call is a complex call, composed of one to three (usually two) long pulsed notes and followed (or not) by one to three (usually two) short, single-pulsed notes (Fig. 23 A–G). The long note had a mean duration of 0.634 s (SD = 0.091) and consisted on average of 32.24 (SD = 7.504) distinct pulses, partly fused, without silent intervals (amplitude modulation close but less than 100%). The amplitude of the long note increases gradually towards the middle of the note after which it decreases gradually by the end. The short notes had a mean duration of 0.072 s (SD = 0.020) and the inter-note interval was on average of 0.549 s (SD = 0.214). The mean dominant frequency of the call was 1496.7 Hz (SD = 38.643), with a mean 90% bandwidth frequency of 1366.7–1637.9 Hz, having the highest values recorded among the analyzed species. The fundamental frequency is recognizable; when visible, 5 to 7 harmonics are distinguishable. Comparisons. The advertisement call of Gastrotheca yacuri is most similar to that of G. pseustes, but G. yacuri has a much shorter call duration, shorter long notes duration, shorter inter-note interval, a higher dominant frequency, and a higher 90% bandwidth frequency compared with the call of G. pseustes (Table 5). Also, G. yacuri emits usually only two short notes compared with 4–6 short notes of G. pseustes. The call of G. yacuri can be easily distinguished form those of G. lojana and G. testudinea by the amplitude modulation of the longer note, longer call duration, lower short note rate, larger number of pulses, higher pulse rate, a much higher dominant frequency, and higher 90% bandwidth frequency. Also, G. yacuri emits up to three (usually two) long notes per call compared with the only one emitted by G. lojana and G. testudinea (Table 5). Distribution and ecology. Gastrotheca yacuri is only known from three nearby localities 2.8 km apart (maximum distance) at elevations of 2712–3492 m in the Cordillera Oriental de los Andes in southern Ecuador. One locality is within Parque Nacional Yacuri and the others are in El Salado de Jimbura in Loja Province (Fig. 9). This nocturnal, semiarboreal species inhabits paramo and subparamo in the Evergreen Montane Forest of Catamayo-Alamor (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 895–1159 mm and the average annual temperature is 10.4–14.6 °C (Fick & Hijmans 2017). The male holotype was found near the road, calling from the branches of a bush, at about 3.5 m above ground. In this same location, choruses were heard from April to late July 2016; the males were calling usually from the vegetation nearby the road. An additional subadult was found during the day in a large terrrestrial bromeliad (Fig. 24A, B). Females and tadpoles are unknown. In this location, in Parque Nacional Yacuri, Gastrotheca yacuri is syntopic with G. turnerorum. Conservation status. We suggest that Gastrotheca yacuri should be assigned to the Data Deficient category according to guidelines of the IUCN (2001), given the inadecuacy of the current information to establish its status. Further searches and definition of its distribution, area of occupancy, habitat use, and biological information are required. Currently its known area of occurrence is less than 10 km 2, thus it could qualify for a Critically Endangered category; however, we can reasonably predict that if further surveys are conducted within the protected Yacuri National Park, its area will increase to more than 100 km 2. In any case, Lagunas Negras de Jimbura have suffered the synergistic effects of agriculture, cattle and sheep raising, fires, introduced species such as trout, pesticide use, and unregulated tourism activitites. Areas in the Salado de Jimbura that are out of the Yacuri National Park already have been heavily modified by human actions. Etymology. The specific name yacuri is a noun in apposition and refers to the Yacuri National Park, where this species has been found. According to Chamba-Troya (2017), Yacuri takes its name from the Quechua words Yacu meaning water and Quri that means gold. Ecuador's southernmost national park ranges from 2000 to 3700 meters in the Cordillera Oriental. This park of 43,090,60 hectares is unique by sheltering a series of 48 glacial lakes, and many endemic species of flora and fauna. It is a Ramsar site (The Convention on Wetlands) and part of the Podocarpus-El Cóndor Biosphere Reserve declared by UNESCO. Comments. Gastrotheca yacuri belongs to the subgenus Gastrotheca, and is reported as Species D in Duellman (2015: Fig 12.1).

Gastrotheca elicioi sp. nov. urn:lsid:zoobank.org:act: 3D26B088-B527-4362-94E8-647FF26C8419 Holotype. CJ 1402 (Fig. 12), an adult male (collected as tadpole in the field and captive raised), from Cajanuma, entrance to Parque Nacional Podocarpus in the Loja-Vilcabamba road, 2456 m (04�� 05' 25.51" S, 79�� 12' 09.97" W), Loja Province, Ecuador, one of a series obtained on 14 June 2011 by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa. Paratypes. (Total 18: 11 males, 3 females, 4 juveniles). Ecuador: Loja: CJ 413���4 (juvenile, male), 1398���401 (four males), 1941 (male) collected with the holotype; KU 142603 ���5 (female, subadults), from 5.5 km W Loja, 2330 m (04�� 00' 48.99" S, 79�� 13' 50.99" W), on 17 July 1971 by William E. Duellman and Linda Trueb; KU 142607 ���8 (subadult, female), 148549���51 (males, female), from 5.5 Km W Loja, 2330 m (04�� 00' 48.99" S, 79�� 13' 50.99" W), on 23 July 1971 by William E. Duellman and Linda Trueb; KU 202688 (male), from 5.2 km W Loja, 2310 m (03�� 58' 58.73" S, 79�� 16' 02.21" W), on 10 March 1984 by William E. Duellman; KU 217511 ���2 (males), from 6.8 km E Loja ca. Loja-Zamora line (03�� 59' 19" S, 79�� 09' 21.99" W), on 9 January 1990 by David Kizirian, John J. Wiens, and Luis A. Coloma. Referred specimens. Ecuador: Loja: QCAZ 22370 (male), from Zamora Huayco, Loja, 3018 m (04��0 5' 49.98" S, 79�� 10' 02.5" W), on 2 December 2002 by Diego Almeida-Reinoso; QCAZ 46319���20 (males), from Zamora Huayco, Loja, 3018 m (04�� 05' 49.98" S, 79�� 10' 02.5" W), on 5 December 2009 by Diego Almeida- Reinoso and Samael D. Padilla; CJ 1841���902 (62 juveniles), from Loja, Barrio La Palmera, Parroquia Sucre, on 22���25 April 2013 by Elicio E. Tapia. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (up to 67.5 mm SVL in female, 46.4���67.8 mm SVL in males, n = 13) with tibia length 42��50% SVL, slightly longer than foot; (2) interorbital distance larger than the width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth to slightly granular; (7) Fingers I and II about equal in length, width of discs about twice width of the digits proximal to discs; (8) fingers unwebbed; (9) webbing between external toes extending nearly to antepenultimate subarticular tubercle on Toe IV and proximal to penultimate subarticular tubercle on Toe V; (10) in life, dorsum green, tan, brown, or reddish-brown with or without dark paravertebral marks; (11) head markings consisting of a pale labial stripe on the posterior margin of the lip, a black canthal stripe and a dark triangular interorbital mark in some individuals; (12) a fragmented, narrow, pale dorsolateral stripe present; (13) flanks bronze-brown, tan, or green with or without small pale spots; groin and anterior and posterior surfaces of thighs lightly mottled with tinges of green or blue; (14) venter cream with small brown flecks or spots; (15) brood pouch single, dorsal. In comparison with similar species Gastrotheca elicioi is most like G. lojana, G. cuencana, G. litonedis, G. turnerorum, and G. pseustes in Ecuador and G. monticola in Peru. Gastrotheca elicioi differs from G. lojana, G. cuencana, and G. litonedis in color pattern as follows (compare these species in Fig. 10). Gastrotheca elicioi and G. litonedis have distinct dark canthal stripes, which are absent in G. lojana and inconspicuous or absent in G. cuencana. The groin and anterior and posterior surfaces of the thighs are slightly mottled in G. elicioi, whereas in G. lojana they are heavily mottled; these surface are translucent cream without marks in G. cuencana and usually brown in G. litonedis. If any marks are present on the dorsum in G. elicioi, they usually consist of a triangular interorbital mark, which is connected or not with two narrow and curved paravertebral marks. In contrast, in G. lojana the interorbital mark is a transverse bar usually connected with two broad paravertebral marks; in G. cuencana and G. litonedis an interorbital mark is absent. Gastrotheca lojana and G. cuencana have a conspicuous, elevated row of dorsolateral warts, whereas they are barely raised in G. elicioi and G. litonedis. Additionally, in G. elicioi the dark bars on the limbs, when present, are shorter, thinner and less defined than in G. lojana, whereas in G. cuencana and in some G. litonedis the bars are replaced by irregular blotches. Gastrotheca elicioi has a cream venter with dark flecks or marks, whereas the venter is uniform creamy white in G. cuencana and pale brownish gray in G. litonedis. Gastrotheca cuencana also differs from G. elicioi by having cream dorsal surfaces of the fingers, whereas they are brown, green, or tan in G. elicoi. Gastrotheca turnerorum and G. pseustes differ from G. elicioi (compare these species in Fig. 10) by having the skin on dorsum areolate and weakly areolate respectively, whereas in G. elicioi the skin on dorsum is finely granular. Also, they differ by lacking an interorbital mark, and bars on limbs, which can be present G. elicioi. Gastrotheca pseustes is sympatric with G. elicioi, from which it differs notably by having smaller digital discs (larger in G. elicioi) and lacking a pale dorsolateral and supracloacal stripes, which are fragmented in G. elicioi. Gastrotheca monticola is the sister species of G. elicioi and their minimum genetic divergence is 3.0% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca monticola differs from G. elicioi by having the axilla and groin, concealed surfaces of the thighs, and the dorsal surfaces of the feet green with well-defined black spots on a green or tan background, whereas in G. elicioi the groin and anterior and posterior surfaces of thighs are lightly mottled with tinges of green or blue. Finally, the advertisement call of G. elicioi is unique amongst the calls of Gastrotheca in southern Ecuador, in that the long, pulsed notes are produced after the short ones, whereas the other species tend to produce the long, pulsated notes before the short ones. Description of the holotype. An adult male (Fig. 12) collected as a tadpole and raised in the laboratory; body moderately robust; SVL 46.4 mm; head wider that long; snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis round in section; loreal region concave, lips rounded; top of head flat; interorbital distance 142% of width of upper eyelid; internarial area elevated; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye in a distance approximately one and a half the diameter of tympanum; tympanic annulus and membrane smooth; supratympanic fold moderately weak, extending from corner of eye to tympanum and on to insertion of forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing three teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 31% of SVL), unwebbed, with distinct narrow lateral fringes; discs much wider than digits, slightly truncated, width of disc on Finger III greater that diameter of tympanum; relative lengths of fingers I=IIColoration in life (See also under Comments). The dorsum is uniform brown becoming yellowish brown on the flanks. The dorsal surfaces of limbs have an olive-green tinge. A bronze labial stripe is present at the margin of the lip and continues to the insertion of the forelimb. A bronze inconspicuous and highly fragmented dorsolateral stripe is present; a black canthal stripe is present. The flanks are bronze-brown; the groin and anterior and posterior surfaces of the thighs are pale green with white and black spots. The venter is brownish gray with black and white marks; the gular region is gray dark. The iris is copper colored with a few black reticulations. Coloration in preservative. The dorsum is gray without markings. A narrow pale, fragmented dorsolateral stripe is present. The dorsal surfaces of the limbs are tan with irregular gray blotches. A short pale labial stripe extends from the margin of the lip to the level of the insertion of forelimb; a dark canthal stripe is present. The flanks are dark gray anteriorly and pale gray posteriorly; the groin and anterior and posterior surfaces of the thighs are pale gray with black marks. The venter is cream with dark gray spots; the gular region is gray. Measurements (in mm). SVL: 46.4, TIBL: 19.8, FL: 20.0, HL: 14.1, HW: 16.2, IOD: 4.9, EW: 4.6 IND: 3.3, ED: 5.6, EN: 4.1, TD: 2.2, FFL: 7.9, TFL: 14.6, TFD: 2.6. Variation. Morphometric variation of one female and 13 males is summarized in Table 2. The female is larger than most males (67.5 mm; 56.0�� 8.1 mm). The skin on the dorsum is finely granular. The tympanic annulus usually is smooth but is slightly granular in some individuals. Each dentigerous vomerine process has 6���8 teeth (6.3��1.5, n = 3). Color variation in preservative. Preserved specimens have a bluish gray, pale gray, or brownish gray dorsum. Interorbital and paravertebral marks usually are present but the paravertebral marks may be absent. The dark gray interorbital mark, when present, is triangular in most specimens, but it is divided into two blotches in some specimens. A dark canthal stripe is present in all specimens. A short, pale labial stripe is inconspicuous in most specimens. The flanks are gray, darkest anteriorly, with or without white and dark flecks. The groin and anterior surfaces of thighs are pale gray with slight mottling in some; the posterior surfaces of the thighs are pale gray with black flecks and marks usually only distally. In some specimens, the dorsal surfaces of arms and limbs present short dark gray bars. The venter is white with evenly, or densely distributed dark marks. Color variation in life. (Figs. 10H, 10I, 13���14). In living individuals the dorsum is uniform green (CJ 1843), reddish-brown (CJ 1400), or tan (CJ 1842), with or without contrasting dark gray paravertebral marks. When present, these paravertebral marks usually are narrow, curved, and fragmented; these marks are adherent at the scapular level and may be connected with the interorbital mark. A short cream labial stripe is present on the margin of the lip; the stripe continues posteriorly to the insertion of the forelimb. A dark brown canthal stripe is always present, but it is inconspicuous in some individuals with a dark ground color. The tympanum is brown, tan, or olive green. The iris is usually copper almost without black reticulations. A bronze highly fragmented dorsolateral stripe is present in most individuals. The flanks are brown, bronze-brown, or green. The groin has bluish coloration with slight mottling. The posterior surfaces of the thighs usually are pale green with black flecks and some specimens show black fringes. When present, supracloacal and heel stripes are cream and fragmented. The ventral surfaces usually are cream with dark marks, but some specimens lack dark marks. The gular surface varies from brownish gray to white with black spots. Duellman (1974) provided detailed descriptions of coloration in life of four adult males (KU 148549 ���51 and 142603, under the name Gastrotheca lojana). Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36 (CJ 4311), from a series of 50 tadpoles (CJ 4311) obtained from a pond at Puntzar�� Alto, near Loja city, 2311 m, Province of Loja, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016. Total length 51.4; body length 19.4 (38% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 11.9, and height at same position 9.9; head width at level of eyes 11.1. Lateral line system present but barely visible, supraorbital and not evident at level of snout, infraorbital line present at level of the eye, touching the inferior portion of the orbit, and making contact with supraorbital line immediately behind the eye. Inferior oral line visible at the eye level, where it contacts angular line, which descends from eye level. Supraorbital line represented for scattered stiches, which does not make contact with the orbit. Posorbital line forming a circle of stitches, just behind the eye. Anterior oral line and loreal lines not visible; dorsal body and middle bodylines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 3.5; internarial distance 2.9. Eye directed dorsally; eye length 2, eye width 1.8; interorbital distance 5. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 13.0; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; spiracle length 2.9, tube transverse width 2.9. Vent tube dextral, opening oriented posteriorly, tube length 3.4, vent tube transverse width 3. Tail length 32.4; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 4.4, width 3.4; caudal fins well developed and proportional, arising abruptly near tail-body junction and forming a notorious hump, which makes and arc in the body plane. Dorsal fin height 4.18, ventral fin height 3.7; maximum height of tail 11.9; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 4.9. It is surrounded by an uniserial row of marginal papillae, interrupted medially on upper lip; lower lip papillae alternate in orientation, giving appearance of two rows. Upper lip with 23 papillae on right side and 20 papillae on left side; lower lip bearing 52 marginal papillae; upper jaw sheath medium-sized, forming a smooth arch and finely serrated, transverse width 2.6 (53% of oral disc width) height 0.4; lower jaw sheath V- shaped, open and finely serrated, width 4.1, height 1.3. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 4.15, A2: 4.0, P1 right row 1.75, P1 left row 1.75, P1 gap 0.1, P2: 3.75, P3: 3.70. (Fig. 6D). Color in life. Based on a specimen (CJ 4312a) in Stage 37 from a series (CJ 4312) obtained at Loja (neigborhood Puntzar�� Grande), Loja Province, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016 (Fig. 5D). In dorsal and ventral views, body dark brown. Snout and flanks dark brown; guts gray; red gills visible through the throat. Venter gray. Caudal musculature reddish brown with brown stippling, distal portion lacking pigments; dorsal and ventral fins gray with minute cream stippling. Vent tube translucent. Iris reddish-gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 31���40 (CJ 4306, 4310���13) are shown in Table 6. Total length varies between 15.8 (Stage 31) and 73.2 (Stage 39) and tail length proportion varies from 57% to 70% until Stage 38. Number of marginal papillae varies among specimens and Gosner stages; variation in number of ventral papillae at lower lip is high (43���64). Based on specimens CJ 1950���2 from a series obtained at Loja (neigborhoods La Palmera and Puntzar�� Grande), Loja Province, Ecuador, by Elicio E. Tapia on 22���25 April 2013 (Fig. 15). We documented changes in coloration during ontogenetic development of one, mostly brown individual (CJ 1950) (Figs. 15 A���B). At Stage 40, the dorsum and flanks were cream and brown with a diffuse pattern of brown-gray paravertebral marks and a dark gray stripe bordering the canthus and body dorsolaterally, extending to about level of midbody, bordered dorsally by a diffuse cream area. The caudal musculature is pink proximally, with a gray suffusion distally. The caudal fins are pale gray. The iris is pale red. By Stage 46, the markings on the dorsum of body and limbs are diffuse gray and green on a dark brown background; the fingers and toes are yellowish cream dorsally. The flanks have a wide, brown-gray band. There are well-defined creamy white dorsolateral and labial stripes and supracloacal white marks. Color variation in six additional metamorphs (CJ 1951���2, CJ 4313) from Loja and Loja���Abra de Zamora, in Stage 46 is depicted in Figure 16. They vary from plain dark brown to plain green, and have a complete supracloacal stripe. ������.continued on the next page ������.continued on the next page Comparisons. Tadpoles of Gastrotheca elicioi may occur in sympatry with those of G. lojana, G. psychrophila, G. pseustes, and G. turnerorum in the Loja-Abra de Zamora region. Gastrotheca elicioi differs from all of them by having a dorsal gray-pigmented fin that abruptly arises from the body, whereas is nearly translucent and arises gradually in the other species (compare in Fig 5). The tadpole described as G. psychrophila by Duellman (2015) is G. elicioi. It has a similar dorsal fin and fit well our description of G. elicioi tadpoles. Nonetheless, until tadpoles of G. psychrophila are described and unequivocally assigned to its species, some doubts will remain. Vocalization. Seven individuals of Gastrotheca elicioi were recorded from three locations in Loja Province (one individual from the old Loja-Catamayo road, three from Loja, Parque Universitario, Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 34-48, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Duellman, W. E. (1974) A systematic review of the marsupial frogs (Hylidae: Gastrotheca) of the Andes Ecuador. Occasional Papers of the Museum of Natural History, The University of Kansas, Lawrence, 22, 1 - 27.","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp.","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","Pollet, D. (2003) A la Sombra de Otonga. Serie de divulgacion 4. Centro de Biodiversidad y Ambiente, Pontificia Universidad Catolica del Ecuador, Quito, 159 pp.","Paez-Vacas, M. I., Coloma, L. A. & Santos, J. C. (2010) Systematics of the Hyloxalus bocagei complex (Anura: Dendrobatidae), description of two new cryptic species, and recognition of H. maculosus. Zootaxa, 2711 (1), 1 - 75. https: // doi. org / 10.11646 / zootaxa. 2711.1.1"]}

Gastrotheca cuencana sp. nov. urn:lsid:zoobank.org:act: 82F87E67-2940-424C-AC03- BC751224 D5E4 Holotype. CJ 1391 (Fig. 2), an adult male, from the city of Cuenca, 2579 m (02° 53' 57.91" S, 79° 01' 52.79" W), Azuay Province, Ecuador, one of a series obtained on 8 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara and Henry Grefa. Paratypes. (Total 48: 10 adult males, 21 adult females, 17 juveniles). Ecuador: Azuay: CJ 1392–7, collected with the holotype; KU 120676 (female), 120683 –4, 120690, 120695 (4 juveniles), 120699 (male), 120705 (male), 120709 –10 (males), 120713 (female), 120718 –9 (female, male), 120721 (female), 120722 (male) from Cuenca, 2600 m, (02° 53' 57.91" S, 79° 01' 52.79" W), on 19 June 1968 by John D. Lynch; KU 138616 (female), 138619– 21 (male, females) from 4 km E: Cuenca 2540 m (02° 52' 59.88" S, 78° 22' 0.11" W), on 10 June 1970 by Thomas H. Fritts; KU 141572 (female) from 2.1 km S Cutchil, 2720 m (03° 06' 00"S, 78° 47' 59.99"W), on 16 May 1971 by Richard R. Montanucci; KU 141579 (female) from 3.5 km S Cutchil, 2785 m (03° 04' 59.87" S, 78° 47' 59.99" W), on 14 May 1971 by Richard R. Montanucci; KU 141582 (juvenile) from 8 km NW Cuenca, 2803 m (02° 51' 47.88" S, 78° 58' 59.88" W), on 16 May 1971 by Richard R. Montanucci; KU 141583 (female) from 8.8 km NW Cuenca, 3820 m (02° 52' 00.12" S, 79° 04' 00.12" W), on 15 May 1971 by Richard R. Montanucci; KU 203441 (male) from Laguna de Zurucuchu, 16 km NW Cuenca, 3200 m (02° 50' 23.99" S, 79° 07' 47.99" W), on 5 March 1984 by William E. Duellman; KU 129779 –82 (4 females), 129783–91 (9 juveniles), 129793–94 (juveniles), 129795 (female), 129796 (juvenile) from Río Matadero, 12 km E Cuenca (02° 52' 59.88"S, 78° 58' 00.12" W), on 17 November 1968 by Craig E. Nelson. Cañar: KU 141571 (male) and KU 141573 (female) from Biblián, 2620 (02° 42' 00" S, 78° 52' 00.12" W), on 17 and 23 May 1971, respectively, by Richard R. Montanucci; KU 142620 –24 (4 females, male), from Biblián, 2620 m (02° 42' 00" S, 78° 52' 00.12" W), on 25 July 1971 by William E. Duellman; KU 147113 (female) from Biblián, 2620 m (02° 42' 00" S, 78° 52' 00.12" W), on 15 January 1972 by John E. Simmons. Referred specimens. (Total 36, 28 adult males, 8 adult females). Ecuador: Azuay: QCAZ 1239 (male), from Sigsig, 2480 m (03° 03' 08.4" S, 78° 48' 10.56" W), on 24 July 1989 by Luis A. Coloma and Luis E. López; QCAZ 26309 (female), from Cumbe, 2740 m (03° 05' 55.64" S, 79° 00' 28.73" W), on 8 August 2003 by Patricio Vargas; QCAZ 26348–50 (males), 26353 (female), 26357–64 (8 males), from Cumbe, 2740 m (03° 05' 55.64" S, 79° 00' 28.73" W), on 9 August 2003 by Luis A. Coloma, Ítalo G. Tapia, Andrés Merino Viteri, Erik Wild and Patricio Vargas Mena; QCAZ 26354 (female), from Cumbe, 2740 m (03° 05' 55.64" S, 79° 00' 28.73" W), on 9 August 2003 by Ítalo G. Tapia; QCAZ 26357–8 (males), from Cumbe, 2740 m (03° 05' 55.64" S, 79° 00' 28.73" W), on 9 August 2003 by Andrés Merino Viteri; QCAZ 31477 (male), from Sigsig, 2424 m (03° 03' 08.4" S, 78° 48' 10.56" W), on 8 March 2006 by Ítalo G. Tapia; QCAZ 31509, 31511 (males), from Sigsig, 2424 m (03° 03' 8.4" S, 78° 48' 10.56" W), on 9 March 2006 by Ítalo G. Tapia and Giovanni Onore; QCAZ 34131 (female), from Cuenca, 2579 m (02° 53' 57.91" S, 79° 01' 52.79" W), on 27 February 1979 by Fernando I. Ortiz; QCAZ 37375–6, 37379–81, QCAZ 37386 (6 males), from unknown locality, on 31 May 2007 by Zoológico Amaru; QCAZ 38232 (male), from Carmen del Guzho, 2666 m (02° 56' 00" S, 79° 03' 00" W), on 16 June 2007 by Hari González Maldonado; QCAZ 39384–5 (female, male), from Tarqui, Patapamba, 2600 m (03° 00' 38.16" S, 79° 01' 53.43" W), on 21 June 2007 by Ernesto Arbeláez Ortiz; QCAZ 42720–1 (males) from Cuenca, 2579 m (02° 53' 57.91" S, 79° 01' 52.79" W), on 3 October 2007 by Sofía Carvajal-Endara and Flavio Jaramillo; QCAZ 47106 (male), from Tarqui, 2741 m (03° 00' 57.17" S, 79° 02' 40.13" W), on 21 June 2007 by Ernesto Arbeláez Ortiz. Cañar: QCAZ 42826, 42835, 42841 (females), from Papaloma de la Nube, 3011 m (02° 40' 21.54" S, 78° 54' 21.28" W), on 12–14 August 2008 by Sofía Carvajal-Endara. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (49.0– 61.6 mm SVL in females, n = 28; 45.2–53.7 mm SVL in males, n = 38), with tibia length 39¯50% SVL, slightly shorter than length of foot; (2) interorbital distance slightly larger than width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus weakly granular to smooth; (7) Fingers I and II about equal in length, width of discs greater than digits; (8) fingers unwebbed; (9) webbing between external toes extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum green, tan, brown, or reddish-brown with or without dark paravertebral marks; (11) head markings consisting of pale labial stripe and, in some individuals, an inconspicuous canthal stripe; (12) pale white or cream dorsolateral stripe present; (13) flanks tan, brown, or green with cream flecks or spots; groin and anterior and posterior surfaces of thighs translucent cream without marks and with a faint pale blue tinge in some; (14) venter uniform creamy white; (15) in females, brood pouch single, dorsal. In comparison with similar species in Ecuador, G. cuencana is most like G. litonedis, G. lojana, G. elicioi, and G. riobambae. It differs from these species by having a uniform creamy white venter; whereas the venter is pale gray in G. litonedis, and cream with dark flecks, spots or mottling in G. lojana, G. elicioi, and G. riobambae. In G. cuencana the dorsal surfaces of fingers are cream, whereas in the other species dorsal surfaces of fingers are of the same color as the rest of the body (tan, brown, or green). Gastrotheca cuencana also differs from G. litonedis, G. elicioi, and G. riobambae by lacking a dark canthal stripe. Moreover, in G. cuencana the groin and anterior surfaces of thighs are mostly translucent cream without marks; whereas, in G. litonedis the groin and anterior surface of thighs are usually bronze-brown or tan with cream or darks flecks and spots. In G. lojana and G. riobambae the groin and anterior surfaces of the thighs are usually cream with black mottling; in G. elicioi the groin and anterior surfaces of thighs usually have dark and cream flecks and spots. In profile, the snout in G. cuencana is rounded above and inclined anteroventrally, whereas the snout in G. elicioi, G. lojana, and G. litonedis is bluntly rounded. Furthermore, G. cuencana has a conspicuous, elevated row of dorsolateral warts whereas these are barely raised in G. litonedi s; a narrow, cream supracloacal stripe is present in G. litonedis, G. lojana, and G. riobambae; whereas the stripe is fragmented in G. elicioi and absent in most G. cuencana. Gastrotheca lojana and G. riobambae also differ from G. cuencana and G. litonedis by having a complex call, whereas the call in the latter two species is a simple call. Additionally, the call of G. cuencana has a lower note rate, shorter note duration, longer inter-note interval and a lower dominant and 90% bandwidth frequency compared with G. litonedis. Finally, G. cuencana differs from its sister species (G. litonedis) by having a genetic distance of 1.2% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca cuencana occurs in syntopy with G. pseustes (sensu lato) throughout most of its range. Gastrotheca pseustes differs from G. cuencana by having coarsely granular skin on the dorsum, in contrast to the usually finely granular dorsum in G. cuencana. In G. cuencana the venter is uniform creamy white, whereas it is cream or white with dark flecks or spots in G. pseustes; also the call of G. cuencana is simple, whereas in G. pseustes the call is complex. Description of holotype. An adult male (Fig. 2); body moderately robust; SVL 50.6 mm; head wider that long; snout rounded in dorsal view, rounded and inclined anteroventrally in profile; canthus rostralis round in section; loreal region concave, lips rounded; top of head flat; interorbital distance 104% of width of upper eyelid; internarial area slightly elevated; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from eye by distance about equal to diameter of tympanum; tympanic annulus and membrane weakly granular; supratympanic fold moderately weak, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing four teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 35% of SVL), unwebbed, with distinct narrow lateral fringes; discs moderately large and rounded, width of disc of Finger III greater that diameter of tympanum; relative lengths of fingers I=II Skin on dorsum finely granular; skin on flanks coarsely granular; skin on throat, venter surfaces of thighs, and arms granular; skin on belly areolate; skin on venter surface of shanks smooth; numerous small tubercles lateral to cloacal opening. Vocal sac single, median, subgular. Vocal slits present at posterior lingual margins of mandibles. Tongue broad, suboval, not notched posteriorly, fully attached to mouth floor. Coloration in life (Fig. 2). The dorsum is yellowish brown with two darker, black delineated, continuous, paravertebral markings. These markings extend to the sacrum where they bifurcate and become two rows of irregular blotches at each side of the vertebral axis. The dorsal surfaces of the limbs are yellowish brown with irregular gray blotches. Cream dorsolateral and labial stripes are present; the latter continues to the insertion of forelimb; a faint brown canthal stripe is present. The flanks are bronze-brown with small black spots. The venter, groin, and anterior surfaces of the thighs are uniform creamy white; the posterior surfaces of the thighs have a faint bluish coloration. The dorsal surfaces of the fingers are cream. The gular region is creamy gray laterally. The iris is reddish bronze with black reticulations. Coloration in preservative. The dorsum is tan with two well-defined, gray paravertebral markings, which bifurcate at the level of sacrum to become two rows of irregular blotches on each side of the vertebral axis. A white dorsolateral stripe is present. The dorsal surfaces of limbs are tan with irregular gray blotches. A cream labial stripe continues to the insertion of the forelimb; a faint canthal is present. The flanks are gray; the venter, groin, and anterior surfaces of thighs are uniform creamy white; the posterior surfaces of thighs have a faint bluish coloration. The gular region is creamy gray. Measurements (in mm). SVL: 50.4, TIBL: 21.9, FL: 24.9, HL: 16.9, HW: 19.1, IOD: 5.2, EW: 5.0 IND: 3.3, ED: 5.6, EN: 4.1, TD: 2.8, FFL: 10.0, TFL: 17.5, TFD: 3.4. Variation. Morphometric variation of 28 females and 38 males is summarized in Table 3. Females are larger than males (54.2± 2.9 mm; 49.6± 2.4 mm). The skin on the dorsum is finely granular in most, but it varies in some specimens from weakly areolate to coarsely granular. All adults have a supratympanic fold that usually extends from the posterior part of the tympanum to point above the insertion of forelimb. The tympanic annulus is usually smooth, but some individuals (e.g., CJ 1395, 1940) have a slightly granular annulus (Fig. 3A, C). Each dentigerous vomerine process has 2–7 teeth (4.4±1.1, n = 39). Color variation in preservative. Preserved specimens have bluish gray dorsum (tan and reddish-brown in CJ 1397). Faint paravertebral marks are present. A canthal stripe is absent in most specimens; the flanks and posterior surfaces of the thighs are dark gray with white spots evenly distributed in some specimens. In all specimens, a prominent white labial stripe extends from the posterior margin of the lip to the insertion of the forelimb. A narrow white dorsolateral stripe is evident in most specimens. The groin and the anterior surfaces of the thighs are uniform creamy gray. The ventral surfaces are uniform creamy white. The posterior surfaces of thighs usually are creamy gray, but they are pale blue in some specimens. Some males have yellowish brown nuptial pads on the medial surface of the thumb. Color variation in life. (Figs. 3–4). The dorsum is uniform green (CJ 1393 ), reddish brown (CJ 1397), or tan (CJ 1392) with (CJ 1391) or without (CJ 1392) darker paravertebral marks. When present, the paravertebral markings usually are well defined; at the level of the sacrum they become two rows of irregular blotches at each side of the vertebral axis. The ventral surfaces are uniform creamy white. A cream labial stripe is extends from the posterior margin of the lip to the insertion of the forelimb; a dark brown canthal stripe is absent in most individuals. The tympanum is brown, tan, or olive green. The iris is reddish bronze with black reticulations. A creamy white dorsolateral stripe is evident in most individuals. The flanks are cream, bronze brown, or green. The groin and anterior surfaces of the thighs are translucent cream without dark markings (Fig. 3). The posterior surfaces of the thighs are translucent, cream, or pale blue or green. When present, supracloacal and heel stripes are cream or tan. The ventral surfaces of the shanks have a faint tinge of pale blue. The gular surface varies from white to creamy gray. ……continued on the next page ……continued on the next page Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 38 (CJ 1945a), from a series of eight tadpoles (CJ 1945) obtained from a pond at south Cuenca, 2579 m, Azuay Province, Ecuador, by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa on 8 June 2011 (Fig. 5A). Total length 44.6; body length 17.3 (39% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 11.5, and height at same position 9.3, head width at level of eyes 8.5. Lateral line system present but barely visible, supraorbital and infraorbital lines both rising at level of tip of snout, extending parallel to the eye and making contact immediately behind eye; angular line descending vertically just porterior of eye to throat, it dorsally contacts with post infraorbital line; anterior oral line descending vertically from oral disc level and behind nares level to throat; it makes a curve that parallels infraorbital line, forming a circuit continuous with angular and loreal lines; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 2.3; internarial distance 2.4. Eye directed dorsally; eye length 1.8, eye width 1.8; interorbital distance 4.3. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 11.0; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; tube length 3.1, tube transverse width 2.0. Vent tube dextral, opening oriented posteriorly, tube length 1.7, tube transverse width 1.8. Tail length 27.2; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 3.7, width 3,2; caudal fins well developed and proportional, arising near tail-body junction, forming a low hump; dorsal fin height 3.0, ventral fin height 2.8; maximum height of tail 9.3; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 4.7. It is surrounded by a uniserial row of marginal papillae, interrupted medially on upper lip; lower lip papillae alternate in orientation, giving appearance of two rows; upper lip with 29 papillae on right side and 27 papillae on left side; lower lip bearing 62 marginal papillae; upper jaw sheath medium-sized, forming a smooth arch and finely serrated, height 0.36, transverse width 2.8 (60% of width of oral width); lower jaw sheath V- shaped, open and finely serrated, width 1.92, height 0.7. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 3.6, A2: 3.5, P1 right row 1.5, P1 left row 1.6, P1 gap 0.2, P2: 3.4, P3: 3.0. (Fig. 6B). Color in preservative. Dorsum dull gray with darker areas on flanks, above eye, and on throat; body contour and snout translucent. Caudal musculature and fins with scattered cream dots, with higher suffusion near tail base, otherwise translucent. Venter pale, translucent in belly region, guts exposed; eyes lavender gray; oral apparatus translucent. Color in life. In dorsal view, body olive-cream; snout and flanks paler olive-cream. In ventral view, guts dark gray in belly area; reddish gills visible throughout throat. Caudal musculature covered by a nearly continuous line of cream marks, proximal half of musculature reddish cream with myomeres and medial line well defined, distal half translucent; dorsal and ventral fins translucent, with scattered cream dots, heavily suffused near tail-body junction. Spiracle, oral apparatus, and legs olive-cream. Iris gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 37–39 (CJ 1945) are shown in Table 4. Total length varies

We review the systematics of the species of Gastrotheca (Anura: Hemiphractidae) in the highlands of the southern Ecuadorian Andes. We analyzed phylogenetic, morphological, ecological, and acoustic data from populations in the region. We provide an updated phylogenetic hypothesis inferred from a database that contains 42 species of Gastrotheca, and sequences from mitochondrial (12S, 16S, and ND1) and nuclear (POMC and RAG-1) genes. We describe four new endemic biphasic species for Ecuador (Gastrotheca cuencana, G. elicioi, G. turnerorum, and G. yacuri). In addition, we redefine G. lojana, recently resurrected from the synonymy of G. monticola, and redescribe G. litonedis. Furthermore, we describe the tadpoles and ontogenetic color changes in six species (G. cuencana, G. elicioi, G. litonedis, G. lojana, G. pseustes, and G. turnerorum). Also, we describe the calls of seven species (G. cuencana, G. elicioi, G. litonedis, G. lojana, G. pseustes, G. testudinea, and G. yacuri). The phylogenetic relationships estimated here are congruent with previous phylogenetic hypotheses for the group, except for the placement of G. galeata, G. plumbea, G. orophylax, G. nicefori, and G. griswoldi. Because most sister species in Gastrotheca are allopatric, the evolutionarly scenarios that likely have intervened in shaping the diversity are the uplift of the Andes and the formation of cross-Andean river valleys, which probably promoted conditions that acted as dispersal barriers that led to speciation. Many species of Gastrotheca (44%) are highly threatened. Most monophasic species have not been seen since the late 1980s, whereas the highland biphasic species have survived the catastrophic events that led to the disappearance of many other sympatric anurans. Research and conservation actions are urgently needed for all surviving species treated herein, five of which are either Critically Endangered or Endangered because of their restricted distributions (10–1600 km2), habitat destruction and fragmentation, climate change, and pathogens. Two of the new species, G. cuencana, and G. elicioi, mostly are known from the cities of Cuenca and Loja, respectively, where urbanization threatens their survival. In order to call attention to conservation issues, we name one new species to refer to the Yacuri National Park, and the other three in honor of the people from Cuenca, the Turner family, and Elicio E. Tapia, whose actions are helping to halt amphibian extinctions.

Gastrotheca turnerorum sp. nov. urn:lsid:zoobank.org:act: B72F71A3-671F-4F89-91D5-2410C1D0B6DF Holotype. CJ 393 (Fig. 18), an adult female (collected as tadpole and raised in captivity), from Laguna Negra de Jimbura, Parque Nacional Yacuri, Amaluza, 3406 m (04° 42' 44.24" S, 79° 25' 42.38" W), Loja Province, Ecuador, one of a series collected on 12 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara and Henry Grefa. Paratypes. (Total 9: 6 males, 2 females, 1 juvenile). Ecuador: Loja: CJ 394–5 (female, male), 415–7 (males, juvenile), 1386 (female), and KU 335390 (male), collected with the holotype. CJ 7823 (male, collected as tadpole and captive raised), from Parque Nacional Yacuri, Amaluza, 3331 m (04° 43' 25.22" S, 79° 26' 15.47" W), Loja Province, Ecuador, one of a series collected on 7 July 2016 by Dan Cogǎlniceanu, Diana Székely and Paul Székely. Zamora Chinchipe: MUTPL 221, male from Reserva Tapichalaca, 3073 m (04° 28' 55.58" S, 79° 09' 29.76" W), collected on 9 December 2014 by Diego Armijos-Ojeda. Referred specimens. Ecuador: Zamora Chinchipe: QCAZ 9575 (captive raised tadpoles and juveniles) from Parque Nacional Podocarpus, Lagunas del Compadre, 3205 m (04° 10' 29.24" S, 79° 06' 59.54" W), on 1 December 1994 by Jenny Rudston; QCAZ 47299 (tadpole), QCAZ (sc 29154) (female) from Parque Nacional Podocarpus, Lagunas del Compadre, 3205 m (04° 10' 29.24" S, 79° 06' 59.54" W), on 20 October 2009 by Elicio E. Tapia. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (54.0– 58.2 mm SVL in females, n = 3; 50.0– 55.6 mm SVL in males, n = 4), with tibia length 37¯42% SVL, shorter than foot; (2) interorbital distance slightly greater than width of upper eyelid; (3) skin on dorsum areolate, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I slightly shorter than Finger II, width of discs much wider than digits; (8) fingers unwebbed; (9) foot webbing between external toes extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum uniform green without dark paravertebral marks; (11) head markings consisting of pale labial and canthal stripes formed by a series of small bronze dotes; (12) dorsolateral stripe present, also consisting in a series of small bronze dotes; (13) flanks brown and groin green both with or without a few cream spots; anterior surfaces of thighs dark brown, posterior surfaces of thighs bluish brown with numerous cream tubercles proximal to the vent; (14) venter dark brown with uniformly distributed cream spots; gular region dark brown; (15) brood pouch single, dorsal. Gastrotheca turnerorum most closely resembles three other species in southern Ecuador, G. pseustes, G. elicioi, and G. litonedis. Gastrotheca turnerorum differs from all of them by having a different skin texture on the dorsum, and a distinctive color pattern (compare Figs. 10J, 19–20 vs 10A–I, K, L). The dorsal skin is areolate in G. turnerorum, whereas it is weakly areolate or smooth in G. pseustes, finely granular in G. elicioi, and smooth in G. litonedis. In G. turnerorum the venter is dark brown with uniformly distributed cream spots, whereas it is cream with dark marks in G. pseustes and G. elicioi and pale brown-gray in G. litonedis. The labial stripe consists of a series of small bronze dots in G. turnerorum, and mostly uniform cream in G. pseustes, G. elicioi, and G. litonedis. Gastrotheca pseustes and G. elicioi have a dark brown canthal stripe, whereas G. turnerorum has a canthal stripe formed by a series of small bronze dots. Gastrotheca turnerorum is the sister species of G. aguaruna + G. yacuri, with a genetic distance of at least 2.85 % (in a DNA dataset of 438 bp, 16S gene). The Peruvian G. aguaruna differs from G. turnerorum by having a weakly granular to smooth dorsal skin (areolate in G. turnerorum), Fingers I and II equal in length (Finger I shorter in G. turnerorum), and a cream venter with dark fleck or spots (venter dark brown with uniformly distributed cream spots in G. turnerorum). Description of the holotype. An adult female (Fig. 18) that was collected as a tadpole and raised in the laboratory; body moderately robust; SVL 54.0 mm; head wider that long; snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis round in section; loreal region concave; lips rounded; top of head flat; interorbital distance 89% of width of upper eyelid; internarial area flat; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye by distance about equal to diameter of tympanum; tympanic annulus barely evident;, supratympanic fold moderately weak, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing five teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 34% of SVL); fingers unwebbed; discs large and rounded, width of disc of Finger III greater that diameter of tympanum; relative lengths of fingers I Skin on dorsum weakly areolate; skin on flanks coarsely areolate; skin on throat, venter surfaces of thighs, and arms heavily areolate; skin on belly areolate; skin on venter surface of shanks smooth; numerous white tubercles ventrolateral to cloacal opening. Tongue broad, suboval, not notched posteriorly, fully attached to mouth floor. Pouch opening V- shaped with anterior border at level of posterior edge of sacrum. Coloration in life. The dorsal surfaces of the head, body, and limbs are green; the inner and outer surfaces of the forearm, thighs, and shanks are brown (Fig. 18). Head markings are canthal and labial stripes that consist of a series of small bronze dots. The labial stripe continues to the insertion of the forelimb. The dorsolateral stripe also consists of a series of small bronze dots. The flanks are bronze-brown with some small white stippling; the groin is green. The anterior surfaces of the thighs are dark brown with some cream stippling; the posterior surfaces of the thighs are dark brown with cream tubercles lateral and below the vent. The ventral surfaces are dark brown with uniformly distributed cream spots. The iris is yellow-cream with brown reticulations and a brown horizontal bar across the eye. Coloration in preservative. The dorsum is uniform bluish-gray. The pale labial, canthal, and dorsolateral stripes formed by a series of small bluish gray dots. The labial stripe continues to the insertion of the forelimb. The flanks, groin, and anterior surfaces of thighs are uniform dull gray; the posterior surfaces of thighs are dark gray with numerous white tubercles lateral to, and below, the cloacal opening. The ventral surfaces are uniform dark bluish-gray. Measurements (in mm). SVL: 54.0, TIBL: 20.5, FL: 24.7, HL: 17.8, HW: 20.5, IOD: 4.9, EW: 5.6 IND: 3.4, ED: 5.6, EN: 4.3, TD: 2.3, FFL: 9.9, TFL: 18.5, TFD: 3.4. Variation. Morphometric variation of three females and three males is summarized in Table 3. Females are larger than males (55.9± 2.1 mm; 52.7.± 2.5 mm). The skin on the dorsum varies from weakly to coarsely areolate. All adults have a supratympanic fold, which usually extends from the posterior edge of the tympanum to a point above the insertion of the forelimb. Color variation in preservative. All preserved specimens have a uniformly bluish gray dorsum; labial, canthal, and dorsolateral stripes are formed by a series of small brown dots. The dorsal surfaces of hands and feet also have brown dots. The flanks, groin, and anterior surfaces of the thighs are uniform dull gray, but in some specimens the flanks are covered by small dark gray dots. The posterior surfaces of the thighs are dark gray with numerous white tubercles lateral to, and below the cloacal opening; a short, fragmented supracloacal stripe is present in some individuals. The ventral surfaces are uniform dark bluish-gray. Color variation in life. (Figs. 10J, 18–19). The dorsum is uniform green (e.g., CJ 1386) or brown (e.g., CJ 7823). In CJ 1386, the dorsal surfaces of limbs are green, whereas the inner and outer surfaces of the forearms, thighs, and shanks are brown. Dorsal surfaces of hands and feet have a pattern of brown dots over a lighter background. Head markings are like those of the holotype; a dorsolateral stripe is present in all specimens. The flanks are bronze-brown with or without white stippling; the axillae and groin are green, with a bluish suffusion in some specimens. The anterior surfaces of thighs are dark brown with or without cream stippling and, and the posterior surfaces of thighs are dark brown with cream tubercles lateral to, and below, the vent. The ventral surfaces are dark brown with uniformly distributed cream dots, but some specimens have larger cream spots, thereby resulting in a paler venter. Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 39, from a series (CJ 1959) obtained from a pond at Laguna Negra de Jimbura, Parque Nacional Yacuri, 3406 m, Loja Province, Ecuador, by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa on 12 June 2011 (Fig. 5F). Total length 61.3, body length 23.5 (38% of total length). Body robust, ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile; body width at level of spiracle 14.7, and height at same position 11.9; head width at level of eyes 12.9. Lateral-line system not evident. Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 3.7. Eyes positioned and directed dorsally, eye length 2.4, eye width 2.2. Spiracle sinistral, located at midbody level; spiracular opening oriented posteriorly; distance from tip of snout to spiracle opening 15.9; spiracle end rounded, not free, attached to body wall, inner wall of spiracle not evident; tube length 3.4, tube transverse width 1.4. Vent tube dextral, the opening oriented posteriorly, tube length 4.0, tube transverse width 2.6. Tail length 38.8, caudal musculature robust in the two-thirds proximal to body, narrowing gradually until tail terminus; tail muscle height 5.4, tail muscle width 4.8; caudal fins thin at proximal half of tail, getting higher on distal half and raising near tail–body junction, dorsal fin height 3.5, ventral fin height 3.7; maximum height of tail 12.7; tail tip rounded, tail musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 5.6. It is surrounded by an uniserial row of marginal papillae, interrupted medially in upper lip; lower lip papillae alternating in and out, giving the appearance of two series; upper lip with 18 papillae on right side and 17 papillae on left side; lower lip with 42 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, height 0.4, transverse width 3.3 (59% of oral disc width); lower jaw sheath V- shaped, open and finely serrated, width 2.4, height 0.7. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.7, A2: 4.0, P1 right row 1.9, P1 left row 2.1, P1 gap 0.1, P2: 3.6, P3: 3.5. (Fig. 6E). Color in preservative. Dorsum dull gray, with darker areas on flanks, above the eyes and on the throat; body contour and snout translucent. Caudal musculature and fins with scattered, white spots, densely arranged near tailbody junction; fins otherwise translucent. Venter dark gray; eyes lavender gray, oral apparatus translucent. Color in life. (CJ 1957, Stage 36). Fig. 5F. In dorsal and lateral views, body tan, speckled with black; areas around the snout are lighter. Venter cream with black markings, semi-translucent; guts visible as a darker area; throat translucent with small cream flecks; gills evident as a red hue. Caudal musculature reddish-pink in proximal half, decreasing its intensity towards distal half; myomeres and nerves barely visible; caudal musculature and fins with cream marks, clustered in dorsal line of caudal musculature, otherwise caudal fins translucent. Legs cream. Oral apparatus light cream. Iris copper-yellow, with small black reticulations. Variation. Variation of 26 meristic characters of tadpoles in Stages 35–39 (CJ 1959) are shown in Table 7. Total length varies between 61.3 (Stage 39) and 75.0 (Stage 35); tail length proportion varied from 63.2 to 67.0 until Stage 39; labial tooth row formula was 2/3(1). Number of marginal papillae varied among specimens and Gosner stages, variation in lower lip papillae is high (42–57). ......continued on the next page We documented changes in coloration during ontogenetic development of CJ 1958–9 (Fig. 20). At Stage 36, the dorsum and flanks are pale brown with dark brown areas on posterior body and flanks. By Stage 42, dorsal surfaces of body and limbs are uniform green; fine, cream, dorsolateral stripes are present, extending to midbody. Cream stripes border the outer dorsal margins of the limbs. At Stage 46, dorsal surfaces of body and limbs have more well-defined cream stripes, and a cream labial stripe; the canthus rostralis is green, and remaining flanks are mostly dark brown with an upper diffuse black-stripe. Dorsal surfaces of fingers and toes are brown except on finger and toe discs, which are yellowish-cream. The iris is copper-yellow, lacking the brown horizontal band that is observed in adults. Comparisons. Tadpoles of Gastrotheca turnerorum may occur in sympatry with those of G. elicioi, G. lojana, G. psychrophila, and G. pseustes, in the Loja-Abra de Zamora region. Gastrotheca turnerorum differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body; from G. lojana by having a more rounded tail terminus and having bold cream marks in a dorsal line of caudal musculature, and from G. pseustes by having a more rounded tail terminus (compare in Fig. 5). For G. psychrophila, see remarks under G. elicioi tadpole account. Distribution and ecology. Gastrotheca turnerorum only is known from three localities in the Cordillera Oriental of the Andes in southern Ecuador. These localities are in Parque Nacional Yacuri, Parque Nacional Podocarpus, and Reserva Tapichalaca in Zamora Chinchipe and Loja provinces. Its elevational range is 3073–3406 m in an area of extent of occurrence of about 450 km 2. This mostly nocturnal, semiarboreal species inhabits mainly paramos and a few forests in the Evergreen Shrub and Herbazal of Paramo, and the Evergreen Montane Forest of Catamayo-Alamor (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 895–1278 mm and the average annual temperature is 10.4– 13.5 °C (Fick & Hijmans 2017). At the Lagunas del Compadre in the Parque Nacional Podocarpus, a gravid female was collected on 1 December 1994. It deposited tadpoles on 12 December in the laboratory, some of them were raised, and others were preserved. On 20 September 2009, another female with eggs in her pouch was found at the same locality. It was sitting on moss at approximately 2 m from the Río Sabanilla, which is about 1.5 km from Lagunas del Compadre. The holotype, from Laguna Negra de Jimbura in the Parque Nacional Yacuri (Fig. 11C), was found at the edge of a pond about 3 x 2 m at the edge of a lagoon, at 14:00 h. The water temperature was 16.4° C, and water pH was 5.3. Eighteen tadpoles were collected at this pond. Another group of tadpoles was found at 18:00 h; at the same locality in another lagoon, the water temperature was 7.4 ° C. Strong winds blow across the lagoon, and only a small patch of forest remained at one border (Elicio E. Tapia field notes, 12 June 2011). A brooding female is depicted in Figure 10J. Conservation status. We suggest that Gastrotheca turnerorum should be considered as an Endangered species according to criteria B2ab(i,ii,iii) of the IUCN Red List. Although this species occurs at the Yacuri and Podocarpus National Parks, we suggest this conservation status because of its small known area of occurrence (159 km 2) that is vulnerable to improperly regulated tourism activities (Aguirre 2001), introduced species such as trout, climate change, and pathogens. Laguna Negra de Jimbura has suffered the synergistic effects of agriculture, cattle and sheep raising, fires, introduced species, pesticide use, and unregulated tourism activitites. Etymology. The specific name turnerorum is a latinized word that honors the Turner family. As unique as this marsupial frog is to the amphibian world, the Turner family stands out in their exceptional, unwavering and ever- growing commitment to conservation of the world natural resources. From Ted Turner’s pioneering efforts to protect very large land areas and restore them to their original and natural states of clean water and abundant fauna and flora to granddaughter Elizabeth’s awareness-raising book “Our friends the frogs,” there are multiple generations of active conservationists with a true appreciation for the value of all of our natural resources. In 1991, Laura Turner Seydel (with father Ted) co-founded the Captain Planet Foundation and continues to work diligently to inspire the same appreciation for conservation and sustainable living in the next generation of young people who will surely be making big decisions that will affect the health of our planet for themselves and their own grandchildren. Through the camera’s eye, Rhett Turner has managed to put conservation issues such as the pollinator perils and amphibian crisis in front of millions of people. Through the Turner Endangered Species Fund, there is hope for dozens of threatened species of plants and animals in the US and abroad that was not there before. Together, the Turner family has put their hearts, souls, and personal resources into helping make th

Gastrotheca yacuri sp. nov. urn:lsid:zoobank.org:act: 6261932E-02AB-4C9E-B2BD-BD213BBDC771 Holotype. CJ 7822 (Figs. 21���22), an adult male, from El Salado de Jimbura, at about 2.5 km (by road) west from the entrance in the Parque Nacional Yacuri, 2914 m (04�� 42' 08.37" S, 79�� 27' 00.09" W), Loja Province, Ecuador, collected on 9 April 2016 by Paul Sz��kely and Diana Sz��kely. Paratype. Ecuador: Loja: QCAZ 21105, adult male, from El Salado de Jimbura, 2712 m, on 26 June 2002 by Fernando Nogales Sornoza. Referred specimen. Ecuador: Loja: Not collected (Fig. 11O), subadult, 43 mm SVL, from nearby of Lagunas Negras, Parque Nacional Yacuri, 3492 m (04�� 42' 31.81" S, 79�� 25' 49.61" W). Diagnosis. Included in the genus Gastrotheca by molecular evidence and general morphological similarity with species of Gastrotheca. Although females are unknown, we infer presence of dorsal brood pouch because all its closely related species have it. A moderately large species (57.4���58.9 mm SVL in males, n = 2) with tibia length 47��49% SVL, larger than foot; (2) interorbital distance greater than width of upper eyelid; (3) skin on dorsum weakly granular to smooth, co-ossified with skull, having a transverse ridge at occipital region; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I shorter than Finger II, width of discs much wider than digits; (8) fingers unwebbed; (9) toes one-half webbed, webbing extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum uniform green or brown with brown paravertebral marks; (11) head markings consisting of brown labial stripe with cream glandular areas at posterior end, canthal and supraciliary stripe of black, brown, and bronze; (12) dorsolateral stripe absent or present, when present above consisting in a series of bronze and cream warts, bordered below with a black and brown line, (13) flanks brown and green with a blue and black reticulum towards the lower flanks, axillae, groin, anterior and posterior surfaces of thighs, shanks, and tarsus; brown pelvic patch; (14) venter cream with nearly uniformly distributed, small, dark brown marks or brown with cream marks; gular region dark brown in males. Gastrotheca yacuri differs from all other species of Gastrotheca in southern Ecuador by having a blue and black reticulated pattern on flanks, axilla, groin, thighs, and shanks. It most closely resembles two other species, G. pseustes and G. psychrophila by having the blue colors, but they lack a blue and black reticulum (compare these species in Fig. 10). Gastrotheca yacuri differs from its sister species, G. aguaruna, from the Cordillera Central in northern Peru, by lacking blue colors, having the skin co-ossified with skull, and being smaller (SVL of males of G. yacuri 57.4���58.9 mm vs 41.6���46.8 mm in G. aguaruna). The genetic distance between them is of at least 2.8 % (in a DNA dataset of 438 bp, 16S gene). Description of the holotype. An adult male (Figs. 21���22); body moderately robust; SVL 57.4 mm; head wider that long; snout slightly acuminate in dorsal view, bluntly rounded in profile; canthus rostralis angular in section; loreal region concave; lips rounded; top of head flat; interorbital distance 92% of width of upper eyelid; internarial area flat; nostrils not protuberant, directed anterolaterally, at about level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye by distance about equal to diameter of tympanum; tympanic evident; supratympanic fold elevated, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing four teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 28% of SVL); fingers unwebbed bearing fringes; discs large and rounded, width of disc of Finger III lesser that diameter of tympanum; relative lengths of fingers IColoration in life. The dorsal surfaces of the head, body, and limbs are green; the hidden surfaces of the forearm, thighs, and shanks are reticulated with black and blue markings (Figs. 21���22). Head markings are brown canthal and labial stripes. The canthal stripe is diffuse; the labial stripe varies from brown anteriorly to white posteriorly until the insertion of the forelimb. The dorsolateral stripe is thin and consists of a series of bronze and cream warts, bordered below with a black and brown line. The flanks are brown and green with a blue and black reticulum towards the lower flanks, axillae, groin, anterior and posterior surfaces of thighs, shanks, and tarsus; the pelvic patch is brown. The venter is cream with nearly uniformly distributed, small, dark brown marks; gular sac is nearly uniform grayish-brown with small dark brown marks. Palms and soles are pinkish gray; ventral fingers and toes are fleshy. The iris is copper with abundant black reticulations. Coloration in preservative. Similar to coloration in life, but green and blue surfaces have begun to fade. Measurements (in mm). SVL: 57.4, TIBL: 27.2, FL: 24.8, HL: 17.1, HW: 19.2, IOD: 6.9, EW: 5.2, IND: 2.8, ED: 6.0, EN: 3.0, TD: 3.3, FFL: 8.2, TFL: 16.2, TFD: 2.8. Variation. Morphometric variation of the male paratype is as follows: SVL: 58.9, TIBL: 28.6, FL: 28.3, HL: 17.9, HW: 20.1, IOD: 6.7, EW: 4.3, IND: 2.8, ED: 5.6, EN: 5.0, TD: 3.0, FFL: 10.8, TFL: 19.0, TFD: 2.7. The skin on the dorsum varies in the subadult by lacking a smooth tranverse occipital ridge. Color variation in life. The referred subadult specimen varies from the holotype in that it is mostly brown; it has a light reddish-brown dorsum of body and limbs with darker, well-defined stripes, two paravertebral and one at the coccyx position; scattered black spots are present on dorsum towards the flanks (Fig. 10O). The loreal region is brown, and a thin dark brown canthal line is present; the labial stripe is dark brown and bordered along its upper margin by a thin pale stripe that varies from brown anteriorly to cream posteriorly until the insertion of the forelimb. There is no dorsolateral stripe but the reddish-brown dorsal color contrasts with a broad dark brown band on the flanks. The flanks are brown with black and cream marks (without blue colors). The venter is mostly brown with cream marks that are larger towards the chest; the gular region is slightly darker that the venter. The palms and soles are gray. The iris is cooper with black reticulations and has a shade of red in the lower half. Vocalization. Four individuals of Gastrotheca yacuri were recorded at El Salado, Loja Province (Appendix III). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call is a complex call, composed of one to three (usually two) long pulsed notes and followed (or not) by one to three (usually two) short, single-pulsed notes (Fig. 23 A���G). The long note had a mean duration of 0.634 s (SD = 0.091) and consisted on average of 32.24 (SD = 7.504) distinct pulses, partly fused, without silent intervals (amplitude modulation close but less than 100%). The amplitude of the long note increases gradually towards the middle of the note after which it decreases gradually by the end. The short notes had a mean duration of 0.072 s (SD = 0.020) and the inter-note interval was on average of 0.549 s (SD = 0.214). The mean dominant frequency of the call was 1496.7 Hz (SD = 38.643), with a mean 90% bandwidth frequency of 1366.7���1637.9 Hz, having the highest values recorded among the analyzed species. The fundamental frequency is recognizable; when visible, 5 to 7 harmonics are distinguishable. Comparisons. The advertisement call of Gastrotheca yacuri is most similar to that of G. pseustes, but G. yacuri has a much shorter call duration, shorter long notes duration, shorter inter-note interval, a higher dominant frequency, and a higher 90% bandwidth frequency compared with the call of G. pseustes (Table 5). Also, G. yacuri emits usually only two short notes compared with 4���6 short notes of G. pseustes. The call of G. yacuri can be easily distinguished form those of G. lojana and G. testudinea by the amplitude modulation of the longer note, longer call duration, lower short note rate, larger number of pulses, higher pulse rate, a much higher dominant frequency, and higher 90% bandwidth frequency. Also, G. yacuri emits up to three (usually two) long notes per call compared with the only one emitted by G. lojana and G. testudinea (Table 5). Distribution and ecology. Gastrotheca yacuri is only known from three nearby localities 2.8 km apart (maximum distance) at elevations of 2712���3492 m in the Cordillera Oriental de los Andes in southern Ecuador. One locality is within Parque Nacional Yacuri and the others are in El Salado de Jimbura in Loja Province (Fig. 9). This nocturnal, semiarboreal species inhabits paramo and subparamo in the Evergreen Montane Forest of Catamayo-Alamor (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 895���1159 mm and the average annual temperature is 10.4���14.6 ��C (Fick & Hijmans 2017). The male holotype was found near the road, calling from the branches of a bush, at about 3.5 m above ground. In this same location, choruses were heard from April to late July 2016; the males were calling usually from the vegetation nearby the road. An additional subadult was found during the day in a large terrrestrial bromeliad (Fig. 24A, B). Females and tadpoles are unknown. In this location, in Parque Nacional Yacuri, Gastrotheca yacuri is syntopic with G. turnerorum. Conservation status. We suggest that Gastrotheca yacuri should be assigned to the Data Deficient category according to guidelines of the IUCN (2001), given the inadecuacy of the current information to establish its status. Further searches and definition of its distribution, area of occupancy, habitat use, and biological information are required. Currently its known area of occurrence is less than 10 km 2, thus it could qualify for a Critically Endangered category; however, we can reasonably predict that if further surveys are conducted within the protected Yacuri National Park, its area will increase to more than 100 km 2. In any case, Lagunas Negras de Jimbura have suffered the synergistic effects of agriculture, cattle and sheep raising, fires, introduced species such as trout, pesticide use, and unregulated tourism activitites. Areas in the Salado de Jimbura that are out of the Yacuri National Park already have been heavily modified by human actions. Etymology. The specific name yacuri is a noun in apposition and refers to the Yacuri National Park, where this species has been found. According to Chamba-Troya (2017), Yacuri takes its name from the Quechua words Yacu meaning water and Quri that means gold. Ecuador's southernmost national park ranges from 2000 to 3700 meters in the Cordillera Oriental. This park of 43,090,60 hectares is unique by sheltering a series of 48 glacial lakes, and many endemic species of flora and fauna. It is a Ramsar site (The Convention on Wetlands) and part of the Podocarpus-El C��ndor Biosphere Reserve declared by UNESCO. Comments. Gastrotheca yacuri belongs to the subgenus Gastrotheca, and is reported as Species D in Duellman (2015: Fig 12.1)., Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 56-61, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","IUCN (2001) Red list categories. Fersion 3.1. IUCN Species Survival Comission, Gland, 32 pp","Chamba-Troya, C. A. (2017) Faloracion del servicio ecosistemico de recreacion y belleza escenica del Parque Nacional Yacuri, 2015. Tesis de Economista, Universidad Tecnica Particular de Loja, Loja, 80 pp.","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp."]}

Gastrotheca lojana Parker 1932 Gastrotheca marsupiata lojana Parker, 1932:25.��� Holotype: BM 1947.2.31.13 (Fig. 25B) from Loja, Loja Province, Ecuador. Gastrotheca monticola (part)��� Duellman & Hillis 1987:158. Gastrotheca (Duellmania) lojana ��� Dubois, 1987:33. Referred specimens. Ecuador: Azuay: CJ 390, 401���2, from O��a, 2272 m (03�� 27' 41.04" S, 79�� 09' 46.47" W), on 15 June 2011 by Elicio E. Tapia, Sof��a Carvajal-Endara and Henry Grefa; KU 138401���3, from Gir��n, 2310 m (03�� 10' 00.12" S, 79�� 07' 59.88" W), on 0 6 June 1970 by Thomas A. Fritts; QCAZ 2692, from O��a, 2272 m (03�� 27' 41.04" S, 79�� 09' 46.47" W), on 1 December 1999 by Luis A. Coloma and Luis E. L��pez; QCAZ 26314���5, 26318, 26322���3, 26327���8, 26334���5, 26337���8, from O��a, 2272 m (03�� 27' 40.43" S, 79�� 09' 45.11" W), on 8 August 2003 by Luis A. Coloma and ��talo G. Tapia, QCAZ 31521���3, from O��a, 2272 m (03�� 27' 41.04" S, 79�� 09' 46.47" W), on 27 December 2005 by ��talo G. Tapia and Giovanni Onore; QCAZ 32212, from O��a, 2272 m (03�� 27' 41.04" S, 79�� 09' 46.47" W), on 2006 by Luis A. Coloma; QCAZ 32571, from O��a, 2272 m (03�� 27' 40.43" S, 79�� 09' 45.11" W), on 8 August 2003 by Luis A. Coloma, Erik R. Wild, Andr��s Merino-Viteri, ��talo G. Tapia and Edwin Patricio Vargas; QCAZ 42725, from San Fernando, Laguna de Busa, 2834 m (03�� 09' 15.80" S, 79�� 15' 49.03" W), on 22 October 2007 by Sof��a Carvajal-Endara, Amaranta Carvajal-Campos, Andrea Carvajal Endara. Loja: BM 1947.2.31.6���10, BM 1947.2.31.13���18, from Loja, 2150 m (04�� 00' 00" S, 79�� 13' 00.12" W), on 1930���1 by Clodoveo Carri��n Mora. KU 120673 ���4, from Loja, 2150 m (04�� 00' 00" S, 79�� 13' 00.12" W), on 9 June 1968 by John D, Lynch; KU 120675, from 2 km E Loja, 2100 m (03�� 59' 56.4" S, 79�� 13' 00.12" W), on 9 June 1968 by John D, Lynch, KU 138233, from 3 km W Loja, 2150 m (03�� 58' 47.99" S, 79�� 12' 54" W), on 21 June 1970 by Thomas H. Fritts; KU 138234, from 10 km W Loja, 2500 m (03�� 59' 56.4" S, 70�� 14' 43.08" W), on 27 June 1970 by Thomas H. Fritts; KU 138235���6, from 5 km N Loja, 2150 m (03�� 52' 00.12" S, 79�� 13' 00.12" W), on 28 June 1970 by Thomas H. Fritts; KU 138404���9, from Saraguro, 2412 m (03�� 36' 00" S, 79�� 13' 00.12" W), on 19���20 June 1970 by Thomas H. Fritts; KU 148568, from Saraguro, 2412 m (03�� 36' 00" S, 79�� 13' 00.12" W), on 22 May 1971 by Richard M. Montanucci; KU 178482���95, from 2 km SSW Saraguro, 2569 m (03�� 38' 22.92" S, 79�� 13' 59.87" W), on 5 January 1978 by John D. Lynch; KU 178496���7, from 2.1 km N Saraguro, 2575 m (03�� 36' 41.76" S, 79�� 13' 00.12" W), on 7 January 1978 by John D. Lynch; QCAZ 22371, from Loja Zamora Huayco, 3018 m (04�� 05' 49.9806" S, 79�� 10' 02.499" W), on 12 December 2002 by Diego Almeida-Reinoso; QCAZ 30788, from Saraguro, 2412 m (03�� 36' 00" S, 79�� 13' 00.01" W), on 20 September 1978 by MC; QCAZ 34505, from 2 km SW Saraguro, 2569 m (03�� 38' 22.99" S, 79�� 14' 24" W), on 5 January 1978 by John D. Lynch. El Oro: QCAZ 32724, from Guanaz��n, 2984 m (03�� 27' 18.43" S, 79�� 29' 23.64" W), on 3 December 2006 by Silvia Ald��s Alarc��n. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (54.1���76.1 mm SVL in females, n = 12; 40.2���61.0 mm SVL in males; n = 24) with tibia length 41��56% SVL, longer than foot; (2) interorbital distance about 1.6 times width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus distinct, smooth; (7) Fingers I and II about equal in length, discs on fingers about twice width of digits proximal to discs; (8) fingers unwebbed; (9) webbing between external toes extending to the antepenultimate subarticular tubercle on Toe IV, to point midway between penultimate and distal subarticular tubercles on Toe V; (10) in life, dorsum green, brown, or tan with dark paravertebral marks; (11) markings on the head consisting of pale labial stripe broader at the posterior half of lip and dark interorbital bar or two blotches connected or not to paravertebral marks; (12) pale cream or white dorsolateral stripe fragmented; (13) flanks dark brown, green or tan with pale cream spots ventrally, anterior and posterior surfaces of thighs, groin, and dorsal surface of foot with contrasting pattern of heavily white mottling on a dark background, with bluish or greenish tinges; dorsal surfaces of limbs with dark brown or green bars; (14) venter creamy white with dark brown dots or marks; (15) brood pouch single, dorsal. Gastrotheca lojana most closely resembles three other species in southern Ecuador (G. elicioi, G. cuencana, and G. litonedis) and one species in northern Peru (G. monticola). Gastrotheca lojana differs from G. elicioi in some coloration patterns (described above; also, compare in Figs. 10L, 29, 30 vs 12���14). A black canthal stripe is absent in G. lojana, but present in G. elicioi. The anterior and posterior surfaces of thighs and groin are heavily mottled in G. lojana, but slightly mottled in G. elicioi. The interorbital mark in G. lojana is a transverse bar that sometimes is divided into two blotches and that is usually connected with two broad paravertebral marks. In G. elicioi dorsal marks, when present, consist of a triangular interorbital mark, which may or may not be connected to two narrow and curved paravertebral marks. Gastrotheca lojana has a conspicuous, elevated row of dorsolateral warts, whereas it is barely raised in G. elicioi. In addition, in G. elicioi the dark bars on limbs, when present, are shorter and less defined than in G. lojana. Gastrotheca lojana differs from its most related species, G. cuencana and G. litonedis (compare Figs. 10L, 29, 30 vs 2���4, 26���27) by having: a cream venter with dark flecks or spots (uniform creamy white in G. cuencana and pale brownish gray in G. litonedis); an interorbital bar or blotches usually connected to paravertebral marks (absent in G. cuencana and G. litonedis); dorsal surfaces of limbs with dark transverse bars (irregular dark blotches in G. cuencana and usually unmarked in G. litonedis); anterior and posterior surfaces of thighs and groin heavily mottled (translucent cream without marks in G. cuencana and usually brown and slightly molted in some G. litonedis); dorsal surfaces of fingers cream or with dark brown spots (cream in G. cuencana and of same color of the rest of the body in G. litonedis); a fragmented dorsolateral stripe (continuous in G. cuencana and G. litonedis); and complex call (simple call in G. cuencana and G. litonedis). The genetic divergence between G. lojana and G. litonedis is 2.5% and between G. lojana and G. cuencana is 2.8% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca lojana differs from G. monticola by having the concealed surfaces of thighs, groin and dorsal surfaces of the feet with white mottling on a dark background, whereas in G. monticola the axilla and groin, concealed surfaces of thighs, and dorsal surfaces of the feet are green with black spots on a green or tan background. A distinctly different species, G. pseustes, occurs sympatrically with G. lojana in part of its range. Gastrotheca lojana differs from G. pseustes by having finely granular skin on the dorsum (smooth to coarsely granular in G. pseustes), larger digital discs, and pale dorsolateral and supracloacal stripes. Also the call of G. lojana can be easily distinguished form the call of G. pseustes by the amplitude modulation of the longer note, shorter call duration, higher short note rate, shorter interval between notes, fewer pulses, lower pulse rate, a much lower dominant frequency, and a lower 90% bandwidth frequency. Variation. Morphometric variation of 12 females and 24 males is summarized in Table 3. Females are larger than males (61.6�� 5.7 mm; 49.4�� 4.5 mm). All adults have a moderate supratympanic fold, which usually extends from the superior part of tympanum to point just above the insertion of forelimb. The dorsolateral row of warts can be less conspicuous in some specimens. Each vomer has four to ten teeth (5.7��1.4, n = 44). Color variation in preservative. In most preserved specimens the dorsum is bluish gray or brown with darker interorbital, paravertebral marks and bars on the limbs. Some specimens also have dark flecks densely distributed on dorsum. All specimens lack a dark canthal stripe and have a cream white supralabial stripe, which is broader at the posterior margin of the lip and that is extended from the posterior margin of the lip to the insertion of the forelimb. Most of the specimens have a dark brown stippling on the supralabial stripe. A fragmented supracloacal and heel stripes are present; also a white mark extends laterally from the vent. The flanks are dark gray with black and white dots; the posterior part of the flanks, groin, concealed surfaces of thighs and dorsal surface of foot are heavily mottled. The venter is white and varies from having a few dark flecks to having a heavily mottled pattern. Color variation in life. (Figs. 10L, 29, 30). In living individuals the color of the dorsum is green (CJ 408), brown (CJ 405) or tan (CJ 407) with darker interorbital and paravertebral marks. The interorbital mark is usually a bar that connects the two eyelids; however, in some individuals it is divided into two blotches (CJ 411). Paravertebral marks are broad and are usually connected with the interorbital mark (CJ 407) or, in some individuals (CJ 405), they are fused at the level of the scapular region. The venter is white in most specimens, some have dark flecks and spots evenly distributed, and others have a heavily mottled pattern. On the head there is a cream supralabial stripe, which is broader at the posterior margin of the lip and extends from the posterior margin of the lip to the insertion of the forelimb. The tympanum is brown, tan or olive green. The iris is reddish bronze with a few black reticulations. The flanks are brown, tan, or green with black and white spots. The posterior part of the flanks, groin, anterior and posterior surfaces of thighs have a heavily mottled pattern, some individuals have green or purple tinge on the groin and posterior surface of the thighs (Fig. 30). Supracloacal and heel stripes are cream or tan. The ventral surfaces of the shanks have a faint tinge of pale blue. Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36, from a series (CJ 1948) obtained from a pond at O��a, 2272 m, Azuay Province, Ecuador, by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa on 15 June 2011 (Fig. 5C). Total length 55.7; body length 19.8 (36% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat shape concave in lateral profile; body width at the level of spiracle 12.3, and height at same position10.2; head width at level of eyes 10.5. Lateralline system barely visible, supraorbital and infraorbital lines both originating at tip of snout, running in parallel to the eye and making contact immediately behind the eye; angular line descending vertically from just posterior of eye to throat; anterior oral line descending vertically from level of oral disc and just anterior to level of nares to throat there it curve to parallel infraorbital line, thereby forming a continuous circuit with angular and loreal lines; dorsal body and middle body lines present. Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 2.8; internarial distance 3.2. Eye positioned and directed dorsally, not visible from below; eye length 2.1, eye width 1.9; interorbital distance 5.7. Spiracle sinistral, located ventrolaterally; spiracular opening directed posteriorly; distance from tip of snout to spiracular opening 13.7; spiracle end rounded, attached to body wall, inner wall of spiracle not evident; tube length 4.0, tube transverse width 2.9. Vent tube dextral, its opening directed posteriorly, tube length 3.7, tube transverse width 2.1. Tail length 37.0, caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 5.2, caudal muscle width 4.3; caudal fins well developed and about the same size, originating near tail-body junction; maximum height of tail 13.6; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 4.1. It is surrounded by a uniserial row of short, marginal papillae, interrupted medially on upper lip; lower lip papillae alternating in and out, giving the appearance of two series; upper lip with 21 papillae on right side and 19 papillae on left side; lower lip with 63 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, height 0.4, transverse width 3.0 (48% of oral disc width); lower jaw sheath V- shaped, open and finely serrated, width 2.1, height 0.7. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.2, A2: 4.1, P1 right row 1.9, P1 left row 1.6, P1 gap 0.3, P2: 3.2, P3: 3.5. (Fig. 6C). Color in preservative. Dorsum dull gray, with paler (translucent) areas on the flanks and snout. Caudal muscles pale gray with small cream flecks; dorsal and ventral fins translucent with evenly distributed cream flecks, except at tip where flecks are absent. Venter translucent on throat and belly regions, guts not exposed, white pigment partially covers the belly, producing a black and white mottling; eyes lavender gray with white flecks; oral apparatus translucent. Color in life. In dorsal and lateral views, body olive-cream; loreal and snout areas palest. Ventrally, guts not visible, belly white with gray marks; gills visible through the throat, with a reddish hue. Caudal musculature cream-gray; proximal third reddish with myomeres and medial line well defined; distal two thirds with small cream flecks; dorsal and ventral fins translucent, suffused with minute cream flecks, clustering in rounded patches near borders of fins; most dense near tail-body junction. Spiracle, oral apparatus, and legs olive-cream. Iris gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 34���41 (CJ 1948) are shown in Table 9. Total length varies between 40.9 (Stage 34) and 61.2 (Stage 41) and tail length proportion varies from 59.1 to 63.7 until stage 41; labial tooth row formula 2/3(1). Number of marginal papillae varies among specimens and Gosner stages, variation in number of ventral papillae at lower lip is moderate (60 and 69). The color of venter varies from nearly plain gray (CJ 4303) (Fig. 31) to gray flecks in a white background (CJ 1948) (Fig. 5C). Duellman (2015) described a tadpole (KU 203548) from 7.9 Km W Loja. Nonetheless, it may also belong to either G. pseustes, G. psychrophila, or G. turnerorum. We documented changes during ontogenetic development of CJ 4303, 4304 (Figs. 31 A���C, 32). At Stage 46, the dorsum is nearly uniform clear brown; the paraverterbal marks and interorbital marks are green and vary from nearly absent to well defined. Comparisons. Tadpoles of Gastrotheca lojana may occur in sympatry with those of G. elicioi, G. psychrophila (tadpole unknown, see Remarks under G. elicioi tadpole account), G. pseustes 1, and G. turnerorum in the Loja- Abra de Zamora region, with G. litonedis and G. pseustes 2 (tadpole not described), in the Laguna de Busa area and with at least G. pseustes in other localities (compare in Fig. 5). Gastrotheca lojana differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body, from G. pseustes 1 by lacking a reticulated pattern on flanks, from G. turnerorum by having a more pointed tail terminus and lacking bold cream marks in a dorsal line of caudal musculature, and from G. litonedis by having a highest tail dorsal fin. Vocalization. Five individuals of Gastrotheca lojana were recorded from one location in Azuay Province (one individual from San Fernando, Laguna de Busa) and from one location in Loja Province (four individuals from O��a) (Appendix III). The advertisement call of G. lojana is a complex call, composed of one long pulsed note and followed (or not) by one or two short, single-pulsed notes (Fig. 17 H���N). The long note had a mean duration of 0.562 s (SD = 0.125) and consisted on average of 17.59 (SD = 2.418) distinct pulses (pulse series), separated by silent intervals (amplitude modulation of 100%). The amplitude of the long note increases gradually towards the end, without falling. The short notes have a mean duration of 0.064 s (SD = 0.027) and the inter-note interval is on average of 0.356 s (SD = 0.107). The mean dominant frequency is 1118.1 Hz (SD = 126.872), with a mean 90% bandwidth of 986.1���1348.4 Hz. The fundamental frequency is not clearly recognizable; when visible, 5 to 7 harmonics are distinguishable in the short notes. Comparisons. The advertisement call of Gastrotheca lojana is most similar to that of G. testudinea, but G. lojana has a shorter call duration, higher short note rate, shorter long notes, short notes duration, shorter inter-note intervals, higher pulse rate, a higher dominant frequency and a higher 90% bandwidth frequency compared with the call of G. testudinea (Table 5). The call of G. lojana can be easily distinguished form the calls of G. yacuri and G. pseustes by the amplitude modulation of the longer note, shorter call duration, higher short note rate, shorter inter-note interval, smaller number of pulses, lower pulse rate, a much lower dominant frequency, and a lower 90% bandwidth frequency. Also, G. lojana emits only one long note, whereas G. yacuri emits up to three (usually 2) long notes per call and G. pseustes emits a larger number of long and short notes (Table 5). Distribution and ecology. Gastrotheca lojana is known in basins within Azuay, Loja, and El Oro Provinces. Its elevational range is 1682���3018 m in an area of extent of occurrence of about 1621.2 km 2. This nocturnal, semiarboreal species inhabits mostly disturbed areas and a few forests in the Evergreen Montane Shrub in the south of the, Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 71-79, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Parker, H. W. (1932) Some new or rare reptiles and amphibians from southern Ecuador. The Annals and Magazine of Natural History, Series 10, 9 (49), 21 - 26. https: // doi. org / 10.1080 / 00222933208673460","Duellman, W. E. & Hillis, D. M. (1987) Marsupial frogs (Anura: Hylidae: Gastrotheca) of the Ecuadorian Andes: resolution of taxonomic problems and phylogenetic relationships. Herpetologica, 43, 141 - 173.","Dubois, A. (1987) Again on the nomenclature of frogs. Alytes, 6 (1 - 2), 27 - 55.","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp.","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","Duellman, W. E. (1974) A systematic review of the marsupial frogs (Hylidae: Gastrotheca) of the Andes Ecuador. Occasional Papers of the Museum of Natural History, The University of Kansas, Lawrence, 22, 1 - 27.","Loaiza-S, C. R. (2012) Clodoveo Carrion Mora (1883 - 1957). Cientifico Insigne de la Provincia de Loja. Universidad Tecnica Particular de Loja, Ediloja, Loja, 255 pp.","Duellman, W. E., Barley, A. J. & Venegas, P. J. (2014) Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru. Zootaxa, 3768 (2), 159 - 177. https: // doi. org / 10.11646 / zootaxa. 3768.2.4"]}

Gastrotheca cuencana sp. nov. urn:lsid:zoobank.org:act: 82F87E67-2940-424C-AC03- BC751224 D5E4 Holotype. CJ 1391 (Fig. 2), an adult male, from the city of Cuenca, 2579 m (02�� 53' 57.91" S, 79�� 01' 52.79" W), Azuay Province, Ecuador, one of a series obtained on 8 June 2011 by Elicio E. Tapia, Sof��a Carvajal-Endara and Henry Grefa. Paratypes. (Total 48: 10 adult males, 21 adult females, 17 juveniles). Ecuador: Azuay: CJ 1392���7, collected with the holotype; KU 120676 (female), 120683 ���4, 120690, 120695 (4 juveniles), 120699 (male), 120705 (male), 120709 ���10 (males), 120713 (female), 120718 ���9 (female, male), 120721 (female), 120722 (male) from Cuenca, 2600 m, (02�� 53' 57.91" S, 79�� 01' 52.79" W), on 19 June 1968 by John D. Lynch; KU 138616 (female), 138619��� 21 (male, females) from 4 km E: Cuenca 2540 m (02�� 52' 59.88" S, 78�� 22' 0.11" W), on 10 June 1970 by Thomas H. Fritts; KU 141572 (female) from 2.1 km S Cutchil, 2720 m (03�� 06' 00"S, 78�� 47' 59.99"W), on 16 May 1971 by Richard R. Montanucci; KU 141579 (female) from 3.5 km S Cutchil, 2785 m (03�� 04' 59.87" S, 78�� 47' 59.99" W), on 14 May 1971 by Richard R. Montanucci; KU 141582 (juvenile) from 8 km NW Cuenca, 2803 m (02�� 51' 47.88" S, 78�� 58' 59.88" W), on 16 May 1971 by Richard R. Montanucci; KU 141583 (female) from 8.8 km NW Cuenca, 3820 m (02�� 52' 00.12" S, 79�� 04' 00.12" W), on 15 May 1971 by Richard R. Montanucci; KU 203441 (male) from Laguna de Zurucuchu, 16 km NW Cuenca, 3200 m (02�� 50' 23.99" S, 79�� 07' 47.99" W), on 5 March 1984 by William E. Duellman; KU 129779 ���82 (4 females), 129783���91 (9 juveniles), 129793���94 (juveniles), 129795 (female), 129796 (juvenile) from R��o Matadero, 12 km E Cuenca (02�� 52' 59.88"S, 78�� 58' 00.12" W), on 17 November 1968 by Craig E. Nelson. Ca��ar: KU 141571 (male) and KU 141573 (female) from Bibli��n, 2620 (02�� 42' 00" S, 78�� 52' 00.12" W), on 17 and 23 May 1971, respectively, by Richard R. Montanucci; KU 142620 ���24 (4 females, male), from Bibli��n, 2620 m (02�� 42' 00" S, 78�� 52' 00.12" W), on 25 July 1971 by William E. Duellman; KU 147113 (female) from Bibli��n, 2620 m (02�� 42' 00" S, 78�� 52' 00.12" W), on 15 January 1972 by John E. Simmons. Referred specimens. (Total 36, 28 adult males, 8 adult females). Ecuador: Azuay: QCAZ 1239 (male), from Sigsig, 2480 m (03�� 03' 08.4" S, 78�� 48' 10.56" W), on 24 July 1989 by Luis A. Coloma and Luis E. L��pez; QCAZ 26309 (female), from Cumbe, 2740 m (03�� 05' 55.64" S, 79�� 00' 28.73" W), on 8 August 2003 by Patricio Vargas; QCAZ 26348���50 (males), 26353 (female), 26357���64 (8 males), from Cumbe, 2740 m (03�� 05' 55.64" S, 79�� 00' 28.73" W), on 9 August 2003 by Luis A. Coloma, ��talo G. Tapia, Andr��s Merino Viteri, Erik Wild and Patricio Vargas Mena; QCAZ 26354 (female), from Cumbe, 2740 m (03�� 05' 55.64" S, 79�� 00' 28.73" W), on 9 August 2003 by ��talo G. Tapia; QCAZ 26357���8 (males), from Cumbe, 2740 m (03�� 05' 55.64" S, 79�� 00' 28.73" W), on 9 August 2003 by Andr��s Merino Viteri; QCAZ 31477 (male), from Sigsig, 2424 m (03�� 03' 08.4" S, 78�� 48' 10.56" W), on 8 March 2006 by ��talo G. Tapia; QCAZ 31509, 31511 (males), from Sigsig, 2424 m (03�� 03' 8.4" S, 78�� 48' 10.56" W), on 9 March 2006 by ��talo G. Tapia and Giovanni Onore; QCAZ 34131 (female), from Cuenca, 2579 m (02�� 53' 57.91" S, 79�� 01' 52.79" W), on 27 February 1979 by Fernando I. Ortiz; QCAZ 37375���6, 37379���81, QCAZ 37386 (6 males), from unknown locality, on 31 May 2007 by Zool��gico Amaru; QCAZ 38232 (male), from Carmen del Guzho, 2666 m (02�� 56' 00" S, 79�� 03' 00" W), on 16 June 2007 by Hari Gonz��lez Maldonado; QCAZ 39384���5 (female, male), from Tarqui, Patapamba, 2600 m (03�� 00' 38.16" S, 79�� 01' 53.43" W), on 21 June 2007 by Ernesto Arbel��ez Ortiz; QCAZ 42720���1 (males) from Cuenca, 2579 m (02�� 53' 57.91" S, 79�� 01' 52.79" W), on 3 October 2007 by Sof��a Carvajal-Endara and Flavio Jaramillo; QCAZ 47106 (male), from Tarqui, 2741 m (03�� 00' 57.17" S, 79�� 02' 40.13" W), on 21 June 2007 by Ernesto Arbel��ez Ortiz. Ca��ar: QCAZ 42826, 42835, 42841 (females), from Papaloma de la Nube, 3011 m (02�� 40' 21.54" S, 78�� 54' 21.28" W), on 12���14 August 2008 by Sof��a Carvajal-Endara. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (49.0��� 61.6 mm SVL in females, n = 28; 45.2���53.7 mm SVL in males, n = 38), with tibia length 39��50% SVL, slightly shorter than length of foot; (2) interorbital distance slightly larger than width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus weakly granular to smooth; (7) Fingers I and II about equal in length, width of discs greater than digits; (8) fingers unwebbed; (9) webbing between external toes extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum green, tan, brown, or reddish-brown with or without dark paravertebral marks; (11) head markings consisting of pale labial stripe and, in some individuals, an inconspicuous canthal stripe; (12) pale white or cream dorsolateral stripe present; (13) flanks tan, brown, or green with cream flecks or spots; groin and anterior and posterior surfaces of thighs translucent cream without marks and with a faint pale blue tinge in some; (14) venter uniform creamy white; (15) in females, brood pouch single, dorsal. In comparison with similar species in Ecuador, G. cuencana is most like G. litonedis, G. lojana, G. elicioi, and G. riobambae. It differs from these species by having a uniform creamy white venter; whereas the venter is pale gray in G. litonedis, and cream with dark flecks, spots or mottling in G. lojana, G. elicioi, and G. riobambae. In G. cuencana the dorsal surfaces of fingers are cream, whereas in the other species dorsal surfaces of fingers are of the same color as the rest of the body (tan, brown, or green). Gastrotheca cuencana also differs from G. litonedis, G. elicioi, and G. riobambae by lacking a dark canthal stripe. Moreover, in G. cuencana the groin and anterior surfaces of thighs are mostly translucent cream without marks; whereas, in G. litonedis the groin and anterior surface of thighs are usually bronze-brown or tan with cream or darks flecks and spots. In G. lojana and G. riobambae the groin and anterior surfaces of the thighs are usually cream with black mottling; in G. elicioi the groin and anterior surfaces of thighs usually have dark and cream flecks and spots. In profile, the snout in G. cuencana is rounded above and inclined anteroventrally, whereas the snout in G. elicioi, G. lojana, and G. litonedis is bluntly rounded. Furthermore, G. cuencana has a conspicuous, elevated row of dorsolateral warts whereas these are barely raised in G. litonedi s; a narrow, cream supracloacal stripe is present in G. litonedis, G. lojana, and G. riobambae; whereas the stripe is fragmented in G. elicioi and absent in most G. cuencana. Gastrotheca lojana and G. riobambae also differ from G. cuencana and G. litonedis by having a complex call, whereas the call in the latter two species is a simple call. Additionally, the call of G. cuencana has a lower note rate, shorter note duration, longer inter-note interval and a lower dominant and 90% bandwidth frequency compared with G. litonedis. Finally, G. cuencana differs from its sister species (G. litonedis) by having a genetic distance of 1.2% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca cuencana occurs in syntopy with G. pseustes (sensu lato) throughout most of its range. Gastrotheca pseustes differs from G. cuencana by having coarsely granular skin on the dorsum, in contrast to the usually finely granular dorsum in G. cuencana. In G. cuencana the venter is uniform creamy white, whereas it is cream or white with dark flecks or spots in G. pseustes; also the call of G. cuencana is simple, whereas in G. pseustes the call is complex. Description of holotype. An adult male (Fig. 2); body moderately robust; SVL 50.6 mm; head wider that long; snout rounded in dorsal view, rounded and inclined anteroventrally in profile; canthus rostralis round in section; loreal region concave, lips rounded; top of head flat; interorbital distance 104% of width of upper eyelid; internarial area slightly elevated; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from eye by distance about equal to diameter of tympanum; tympanic annulus and membrane weakly granular; supratympanic fold moderately weak, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing four teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 35% of SVL), unwebbed, with distinct narrow lateral fringes; discs moderately large and rounded, width of disc of Finger III greater that diameter of tympanum; relative lengths of fingers I=IIColoration in life (Fig. 2). The dorsum is yellowish brown with two darker, black delineated, continuous, paravertebral markings. These markings extend to the sacrum where they bifurcate and become two rows of irregular blotches at each side of the vertebral axis. The dorsal surfaces of the limbs are yellowish brown with irregular gray blotches. Cream dorsolateral and labial stripes are present; the latter continues to the insertion of forelimb; a faint brown canthal stripe is present. The flanks are bronze-brown with small black spots. The venter, groin, and anterior surfaces of the thighs are uniform creamy white; the posterior surfaces of the thighs have a faint bluish coloration. The dorsal surfaces of the fingers are cream. The gular region is creamy gray laterally. The iris is reddish bronze with black reticulations. Coloration in preservative. The dorsum is tan with two well-defined, gray paravertebral markings, which bifurcate at the level of sacrum to become two rows of irregular blotches on each side of the vertebral axis. A white dorsolateral stripe is present. The dorsal surfaces of limbs are tan with irregular gray blotches. A cream labial stripe continues to the insertion of the forelimb; a faint canthal is present. The flanks are gray; the venter, groin, and anterior surfaces of thighs are uniform creamy white; the posterior surfaces of thighs have a faint bluish coloration. The gular region is creamy gray. Measurements (in mm). SVL: 50.4, TIBL: 21.9, FL: 24.9, HL: 16.9, HW: 19.1, IOD: 5.2, EW: 5.0 IND: 3.3, ED: 5.6, EN: 4.1, TD: 2.8, FFL: 10.0, TFL: 17.5, TFD: 3.4. Variation. Morphometric variation of 28 females and 38 males is summarized in Table 3. Females are larger than males (54.2�� 2.9 mm; 49.6�� 2.4 mm). The skin on the dorsum is finely granular in most, but it varies in some specimens from weakly areolate to coarsely granular. All adults have a supratympanic fold that usually extends from the posterior part of the tympanum to point above the insertion of forelimb. The tympanic annulus is usually smooth, but some individuals (e.g., CJ 1395, 1940) have a slightly granular annulus (Fig. 3A, C). Each dentigerous vomerine process has 2���7 teeth (4.4��1.1, n = 39). Color variation in preservative. Preserved specimens have bluish gray dorsum (tan and reddish-brown in CJ 1397). Faint paravertebral marks are present. A canthal stripe is absent in most specimens; the flanks and posterior surfaces of the thighs are dark gray with white spots evenly distributed in some specimens. In all specimens, a prominent white labial stripe extends from the posterior margin of the lip to the insertion of the forelimb. A narrow white dorsolateral stripe is evident in most specimens. The groin and the anterior surfaces of the thighs are uniform creamy gray. The ventral surfaces are uniform creamy white. The posterior surfaces of thighs usually are creamy gray, but they are pale blue in some specimens. Some males have yellowish brown nuptial pads on the medial surface of the thumb. Color variation in life. (Figs. 3���4). The dorsum is uniform green (CJ 1393 ), reddish brown (CJ 1397), or tan (CJ 1392) with (CJ 1391) or without (CJ 1392) darker paravertebral marks. When present, the paravertebral markings usually are well defined; at the level of the sacrum they become two rows of irregular blotches at each side of the vertebral axis. The ventral surfaces are uniform creamy white. A cream labial stripe is extends from the posterior margin of the lip to the insertion of the forelimb; a dark brown canthal stripe is absent in most individuals. The tympanum is brown, tan, or olive green. The iris is reddish bronze with black reticulations. A creamy white dorsolateral stripe is evident in most individuals. The flanks are cream, bronze brown, or green. The groin and anterior surfaces of the thighs are translucent cream without dark markings (Fig. 3). The posterior surfaces of the thighs are translucent, cream, or pale blue or green. When present, supracloacal and heel stripes are cream or tan. The ventral surfaces of the shanks have a faint tinge of pale blue. The gular surface varies from white to creamy gray. ������continued on the next page ������continued on the next page Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 38 (CJ 1945a), from a series of eight tadpoles (CJ 1945) obtained from a pond at south Cuenca, 2579 m, Azuay Province, Ecuador, by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa on 8 June 2011 (Fig. 5A). Total length 44.6; body length 17.3 (39% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 11.5, and height at same position 9.3, head width at level of eyes 8.5. Lateral line system present but barely visible, supraorbital and infraorbital lines both rising at level of tip of snout, extending parallel to the eye and making contact immediately behind eye; angular line descending vertically just porterior of eye to throat, it dorsally contacts with post infraorbital line; anterior oral line descending vertically from oral disc level and behind nares level to throat; it makes a curve that parallels infraorbital line, forming a circuit continuous with angular and loreal lines; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 2.3; internarial distance 2.4. Eye directed dorsally; eye length 1.8, eye width 1.8; interorbital distance 4.3. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 11.0; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; tube length 3.1, tube transverse width 2.0. Vent tube dextral, opening oriented posteriorly, tube length 1.7, tube transverse width 1.8. Tail length 27.2; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 3.7, width 3,2; caudal fins well developed and proportional, arising near tail-body junction, forming a low hump; dorsal fin height 3.0, ventral fin height 2.8; maximum height of tail 9.3; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 4.7. It is surrounded by a uniserial row of marginal papillae, interrupted medially on upper lip; lower lip papillae alternate in orientation, giving appearance of two rows; upper lip with 29 papillae on right side and 27 papillae on left side; lower lip bearing 62 marginal papillae; upper jaw sheath medium-sized, forming a smooth arch and finely serrated, height 0.36, transverse width 2.8 (60% of width of oral width); lower jaw sheath V- shaped, open and finely serrated, width 1.92, height 0.7. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 3.6, A2: 3.5, P1 right row 1.5, P1 left row 1.6, P1 gap 0.2, P2: 3.4, P3: 3.0. (Fig. 6B). Color in preservative. Dorsum dull gray with darker areas on flanks, above eye, and on throat; body contour and snout translucent. Caudal musculature and fins with scattered cream dots, with higher suffusion near tail base, otherwise translucent. Venter pale, translucent in belly region, guts exposed; eyes lavender gray; oral apparatus translucent. Color in life. In dorsal view, body olive-cream; snout and flanks paler olive-cream. In ventral view, guts dark gray in belly area; reddish gills visible throughout throat. Caudal musculature covered by a nearly continuous line of cream marks, proximal half of musculature reddish cream with myomeres and medial line well defined, distal half translucent; dorsal and ventral fins translucent, with scattered cream dots, heavily suffused near tail-body junction. Spiracle, oral apparatus, and legs olive-cream. Iris gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 37���39 (CJ 1945) are shown in Table 4. Total length varies, Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 17-34, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","Amphibian Ark (2013) El Centro de Conservacion de Anfibios en el Zoologico Amaru, Ecuador. Boletin Informativo Amphibian Ark, 25, 13.","Siavichay-Pesantez, F., Maldonado-Cedeno, G. & Mejia-Coronel, D. (2016) Anfibios Urbanos de Cuenca. Editorial Don Bosco, Centro Grafico Salesiano, Cuenca, 108 pp.","Arbelaez Ortiz, E. & Vega Toral, A. (2008) Guia de Anfibios, Reptiles y Peces del Parque Nacional Cajas / Cajas National Park Amphibian, Reptile and Fish Guide. ETAPA, Municipalidad Cuenca, Grafisum, Cuenca, 106 pp.","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp."]}

Gastrotheca litonedis Duellman & Hillis 1987 Holotype. KU 202690 (Figs. 10A, 25A), an adult female, from 10 km (by road) northeast of Gir��n, 2750 m (03�� 07' 11.8" S, 79�� 06' 28.4'' W), Azuay Province, Ecuador, obtained on 7 March 1984 by William E. Duellman. Referred specimens. (Total 28: 13 males, 15 females). Ecuador: Azuay: CJ 386 ��9 (females), 401��4 (females, male), 1404��8 (males), KU 335388 ���9 (male, female), San Fernando, Laguna de Busa, 2834 m (03�� 09' 15.8" S, 79�� 15' 49.03" W), collected on 9 June 2011 by Elicio E. Tapia, Sof��a Carvajal-Endara and Henry Grefa; QCAZ 42734 (female), San Fernando, Laguna de Busa, 2834 m (03�� 09' 15.8" S, 79�� 15' 49.03" W), collected on 27 October 2007 by Sof��a Carvajal-Endara, Amaranta Carvajal-Campos and Andrea Carvajal-Endara; QCAZ 42855 (female), 42857��9 (females), 42861 (female), 42866 (male), 42871 (male), San Fernando, Laguna de Busa, 2834 m (03�� 09' 15.8" S, 79�� 15' 49.03" W), collected on 18���20 August 2008 by Sof��a Carvajal-Endara; QCAZ 49973 ��4 (female, male), 49976��8 (males), San Gerardo, 2854 m (03�� 08' 00.17" S, 79�� 11' 36.78" W), collected on 1 December 2010 by Sof��a Carvajal-Endara. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (53.5��� 62.4 mm SVL in females, n = 22; 48.9���57.4 mm SVL in males, n = 13) with tibia length 40��54% SVL, slightly longer than foot; (2) interorbital distance about 1.5 times width of upper eyelid; (3) skin on dorsum smooth, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus distinct, smooth; (7) Fingers I and II about equal in length, width of discs notably wider than digits; (8) fingers unwebbed; (9) webbing between external toes extending to the penultimate subarticular tubercle on Toe IV, to the distal subarticular tubercle on Toe V; (10) in life, dorsum green, brown, or tan with or without dark paravertebral marks or dark flecks; (11) head markings consisting of a pale cream labial stripe and narrow dark brown canthal stripe; (12) pale creamy white dorsolateral stripe present; (13) flanks, anterior surfaces of thighs and groin pale brown or tan slightly molted in some, posterior surfaces of thighs dark brown���gray with pale cream warts in some; (14) venter pale brownish gray; (15) brood pouch single, dorsal. Gastrotheca litonedis most closely resembles five other species in southern Ecuador��� G. cuencana, G. plumbea, G. pseustes, G. turnerorum, and G. elicioi. Gastrotheca litonedis differs from all of them by having a pale brownish gray venter; whereas, the venter is uniform creamy white in G. cuencana, cream in G. plumbea, cream with dark flecks or spots in G. pseustes and G. elicioi, and bronze or dark brown in G. turnerorum. Gastrotheca litonedis, like G. plumbea, has smooth skin on the dorsum, whereas it is finely granular in G. elicioi and G. cuencana, weakly areolate to smooth in G. pseustes and areolate in G. turnerorum. A pale supracloacal stripe is present in G. litonedis, G. pseustes, and G. turnerorum, but it is absent in G. cuencana and G. plumbea. Gastrotheca pseustes differs further from G. litonedis by having a complex call structure (long and short pulsed-notes), which is simple in G. litonedis (only short pulsed-notes). The sister species of G. litonedis is the parapatric G. cuencana. Distinctive morphological and coloration features (compare Figs. 26���27, vs 2���4) are (character states for G. cuencana in parentheses): Gastrotheca litonedis is larger than G. cuencana (SVL student���s t-test: t = -6.725, df = 99, p G. litonedis has a higher note rate, longer note duration, shorter inter-note interval, higher dominant frequency, and higher 90% bandwidth frequency compared with G. cuencana. Finally, these two species also differ by having a genetic distance of 1.2 % (in a DNA dataset of 438 bp, 16S gene). Gastrotheca litonedis is syntopic with G. lojana and G. pseustes 2, from which it differs notably by the texture of skin on dorsum, ventral coloration, and call structure. The skin on the dorsum is smooth in G. litonedis, whereas it is finely granular in G. lojana and coarsely granular in G. pseustes 2; furthermore, both species have cream venters with dark flecks, spots, or mottling, and a complex call structure with long and short pulsed-notes (G. lojana call in Fig. 17 H���N). Variation. Morphometric variation of 22 females and 13 males (from Duellman 2015) is summarized in Table 3. Females are larger than males (58.0�� 2.4 mm; 53.8�� 2.8 mm). The skin on the dorsum is mostly smooth in most specimens; however, in some specimens it is weakly areolate. All adults have a moderate supratympanic fold, which usually extends from the upper part of the tympanum to a point above the insertion of the forelimb. Some individuals have several small rounded tubercles anterolateral to the tympanum. Each dentigerous vomerine process has 2���10 teeth (5.4��1.6, n = 34). Color variation in preservative. In most specimens the dorsum is bluish gray without marks; in a few specimens faint paravertebral marks are present. The narrow canthal stripe, flanks, and posterior surfaces of thighs are dark brown. A creamy white supralabial stripe from the posterior margin of the lip to the insertion of the forelimb is prominent in all specimens. Creamy white dorsolateral, supracloacal and heel stripes are present. On the posterior part of the flanks the coloration varies from small white flecks to mottling on a dark brown ground color. The anterior surfaces of the thighs are usually gray with black flecks or spots. The ventral surfaces are uniform dull gray in most specimens; some have a pale gray venter with evenly distributed dark flecks and spots, and in CJ 1412 the venter is pale gray with black markings. Some specimens have a faint tinge of pale blue on the ventral surfaces of the shanks. Males have a cream nuptial pad on the medial surface of the thumb. Color variation in life. (Figs. 26, 27). The dorsum varies from uniform green (QCAZ 42858), to brown (CJ 404); in some individuals dark paravertebral marks and dark brown flecks are present (QCAZ 49977). The ventral surfaces are pale gray in most specimens; some have evenly distributed dark flecks and spots, and some have a faint bluish tinge on the ventral surfaces (QCAZ 49977). A narrow dark brown canthal stripe is present; a cream supralabial stripe extends from the posterior margin of the lip to the insertion of the forelimb. The tympanum is brown, tan, or olive green. The iris is reddish bronze with dense black reticulations. The posterior surfaces of the thighs are dark brown or cream with orange flecks. A cream dorsolateral stripe is evident in most individuals. The flanks, groin, and anterior surface of the thighs usually are bronze-brown or tan with cream or dark flecks and spots. In some individuals a pink-orange tinge is present in the groin (Fig. 27). The supracloacal and heel stripes are cream or tan, and the ventral surfaces of the shanks have a faint pale blue tinge. Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 37 (CJ 6558), from a series of 79 tadpoles (CJ 6557) born and reared in laboratory conditions, from a mother collected when gravid in the field (CJ 6018) at Laguna de Busa, 2780 m, Azuay Province, Ecuador, by Manuel A. Morales-Mite on 0 6 January 2016 (Fig. 5A). Total length 52.8; body length 19.4 (37% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 12.5, and height at same position 10.3, head width at level of eyes 10.0. Lateral line system present but barely visible, supraorbital and infraorbital lines not evident at level of snout, infraorbital line present at level of the eye, touching the inferior portion of the orbit, and making contact with supraorbital line immediately behind the eye. Postorbital line represented by three stitches arranged in line and forming a 45-degree angle with the body plane. Angular line, anterior oral line and loreal lines not visible; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 4.2; internarial distance 2.8. Eyes dorsally positioned; eye length 1.9, eye width 1.8; interorbital distance 5.2. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 12.9; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; spiracle length 2.7, spiracle transverse width 3.5. Vent tube dextral, opening oriented posteriorly, tube length 2.5, tube transverse width 2.2. Tail length 33.3; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 4.1, width 3.6; caudal fins well developed and proportional, dorsal fin arising abruptly near tail-body junction; dorsal fin height 4.1, ventral fin height 3.7; maximum tail height 9.2; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 5.8. It is surrounded by a uniserial row of marginal papillae, interrupted medially on upper lip. Lower lip papillae alternate in orientation, giving appearance of two rows; upper lip with 23 papillae on right side and 23 papillae on left side; lower lip bearing 57 marginal papillae. Upper jaw sheath medium-sized, forming a smooth arch and finely serrated, transverse width 2.9 (50% of width of oral width) and height 0.3. Lower jaw sheath V- shaped, open and finely serrated, width 2.6, and height 0.7. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 4.5, A2: 4.4, P1 right row 1.65, P1 left row 2.0, P1 gap 0.2, P2: 4.1, P3: 3.95. (See also Fig. 6A of CJ 5292, Stage 35). Color in life. Based on a specimen in Stage 34 from a series (CJ 1947) obtained from San Fernando, Laguna de Busa, Azuay Province, 2834 m, by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa on 9 June 2011 (Fig. 6A). In dorsal view, body olive-gray, head and tail suffused with pink and cream. Flanks cream anteriorly to olivegray posteriorly. In ventral view, cream anteriorly in gular region; followed posteriorly by red gills visible troughout the throat, and olive-gray guts. Caudal musculature pink proximally, gradually becoming gray posteriorly; dorsal and ventral fins translucent, with olive stippling distributed regularly along length of tail except distally, where stippling diminishes. Spiracle and oral apparatus nearly translucent. Iris gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 33���40 (CJ 5447, 5449, 5464���67, 6557, 6558) are shown in Table 8. Total length varies between 31.4 (Stage 33) and 61.9 (Stage 40) and tail length proportion varies from 58% to 69% until Stage 40. Number of marginal papillae varies among specimens and Gosner stages; variation in number of ventral papillae at lower lip is high (39���66). We documented changes in coloration during ontogenetic development of CJ 1947 (Fig. 28A���C). At Stage 39, the dorsum and flanks are olive-gray with a diffuse pattern of gray paravertebral marks. By Stage 41, dorsum of body and limbs is green with well-defined dark green paravertebral marks; a cream dorsolateral stripe is bordered below by a fine black line; a creamy white stripe is present on face where upper lip will be formed. The flanks are greenish gray. At Stage 46, the dorsum is nearly uniform green; the paraverterbal marks are barely evident. The flanks are brown, and the iris is red. CJ 6813 is similar to the above described but has scattered cream flecks on a green dorsum. Comparisons. Tadpoles of Gastrotheca litonedis may occur in sympatry with those of G. pseustes 2 and G. lojana. Gastrotheca litonedis differs from them by having less pigmentation on dorsum and venter (compare in Fig. 5). It further differs from G. lojana by having a lowest tail dorsal fin.and from G. pseustes by lacking a reticulated pattern on the flanks. Vocalization. A total of nine individuals of Gastrotheca litonedis were recorded from two locations (6 individuals from San Gerardo and 3 from San Fernando, Laguna de Busa) in Azuay Province (Appendix III). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call of G. litonedis is a simple call, composed of a series of short, single-pulsed notes, emitted at regular intervals (Fig. 8 E���H). In our recordings, the calls have between 1 and 21 notes per call. The notes have a mean duration of 0.075 s (SD = 0.025) and a mean inter-note interval of 1.768 s (SD = 0.286). The mean dominant frequency of the call is 1224.0 Hz (SD = 66.415), with mean 90% bandwidth of 1114.7���1308.1 Hz. The fundamental frequency and harmonics are not clearly recognizable. Comparisons. The advertisement call of Gastrotheca litonedis is similar to that of G. cuencana; all the other species of Gastrotheca species in the southern Ecuadorian Andes have complex calls. However, the call of G. litonedis has a higher note rate, longer note duration, shorter inter-note interval, higher dominant frequency, and higher 90% bandwidth frequency compared with G. cuencana (Table 5). Distribution and ecology. Gastrotheca litonedis is known from three localities (ca. Gir��n, San Fernando (Laguna de Busa), San Gerardo) on the eastern slope of the Cordillera Occidental in Azuay Province (Fig. 9). Its elevational range is 2750���2854 m in an area of extent of occurrence of about 10.2 km 2. This nocturnal, semiarboreal species inhabits forest and disturbed areas in the Evergreen Montane Forest of Cordillera Occidental of the Andes (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 611���658 mm and the average annual temperature is 12.2���12.5 ��C (Fick & Hijmans 2017). Most of the specimens collected at Laguna de Busa were found among totora reeds (Schoenoplectus californicus) at the border of the lagoon. Males were calling from totora reeds at approximately 1���2 m above ground. Females were perched on totora reeds approximately 40 cm above ground and along small streams that were close to the lagoon. Some males also were calling from branches of shrubs around the lagoon approximately 2.5 m above ground. During the day a male and a female were basking on leaves of calla lilies (Zantedeschia aethiopica), which were present along irrigation ditches, bordering pastures. At this locality, Gastrotheca litonedis is syntopic with G. pseustes (sensu lato) and G. lojana; all of them have a biphasic mode of development, in which brooding females release tadpoles in ponds, where they complete their development (Figs. 10���11). In all the visits made to this locality (October 2007 and December 2007, August 2008, June 2011), gravid females were found, as well as tadpoles in small streams, temporary ponds, and at the edge of the lagoon. One of the gravid females collected on 9 June 2011 was transported to the laboratory, where it deposited 96 tadpoles. At San Gerardo, most individuals were found on the border of an irrigation ditch in a pasture. Males were calling from the grass at approximately 50 cm above ground. At this locality, G. pseustes (sensu lato) also was found. A brooding female of G. litonedis, showing heliophilic behavior during most of the incubation time, under captive conditions in an outdoor enclosure, is depicted in Figure 10C. Conservation status. IUCN categorizes Gastrotheca litonedis as Endangered (Coloma et al. 2004). However, the current assessment was based on information combining the two species described in this study (G. litonedis and G. cuencana). Thus, we re-categorize this species as Critically Endangered, according with IUCN criteria and sub-criteria B1ab(i,ii,iii,iv). Currently, the known area of occurrence of G. litonedis is extremely small (10.2 km 2), and its area of occupancy is even smaller; although we suspect that its distribution might increase as further searches are conducted. Its populations are in heavily human populated areas. For example, the surroundings of Laguna de Busa have been deforested and modified for human activities (Fig. 11D). The lake contains two introduced, predatory fish (trout and carp) and its surroundings are greatly altered by the introduction of exotic eucalyptus and conifers. Additional threats at this site are unregulated tourisms activities. Comments. In the original description of Gastrotheca litonedis, Duellman and Hillis (1987) included data on nine males and 15 females. As noted by Duellman (2015), of these specimens, only the holotype, KU 202690, is G. litonedis. The other specimens used by Duellman & Hillis (1987) are identified herein as G. cuencana. In addition, the photograph published as the holotype of G. litonedis by Duellman and Hillis (1987:156: Fig. 9) is not the holotype. The frog in the photograph is actually G. lojana (KU 148794, from 2 km SSW of Saraguro, Loja Province, Ecuador). Also, the frog QCAZ 2692 depicted in Duellman (2015: Fig. 11.12B) is G. lojana (not G. litonedis). Genetic sequences generated in this study from tissues of the holotype differ from the sequences of other specimens generated and published by Wiens et al. (2007) (Genbank numbers: DQ679395, DQ679355, DQ679323, DQ679287), which are more similar to sequences of G. cuencana., Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 61-67, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Duellman, W. E. & Hillis, D. M. (1987) Marsupial frogs (Anura: Hylidae: Gastrotheca) of the Ecuadorian Andes: resolution of taxonomic problems and phylogenetic relationships. Herpetologica, 43, 141 - 173.","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp.","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","Coloma, L. A., Ron, S. & Morales, M. (2004) Gastrotheca litonedis. In: IUCN 2013. IUCN Red List of Threatened Species. Fersion 2013.2. Available from: www. iucnredlist. org (accessed 30 October 2018)","Wiens, J. J., Kuczynski, C. A., Duellman, W. E. & Reeder, T. W. (2007) Loss and re-evolution of complex life cycles in marsupial frogs: Does ancestral trait reconstruction mislead? Evolution, 61 (8), 1886 - 1899. https: // doi. org / 10.1111 / j. 1558 - 5646.2007.00159. x"]}

Gastrotheca litonedis Duellman & Hillis 1987 Holotype. KU 202690 (Figs. 10A, 25A), an adult female, from 10 km (by road) northeast of Girón, 2750 m (03° 07' 11.8" S, 79° 06' 28.4'' W), Azuay Province, Ecuador, obtained on 7 March 1984 by William E. Duellman. Referred specimens. (Total 28: 13 males, 15 females). Ecuador: Azuay: CJ 386 ¯9 (females), 401¯4 (females, male), 1404¯8 (males), KU 335388 –9 (male, female), San Fernando, Laguna de Busa, 2834 m (03° 09' 15.8" S, 79° 15' 49.03" W), collected on 9 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara and Henry Grefa; QCAZ 42734 (female), San Fernando, Laguna de Busa, 2834 m (03° 09' 15.8" S, 79° 15' 49.03" W), collected on 27 October 2007 by Sofía Carvajal-Endara, Amaranta Carvajal-Campos and Andrea Carvajal-Endara; QCAZ 42855 (female), 42857¯9 (females), 42861 (female), 42866 (male), 42871 (male), San Fernando, Laguna de Busa, 2834 m (03° 09' 15.8" S, 79° 15' 49.03" W), collected on 18–20 August 2008 by Sofía Carvajal-Endara; QCAZ 49973 ¯4 (female, male), 49976¯8 (males), San Gerardo, 2854 m (03° 08' 00.17" S, 79° 11' 36.78" W), collected on 1 December 2010 by Sofía Carvajal-Endara. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (53.5– 62.4 mm SVL in females, n = 22; 48.9–57.4 mm SVL in males, n = 13) with tibia length 40¯54% SVL, slightly longer than foot; (2) interorbital distance about 1.5 times width of upper eyelid; (3) skin on dorsum smooth, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus distinct, smooth; (7) Fingers I and II about equal in length, width of discs notably wider than digits; (8) fingers unwebbed; (9) webbing between external toes extending to the penultimate subarticular tubercle on Toe IV, to the distal subarticular tubercle on Toe V; (10) in life, dorsum green, brown, or tan with or without dark paravertebral marks or dark flecks; (11) head markings consisting of a pale cream labial stripe and narrow dark brown canthal stripe; (12) pale creamy white dorsolateral stripe present; (13) flanks, anterior surfaces of thighs and groin pale brown or tan slightly molted in some, posterior surfaces of thighs dark brown–gray with pale cream warts in some; (14) venter pale brownish gray; (15) brood pouch single, dorsal. Gastrotheca litonedis most closely resembles five other species in southern Ecuador— G. cuencana, G. plumbea, G. pseustes, G. turnerorum, and G. elicioi. Gastrotheca litonedis differs from all of them by having a pale brownish gray venter; whereas, the venter is uniform creamy white in G. cuencana, cream in G. plumbea, cream with dark flecks or spots in G. pseustes and G. elicioi, and bronze or dark brown in G. turnerorum. Gastrotheca litonedis, like G. plumbea, has smooth skin on the dorsum, whereas it is finely granular in G. elicioi and G. cuencana, weakly areolate to smooth in G. pseustes and areolate in G. turnerorum. A pale supracloacal stripe is present in G. litonedis, G. pseustes, and G. turnerorum, but it is absent in G. cuencana and G. plumbea. Gastrotheca pseustes differs further from G. litonedis by having a complex call structure (long and short pulsed-notes), which is simple in G. litonedis (only short pulsed-notes). The sister species of G. litonedis is the parapatric G. cuencana. Distinctive morphological and coloration features (compare Figs. 26–27, vs 2–4) are (character states for G. cuencana in parentheses): Gastrotheca litonedis is larger than G. cuencana (SVL student’s t-test: t = -6.725, df = 99, p G. litonedis has a higher note rate, longer note duration, shorter inter-note interval, higher dominant frequency, and higher 90% bandwidth frequency compared with G. cuencana. Finally, these two species also differ by having a genetic distance of 1.2 % (in a DNA dataset of 438 bp, 16S gene). Gastrotheca litonedis is syntopic with G. lojana and G. pseustes 2, from which it differs notably by the texture of skin on dorsum, ventral coloration, and call structure. The skin on the dorsum is smooth in G. litonedis, whereas it is finely granular in G. lojana and coarsely granular in G. pseustes 2; furthermore, both species have cream venters with dark flecks, spots, or mottling, and a complex call structure with long and short pulsed-notes (G. lojana call in Fig. 17 H–N). Variation. Morphometric variation of 22 females and 13 males (from Duellman 2015) is summarized in Table 3. Females are larger than males (58.0± 2.4 mm; 53.8± 2.8 mm). The skin on the dorsum is mostly smooth in most specimens; however, in some specimens it is weakly areolate. All adults have a moderate supratympanic fold, which usually extends from the upper part of the tympanum to a point above the insertion of the forelimb. Some individuals have several small rounded tubercles anterolateral to the tympanum. Each dentigerous vomerine process has 2–10 teeth (5.4±1.6, n = 34). Color variation in preservative. In most specimens the dorsum is bluish gray without marks; in a few specimens faint paravertebral marks are present. The narrow canthal stripe, flanks, and posterior surfaces of thighs are dark brown. A creamy white supralabial stripe from the posterior margin of the lip to the insertion of the forelimb is prominent in all specimens. Creamy white dorsolateral, supracloacal and heel stripes are present. On the posterior part of the flanks the coloration varies from small white flecks to mottling on a dark brown ground color. The anterior surfaces of the thighs are usually gray with black flecks or spots. The ventral surfaces are uniform dull gray in most specimens; some have a pale gray venter with evenly distributed dark flecks and spots, and in CJ 1412 the venter is pale gray with black markings. Some specimens have a faint tinge of pale blue on the ventral surfaces of the shanks. Males have a cream nuptial pad on the medial surface of the thumb. Color variation in life. (Figs. 26, 27). The dorsum varies from uniform green (QCAZ 42858), to brown (CJ 404); in some individuals dark paravertebral marks and dark brown flecks are present (QCAZ 49977). The ventral surfaces are pale gray in most specimens; some have evenly distributed dark flecks and spots, and some have a faint bluish tinge on the ventral surfaces (QCAZ 49977). A narrow dark brown canthal stripe is present; a cream supralabial stripe extends from the posterior margin of the lip to the insertion of the forelimb. The tympanum is brown, tan, or olive green. The iris is reddish bronze with dense black reticulations. The posterior surfaces of the thighs are dark brown or cream with orange flecks. A cream dorsolateral stripe is evident in most individuals. The flanks, groin, and anterior surface of the thighs usually are bronze-brown or tan with cream or dark flecks and spots. In some individuals a pink-orange tinge is present in the groin (Fig. 27). The supracloacal and heel stripes are cream or tan, and the ventral surfaces of the shanks have a faint pale blue tinge. Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 37 (CJ 6558), from a series of 79 tadpoles (CJ 6557) born and reared in laboratory conditions, from a mother collected when gravid in the field (CJ 6018) at Laguna de Busa, 2780 m, Azuay Province, Ecuador, by Manuel A. Morales-Mite on 0 6 January 2016 (Fig. 5A). Total length 52.8; body length 19.4 (37% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 12.5, and height at same position 10.3, head width at level of eyes 10.0. Lateral line system present but barely visible, supraorbital and infraorbital lines not evident at level of snout, infraorbital line present at level of the eye, touching the inferior portion of the orbit, and making contact with supraorbital line immediately behind the eye. Postorbital line represented by three stitches arranged in line and forming a 45-degree angle with the body plane. Angular line, anterior oral line and loreal lines not visible; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 4.2; internarial distance 2.8. Eyes dorsally positioned; eye length 1.9, eye width 1.8; interorbital distance 5.2. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 12.9; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; spiracle length 2.7, spiracle transverse width 3.5. Vent tube dextral, opening oriented posteriorly, tube length 2.5, tube transverse width 2.2. Tail length 33.3; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 4.1, width 3.6; caudal fins well developed and proportional, dorsal fin arising abruptly near tail-body junction; dorsal fin height 4.1, ventral fin height 3.7; maximum tail height 9.2; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 5.8. It is surrounded by a uniserial row of marginal papillae, interrupted medially on upper lip. Lower lip papillae alternate in orientation, giving appearance of two rows; upper lip with 23 papillae on right side and 23 papillae on left side; lower lip bearing 57 marginal papillae. Upper jaw sheath medium-sized, forming a smooth arch and finely serrated, transverse width 2.9 (50% of width of oral width) and height 0.3. Lower jaw sheath V- shaped, open and finely serrated, width 2.6, and height 0.7. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 4.5, A2: 4.4, P1 right row 1.65, P1 left row 2.0, P1 gap 0.2, P2: 4.1, P3: 3.95. (See also Fig. 6A of CJ 5292, Stage 35). Color in life. Based on a specimen in Stage 34 from a series (CJ 1947) obtained from San Fernando, Laguna de Busa, Azuay Province, 2834 m, by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa on 9 June 2011 (Fig. 6A). In dorsal view, body olive-gray, head and tail suffused with pink and cream. Flanks cream anteriorly to olivegray posteriorly. In ventral view, cream anteriorly in gular region; followed posteriorly by red gills visible troughout the throat, and olive-gray guts. Caudal musculature pink proximally, gradually becoming gray posteriorly; dorsal and ventral fins translucent, with olive stippling distributed regularly along length of tail except distally, where stippling diminishes. Spiracle and oral apparatus nearly translucent. Iris gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 33–40 (CJ 5447, 5449, 5464–67, 6557, 6558) are shown in Table 8. Total length varies between 31.4 (Stage 33) and 61.9 (Stage 40) and tail length proportion varies from 58% to 69% until Stage 40. Number of marginal papillae varies among specimens and Gosner stages; variation in number of ventral papillae at lower lip is high (39–66). We documented changes in coloration during ontogenetic development of CJ 1947 (Fig. 28A–C). At Stage 39, the dorsum and flanks are olive-gray with a diffuse pattern of gray paravertebral marks. By Stage 41, dorsum of body and limbs is green with well-defined dark green paravertebral marks; a cream dorsolateral stripe is bordered below by a fine black line; a creamy white stripe is present on face where upper lip will be formed. The flanks are greenish gray. At Stage 46, the dorsum is nearly uniform green; the paraverterbal marks are barely evident. The flanks are brown, and the iris is red. CJ 6813 is similar to the above described but has scattered cream flecks on a green dorsum. Comparisons. Tadpoles of Gastrotheca litonedis may occur in sympatry with those of G. pseustes 2 and G. lojana. Gastrotheca litonedis differs from them by having less pigmentation on dorsum and venter (compare in Fig. 5). It further differs from G. lojana by having a lowest tail dorsal fin.and from G. pseustes by lacking a reticulated pattern on the flanks. Vocalization. A total of nine individuals of Gastrotheca litonedis were recorded from two locations (6 individuals from San Gerardo and 3 from San Fernando, Laguna de Busa) in Azuay Province (Appendix III). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call of G. litonedis is a simple call, composed of a series of short, single-pulsed notes, emitted at regular intervals (Fig. 8 E–H). In our recordings, the calls have between 1 and 21 notes per call. The notes have a mean duration of 0.075 s (SD = 0.025) and a mean inter-note interval of 1.768 s (SD = 0.286). The mean dominant frequency of the call is 1224.0 Hz (SD = 66.415), with mean 90% bandwidth of 1114.7–1308.1 Hz. The fundamental frequency and harmonics are not clearly recognizable. Comparisons. The advertisement call of Gastrotheca litonedis is similar to that of G. cuencana; all the other species of Gastrotheca species in the southern Ecuadorian Andes have complex calls. However, the call of G. litonedis has a higher note rate, longer note duration, shorter inter-note interval, higher dominant frequency, and higher 90% bandwidth frequency compared with G. cuencana (Table 5). Distribution and ecology. Gastrotheca litonedis is known from three localities (ca. Girón, San Fernando (Laguna de Busa), San Gerardo) on the eastern slope of the Cordillera Occidental in Azuay Province (Fig. 9). Its elevational range is 2750–2854 m in an area of extent of occurrence of about 10.2 km 2. This nocturnal, semiarboreal species inhabits forest and disturbed areas in the Evergreen Montane Forest of Cordillera Occidental of the Andes (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 611–658 mm and the average annual temperature is 12.2–12.5 °C (Fick & Hijmans 2017). Most of the specimens collected at Laguna de Busa were found among totora reeds (Schoenoplectus californicus) at the border of the lagoon. Males were calling from totora reeds at approximately 1–2 m above ground. Females were perched on totora reeds approximately 40 cm above ground and along small streams that were close to the lagoon. Some males also were calling from branches of shrubs around the lagoon approximately 2.5 m above ground. During the day a male and a female were basking on leaves of calla lilies (Zantedeschia aethiopica), which were present along irrigation ditches, bordering pastures. At this locality, Gastrotheca litonedis is syntopic with G. pseustes (sensu lato) and G. lojana; all of them have a biphasic mode of development, in which brooding females release tadpoles in ponds, where they complete their development (Figs. 10–11). In all the visits made to this locality (October 2007 and December 2007, August 2008, June 2011), gravid females were found, as well as tadpoles in small streams, temporary ponds, and at the edge of the lagoon. One of the gravid females collected on 9 June 2011 was transported to the laboratory, where it deposited 96 tadpoles. At San Gerardo, most individuals were found on the border of an irrigation ditch in a pasture. Males were calling from the grass at approximately 50 cm above ground. At this locality, G. pseustes (sensu lato) also was found. A brooding female of G. litonedis, showing heliophilic behavior during most of the incubation time, under captive conditions in an outdoor enclosure, is depicted in Figure 10C. Conservation status. IUCN categorizes Gastrotheca litonedis as Endangered (Coloma et al. 2004). However, the current assessment was based on information combining the two species described in this study (G. litonedis and G. cuencana). Thus, we re-categorize this species as Critically Endangered, according with IUCN criteria and sub-criteria B1ab(i,ii,iii,iv). Currently, the known area of occurrence of G. litonedis is extremely small (10.2 km 2), and its area of occupancy is even smaller; although we suspect that its distribution might increase as further searches are conducted. Its populations are in heavily human populated areas. For example, the surroundings of Laguna de Busa have been deforested and modified for human activities (Fig. 11D). The lake contains two introduced, predatory fish (trout and carp) and its surroundings are greatly altered by the introduction of exotic eucalyptus and conifers. Additional threats at this site are unregulated tourisms activities. Comments. In the original description of Gastrotheca litonedis, Duellman and Hillis (1987) included data on nine males and 15 females. As noted by Duellman (2015), of these specimens, only the holotype, KU 202690, is G. litonedis. The other specimens used by Duellman & Hillis (1987) are identified herein as G. cuencana. In addition, the photograph published as the holotype of G. litonedis by Duellman and Hillis (1987:156: Fig. 9) is not the holotype. The frog in the photograph is actually G. lojana (KU 148794, from 2 km SSW of Saraguro, Loja Province, Ecuador). Also, the frog QCAZ 2692 depicted in Duellman (2015: Fig. 11.12B) is G. lojana (not G. litonedis). Genetic sequences generated in this study from tissues of the holotype differ from the sequences of other specimens generated and published by Wiens et al. (2007) (Genbank numbers: DQ679395, DQ679355, DQ679323, DQ679287), which are more similar to sequences of G. cuencana.

Gastrotheca elicioi sp. nov. urn:lsid:zoobank.org:act: 3D26B088-B527-4362-94E8-647FF26C8419 Holotype. CJ 1402 (Fig. 12), an adult male (collected as tadpole in the field and captive raised), from Cajanuma, entrance to Parque Nacional Podocarpus in the Loja-Vilcabamba road, 2456 m (04° 05' 25.51" S, 79° 12' 09.97" W), Loja Province, Ecuador, one of a series obtained on 14 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa. Paratypes. (Total 18: 11 males, 3 females, 4 juveniles). Ecuador: Loja: CJ 413–4 (juvenile, male), 1398–401 (four males), 1941 (male) collected with the holotype; KU 142603 –5 (female, subadults), from 5.5 km W Loja, 2330 m (04° 00' 48.99" S, 79° 13' 50.99" W), on 17 July 1971 by William E. Duellman and Linda Trueb; KU 142607 –8 (subadult, female), 148549–51 (males, female), from 5.5 Km W Loja, 2330 m (04° 00' 48.99" S, 79° 13' 50.99" W), on 23 July 1971 by William E. Duellman and Linda Trueb; KU 202688 (male), from 5.2 km W Loja, 2310 m (03° 58' 58.73" S, 79° 16' 02.21" W), on 10 March 1984 by William E. Duellman; KU 217511 –2 (males), from 6.8 km E Loja ca. Loja-Zamora line (03° 59' 19" S, 79° 09' 21.99" W), on 9 January 1990 by David Kizirian, John J. Wiens, and Luis A. Coloma. Referred specimens. Ecuador: Loja: QCAZ 22370 (male), from Zamora Huayco, Loja, 3018 m (04°0 5' 49.98" S, 79° 10' 02.5" W), on 2 December 2002 by Diego Almeida-Reinoso; QCAZ 46319–20 (males), from Zamora Huayco, Loja, 3018 m (04° 05' 49.98" S, 79° 10' 02.5" W), on 5 December 2009 by Diego Almeida- Reinoso and Samael D. Padilla; CJ 1841–902 (62 juveniles), from Loja, Barrio La Palmera, Parroquia Sucre, on 22–25 April 2013 by Elicio E. Tapia. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (up to 67.5 mm SVL in female, 46.4–67.8 mm SVL in males, n = 13) with tibia length 42¯50% SVL, slightly longer than foot; (2) interorbital distance larger than the width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth to slightly granular; (7) Fingers I and II about equal in length, width of discs about twice width of the digits proximal to discs; (8) fingers unwebbed; (9) webbing between external toes extending nearly to antepenultimate subarticular tubercle on Toe IV and proximal to penultimate subarticular tubercle on Toe V; (10) in life, dorsum green, tan, brown, or reddish-brown with or without dark paravertebral marks; (11) head markings consisting of a pale labial stripe on the posterior margin of the lip, a black canthal stripe and a dark triangular interorbital mark in some individuals; (12) a fragmented, narrow, pale dorsolateral stripe present; (13) flanks bronze-brown, tan, or green with or without small pale spots; groin and anterior and posterior surfaces of thighs lightly mottled with tinges of green or blue; (14) venter cream with small brown flecks or spots; (15) brood pouch single, dorsal. In comparison with similar species Gastrotheca elicioi is most like G. lojana, G. cuencana, G. litonedis, G. turnerorum, and G. pseustes in Ecuador and G. monticola in Peru. Gastrotheca elicioi differs from G. lojana, G. cuencana, and G. litonedis in color pattern as follows (compare these species in Fig. 10). Gastrotheca elicioi and G. litonedis have distinct dark canthal stripes, which are absent in G. lojana and inconspicuous or absent in G. cuencana. The groin and anterior and posterior surfaces of the thighs are slightly mottled in G. elicioi, whereas in G. lojana they are heavily mottled; these surface are translucent cream without marks in G. cuencana and usually brown in G. litonedis. If any marks are present on the dorsum in G. elicioi, they usually consist of a triangular interorbital mark, which is connected or not with two narrow and curved paravertebral marks. In contrast, in G. lojana the interorbital mark is a transverse bar usually connected with two broad paravertebral marks; in G. cuencana and G. litonedis an interorbital mark is absent. Gastrotheca lojana and G. cuencana have a conspicuous, elevated row of dorsolateral warts, whereas they are barely raised in G. elicioi and G. litonedis. Additionally, in G. elicioi the dark bars on the limbs, when present, are shorter, thinner and less defined than in G. lojana, whereas in G. cuencana and in some G. litonedis the bars are replaced by irregular blotches. Gastrotheca elicioi has a cream venter with dark flecks or marks, whereas the venter is uniform creamy white in G. cuencana and pale brownish gray in G. litonedis. Gastrotheca cuencana also differs from G. elicioi by having cream dorsal surfaces of the fingers, whereas they are brown, green, or tan in G. elicoi. Gastrotheca turnerorum and G. pseustes differ from G. elicioi (compare these species in Fig. 10) by having the skin on dorsum areolate and weakly areolate respectively, whereas in G. elicioi the skin on dorsum is finely granular. Also, they differ by lacking an interorbital mark, and bars on limbs, which can be present G. elicioi. Gastrotheca pseustes is sympatric with G. elicioi, from which it differs notably by having smaller digital discs (larger in G. elicioi) and lacking a pale dorsolateral and supracloacal stripes, which are fragmented in G. elicioi. Gastrotheca monticola is the sister species of G. elicioi and their minimum genetic divergence is 3.0% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca monticola differs from G. elicioi by having the axilla and groin, concealed surfaces of the thighs, and the dorsal surfaces of the feet green with well-defined black spots on a green or tan background, whereas in G. elicioi the groin and anterior and posterior surfaces of thighs are lightly mottled with tinges of green or blue. Finally, the advertisement call of G. elicioi is unique amongst the calls of Gastrotheca in southern Ecuador, in that the long, pulsed notes are produced after the short ones, whereas the other species tend to produce the long, pulsated notes before the short ones. Description of the holotype. An adult male (Fig. 12) collected as a tadpole and raised in the laboratory; body moderately robust; SVL 46.4 mm; head wider that long; snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis round in section; loreal region concave, lips rounded; top of head flat; interorbital distance 142% of width of upper eyelid; internarial area elevated; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye in a distance approximately one and a half the diameter of tympanum; tympanic annulus and membrane smooth; supratympanic fold moderately weak, extending from corner of eye to tympanum and on to insertion of forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing three teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 31% of SVL), unwebbed, with distinct narrow lateral fringes; discs much wider than digits, slightly truncated, width of disc on Finger III greater that diameter of tympanum; relative lengths of fingers I=II Skin on dorsum finely granular; skin on flanks coarsely granular; skin on throat, venter surfaces of thighs, and arms granular; skin on belly areolate; skin on venter surfaces of shanks smooth; numerous small tubercles lateral to cloacal opening. Vocal sac single, median, subgular. Vocal slits present at posterior lingual margins of mandibles. Tongue broad, suboval, notched posteriorly, fully attached to mouth floor. Coloration in life (See also under Comments). The dorsum is uniform brown becoming yellowish brown on the flanks. The dorsal surfaces of limbs have an olive-green tinge. A bronze labial stripe is present at the margin of the lip and continues to the insertion of the forelimb. A bronze inconspicuous and highly fragmented dorsolateral stripe is present; a black canthal stripe is present. The flanks are bronze-brown; the groin and anterior and posterior surfaces of the thighs are pale green with white and black spots. The venter is brownish gray with black and white marks; the gular region is gray dark. The iris is copper colored with a few black reticulations. Coloration in preservative. The dorsum is gray without markings. A narrow pale, fragmented dorsolateral stripe is present. The dorsal surfaces of the limbs are tan with irregular gray blotches. A short pale labial stripe extends from the margin of the lip to the level of the insertion of forelimb; a dark canthal stripe is present. The flanks are dark gray anteriorly and pale gray posteriorly; the groin and anterior and posterior surfaces of the thighs are pale gray with black marks. The venter is cream with dark gray spots; the gular region is gray. Measurements (in mm). SVL: 46.4, TIBL: 19.8, FL: 20.0, HL: 14.1, HW: 16.2, IOD: 4.9, EW: 4.6 IND: 3.3, ED: 5.6, EN: 4.1, TD: 2.2, FFL: 7.9, TFL: 14.6, TFD: 2.6. Variation. Morphometric variation of one female and 13 males is summarized in Table 2. The female is larger than most males (67.5 mm; 56.0± 8.1 mm). The skin on the dorsum is finely granular. The tympanic annulus usually is smooth but is slightly granular in some individuals. Each dentigerous vomerine process has 6–8 teeth (6.3±1.5, n = 3). Color variation in preservative. Preserved specimens have a bluish gray, pale gray, or brownish gray dorsum. Interorbital and paravertebral marks usually are present but the paravertebral marks may be absent. The dark gray interorbital mark, when present, is triangular in most specimens, but it is divided into two blotches in some specimens. A dark canthal stripe is present in all specimens. A short, pale labial stripe is inconspicuous in most specimens. The flanks are gray, darkest anteriorly, with or without white and dark flecks. The groin and anterior surfaces of thighs are pale gray with slight mottling in some; the posterior surfaces of the thighs are pale gray with black flecks and marks usually only distally. In some specimens, the dorsal surfaces of arms and limbs present short dark gray bars. The venter is white with evenly, or densely distributed dark marks. Color variation in life. (Figs. 10H, 10I, 13–14). In living individuals the dorsum is uniform green (CJ 1843), reddish-brown (CJ 1400), or tan (CJ 1842), with or without contrasting dark gray paravertebral marks. When present, these paravertebral marks usually are narrow, curved, and fragmented; these marks are adherent at the scapular level and may be connected with the interorbital mark. A short cream labial stripe is present on the margin of the lip; the stripe continues posteriorly to the insertion of the forelimb. A dark brown canthal stripe is always present, but it is inconspicuous in some individuals with a dark ground color. The tympanum is brown, tan, or olive green. The iris is usually copper almost without black reticulations. A bronze highly fragmented dorsolateral stripe is present in most individuals. The flanks are brown, bronze-brown, or green. The groin has bluish coloration with slight mottling. The posterior surfaces of the thighs usually are pale green with black flecks and some specimens show black fringes. When present, supracloacal and heel stripes are cream and fragmented. The ventral surfaces usually are cream with dark marks, but some specimens lack dark marks. The gular surface varies from brownish gray to white with black spots. Duellman (1974) provided detailed descriptions of coloration in life of four adult males (KU 148549 –51 and 142603, under the name Gastrotheca lojana). Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36 (CJ 4311), from a series of 50 tadpoles (CJ 4311) obtained from a pond at Puntzará Alto, near Loja city, 2311 m, Province of Loja, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016. Total length 51.4; body length 19.4 (38% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 11.9, and height at same position 9.9; head width at level of eyes 11.1. Lateral line system present but barely visible, supraorbital and not evident at level of snout, infraorbital line present at level of the eye, touching the inferior portion of the orbit, and making contact with supraorbital line immediately behind the eye. Inferior oral line visible at the eye level, where it contacts angular line, which descends from eye level. Supraorbital line represented for scattered stiches, which does not make contact with the orbit. Posorbital line forming a circle of stitches, just behind the eye. Anterior oral line and loreal lines not visible; dorsal body and middle bodylines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 3.5; internarial distance 2.9. Eye directed dorsally; eye length 2, eye width 1.8; interorbital distance 5. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 13.0; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; spiracle length 2.9, tube transverse width 2.9. Vent tube dextral, opening oriented posteriorly, tube length 3.4, vent tube transverse width 3. Tail length 32.4; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 4.4, width 3.4; caudal fins well developed and proportional, arising abruptly near tail-body junction and forming a notorious hump, which makes and arc in the body plane. Dorsal fin height 4.18, ventral fin height 3.7; maximum height of tail 11.9; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 4.9. It is surrounded by an uniserial row of marginal papillae, interrupted medially on upper lip; lower lip papillae alternate in orientation, giving appearance of two rows. Upper lip with 23 papillae on right side and 20 papillae on left side; lower lip bearing 52 marginal papillae; upper jaw sheath medium-sized, forming a smooth arch and finely serrated, transverse width 2.6 (53% of oral disc width) height 0.4; lower jaw sheath V- shaped, open and finely serrated, width 4.1, height 1.3. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 4.15, A2: 4.0, P1 right row 1.75, P1 left row 1.75, P1 gap 0.1, P2: 3.75, P3: 3.70. (Fig. 6D). Color in life. Based on a specimen (CJ 4312a) in Stage 37 from a series (CJ 4312) obtained at Loja (neigborhood Puntzará Grande), Loja Province, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016 (Fig. 5D). In dorsal and ventral views, body dark brown. Snout and flanks dark brown; guts gray; red gills visible through the throat. Venter gray. Caudal musculature reddish brown with brown stippling, distal portion lacking pigments; dorsal and ventral fins gray with minute cream stippling. Vent tube translucent. Iris reddish-gold. Variation. Variation of 28 meristic characters of tadpoles in Stages 31–40 (CJ 4306, 4310–13) are shown in Table 6. Total length varies between 15.8 (Stage 31) and 73.2 (Stage 39) and tail length proportion varies from 57% to 70% until Stage 38. Number of marginal papillae varies among specimens and Gosner stages; variation in number of ventral papillae at lower lip is high (43–64). Based on specimens CJ 1950–2 from a series obtained at Loja (neigborhoods La Palmera and Puntzará Grande), Loja Province, Ecuador, by Elicio E. Tapia on 22–25 April 2013 (Fig. 15). We documented changes in coloration during ontogenetic development of one, mostly brown individual (CJ 1950) (Figs. 15 A–B). At Stage 40, the dorsum and flanks were cream and brown with a diffuse pattern of brown-gray paravertebral marks and a dark gray stripe bordering the canthus and body dorsolaterally, extending to about level of midbody, bordered dorsally by a diffuse cream area. The caudal musculature is pink proximally, with a gray suffusion distally. The caudal fins are pale gray. The iris is pale red. By Stage 46, the markings on the dorsum of body and limbs are diffuse gray and green on a dark brown background; the fingers and toes are yellowish cream dorsally. The flanks have a wide, brown-gray band. There are well-defined creamy white dorsolateral and labial stripes and supracloacal white marks. Color variation in six additional metamorphs (CJ 1951–2, CJ 4313) from Loja and Loja–Abra de Zamora, in Stage 46 is depicted in Figure 16. They vary from plain dark brown to plain green, and have a complete supracloacal stripe. …….continued on the next page …….continued on the next page Comparisons. Tadpoles of Gastrotheca elicioi may occur in sympatry with those of G. lojana, G. psychrophila, G. pseustes, and G. turnerorum in the Loja-Abra de Zamora region. Gastrotheca elicioi differs from all of them by having a dorsal gray-pigmented fin that abruptly arises from the body, whereas is nearly translucent and arises gradually in the other species (compare in Fig 5). The tadpole described as G. psychrophila by Duellman (2015) is G. elicioi. It has a similar dorsal fin and fit well our description of G. elicioi tadpoles. Nonetheless, until tadpoles of G. psychrophila are described and unequivocally assigned to its species, some doubts will remain. Vocalization. Seven individuals of Gastrotheca elicioi were recorded from three locations in Loja Province (one individual from the old Loja-Catamayo road, three from Loja, Parque Universitario

Gastrotheca pseustes Duellman & Hillis 1987 Gastrotheca pseustes was described from 7.1 km by road north of San Lucas, 2940 m (03�� 41' S, 79�� 15' W), Loja Province, Ecuador, in the southern Andean Cordillera. The type locality is located about midway between San Lucas and Saraguro. Current assignment of specimens to this taxon is constrained because: (1) there is a high phenotypic similarity between G. pseustes and G. lateonota from the Cordillera de Huancabamba in Peru (compare both species in Fig. 10 with Duellman 2015: Fig. 12.21); (2). There is no genetic information for topotypic G. lateonota, thereby precluding comparisons with specimens from southern Ecuador; (3) according to Duellman (2015), variation in morphometrics, structural characters, and coloration within G. pseustes presents a geographic mosaic, except for the southernmost population, Saraguro. The frogs from Saraguro are somewhat more distinctive than the others in being larger in size, and in having broader heads and some consistent characteristics of color pattern, including dark flanks with pale spots and uniformly dark posterior surfaces of the thighs; and (4) new molecular phylogeographic data (not shown) suggest that there may be at least two species in what is now recognized as Gastrotheca pseustes. While awaiting ongoing analyses of molecular data and a future report with a more detailed taxonomic and phylogeographic revision of Gastrotheca pseustes, we provide additional data of specimens (adults, tadpoles and metamorphs) from the type locality and surrounding areas in Loja and El Oro provinces. The recognition of Gastrotheca lateonota in southern Ecuador (from El Oro and Loja provinces) by Blackburn & Duellman (2013), Duellman et al. (2014), Y����ez-Mu��oz et al. (2014), and Duellman (2015) was based on specimens identified as that species by Carvajal-Endara and Coloma. Subsequent examination of the specimens from Chilla, El Oro Province, reveals that they do not differ from the holotype (KU 203443, Fig. 10G) and additional specimens (Figs. 10 D���F) of G. pseustes, from 3.7 km S Saraguro, 2800 m. Thus, we do not recognize G. lateonota to occur in Ecuador, and treat the specimens from Chilla as part of a series of populations we consider G. pseustes complex (G. pseustes 1, G. pseustes 2, Fig. 33), occurring from the Equator south to high elevations of Podocarpus National Park in southern Ecuador (see also map in Duellman 2015: Figure 12.47). Conceivably, molecular data from topotypic G. lateonota from 31.5 km [by road] E Canchaque, 2770 m, Cordillera de Huancabamba, Regi��n de Piura, Peru, (Fig. 33), and additional populations from northern Peru would resolve by (1) placement of non-type populations in either G. pseustes or G. lateonota, or (2) placement of G. lateonota as a junior synonym of G. pseustes. Castroviejo-Fisher et al. (2015) included in a phylogenetic analysis a specimen UINHM 94580 from San Rafael, Azuay, Ecuador under the name Gastrotheca riobambae, a species that does not occur in southern Ecuador. Their analysis places it in the G. pseustes complex. We also found a locality named San Rafael in Azuay Province, which is located between Cuenca and Cumbe, in Parroquia Tarqui at about 2700 m asl (not in the lowlands as was stated by Castroviejo-Fisher et al., 2015). This locality is well within the altitudinal and latitudinal range of G. pseustes 2. Herein, we provide meristic data from specimens of Gastrotheca pseustes 1 and 2 (Table 3), images depicting intrapopulation color pattern variation of metamorphs (Fig. 34), of adults of G. pseustes 1 from the type locality (Fig. 10G) and from Chillacocha (Figs. 10 D���F). Data for additional specimens of G. pseustes 1 are: QCAZ 45121, 45124 ��� 5, adult males, and QCAZ 45123 adult female from Chillacocha, ~ 8 km SW Chilla, El Oro Province (03�� 30' 39.1" S, 79�� 36' 52.92" W; 3163 m) collected on 18 August 2009. The frogs were on leaves of Gunnera sp. (Gunneraceae) near a small stream and adjoining marshland about 10 x 5 m. The water temperature was 9.6�� C. Two individuals���CJ 201 adult female and KU 335386 adult male���are from Chillacocha, ~ 8 km SW Chilla, El Oro Province (03�� 30' 15.37" S, 79�� 37' 01.7" W; 3250 m) collected on 10 June 2011. They were in amplexus on a leaf of Gunnera sp. (Gunneraceae) approximately 90 cm above the ground and a small stream. Tadpoles were collected from a puddle next to the road in the vicinity of Chilla, El Oro Province (03�� 27' 29.41" S, 79�� 34' 52.07" W; 2452 m) on 11 June 2011; one of these was raised to be an adult male (CJ 399). Tadpoles were collected from a puddle next to the road at Manu, Loja Province (03�� 33' 17.03" S, 79�� 22' 6.02" W; 2876 m) on 11 June 2011; one of these was raised to be an adult female (CJ 400). Tadpoles were collected from the entrance of Parque Nacional Podocarpus, Loja Province (04�� 05' 25.51" S, 79�� 12' 09.97" W; 2456 m) on 14 June 2011; one of these was raised to be an adult female (KU 335387). Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36, from a series (CJ 1949) obtained from a pond at Chillacocha, 8 km SW Chilla, 3250 m, El Oro Province, Ecuador, by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa on 10 June 2011 (Fig. 5E). Total length 58.2; body length 22.7 (39% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat concave in lateral profile, sloping from anterior margin of snout to belly; body width at level of spiracle 15.3, and height at same position 12.9; head width at level of eyes 11.7. Lateral-line system present but barely visible, supraorbital and infraorbital lines both originating at tip of snout, running parallel to the eye and making contact immediately behind the eye; angular line descending vertically from just posterior of eye to throat; it dorsally contacts with post infraorbital line; post-supraocular present in form of a few stitches, anterior oral line descending vertically from oral disc level and behind nares level to throat, making a curve that parallels infraorbital line, forming a circuit continuous with angular and loreal lines; ventral line surrounds dorsally the spiracle; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 3.9; internarial distance 3.6. Eye positioned and directed dorsolaterally, eye length 2.9, eye width 2.3; interorbital distance 7.1. Spiracle sinistral, located at midbody, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 15.6; end of spiracle rounded, attached to body wall, inner wall of spiracle not evident; tube length 2.7, tube transverse width 3.0. Vent tube dextral, opening directed posteriorly, tube length 2.9, tube transverse width 2.8. Tail length 35.0, caudal musculature robust, narrowing gradually until tail terminus; caudal muscle height 4.5, caudal muscle width 4.3; caudal fins well developed and proportional, dorsal fin originating near tail-body junction, forming low hump; dorsal fin height 4.2, ventral fin height 3.8; maximum height of tail 12.3; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, anteriorly reaching level of tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 5.9, surrounded by an uniserial row of small, marginal papillae, interrupted medially in upper lip; lower lip papillae alternating in orientation in and out, giving appearance of two rows; upper lip with 17 papillae on right side and 16 papillae on left side; lower lip with 52 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, with lateral processes, height 0.56, transverse width 3.7 (63% of oral disc width); lower jaw sheath V- shaped, finely serrated, width 2.9, height 0.68. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.2, A2: 4.5, P1 right row 1.9, P1 left row 2.0, P1 gap 0.3, P2: 4.1, P3: 3.7. (Fig. 6F). Color in preservative. Dorsum gray with darker gray areas on flanks, above eye and on throat; margin of snout paler than adjacent areas. Caudal musculature and fins with a profusion of medium-sized dots that are most dense on upper part of the musculature; fins otherwise translucent. Venter cream, speckled with white, guts not exposed; eyes lavender gray with irregular white markings, oral apparatus translucent. Color in life. In dorsal view, body tan with black flecks. Flanks reticulated with black and tan; areas around the eyes and snout paler tan. Venter cream with black markings; throat translucent with small cream flecks; reddish gills evident. Caudal muscles reddish-pink, more evident on proximal half; myomeres barely visible; caudal muscles and fins having medium-sized cream dots, most dense on upper side of caudal muscles and near tail-body junction, forming a nearly conitnuous stripe along dorsolateral caudal musculature; otherwise caudal fins translucent. Legs cream with black markings on toes. Oral apparatus pale cream. Iris copper-yellow, with black reticulations. ......continued on the next page Variation. Variation of 28 meristic characters of tadpoles in Stages 36���42 (CJ 1949) are shown in Table 10. Total length varies between 55.0 (Stage 40) and 58.2 (Stage 36) and tail length proportion varies from 60% to 70% until Stage 42. Number of marginal papillae varies among specimens and Gosner stages; number of lower lip papillae is high (43���52). We documented changes in coloration during ontogenetic development of one, mostly pale brown individual (CJ 1953) (Figs. 5E, 35). At Stage 41, the dorsum and flanks are yellowish-brown with a poorly defined pattern of brown-gray, elongated paravertebral marks and few blotches on dorsum of hind limbs. A diffuse brown-gray stripe borders the canthal and dorsolateral body, and is bordered above by a creamy-brown stripe. The iris has a reddishgold suffusion. By Stage 45, markings on the dorsum and flanks are better defined with brown markings darkest peripherally; scattered green ill-defined flecks are present on dorsum of head, limbs, and canthus rostralis; tip of fingers are yellowish-cream. At Stage 46, paravertebral green-brown markings on dorsum of body and green flecks on limbs are contrasting to surrounding light brown areas; flanks have a dark brown stripe. Color variation in 6 metamorphs (CJ 1949, 1953���56, QCAZ 45126) in Stage 46 is depicted in Figure 34. They vary from dark brown to green with either well-defined paravertebral marks to a uniformly colored dorsum. Comparisons. Tadpoles of Gastrotheca pseustes sensu stricto (in Loja and El Oro provinces) occur in sympatry with those of G. elicioi in the Saraguro region and with G. elicioi, G. lojana, G. psychrophila, and G. turnerorum in the Loja-Abra de Zamora region. Gastrotheca pseustes differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body; from G. lojana by having a reticulated pattern on flanks, and from G. turnerorum by having a less rounded tail terminus (compare in Fig. 5). For G. psychrophila, see remarks under G. elicioi tadpole account. Vocalization. Four individuals of Gastrotheca pseustes were recorded from three locations in Loja Province (2 individuals from Bosque Washapamba, one from V��a Urdaneta-Tutupali, and 1 from Cerro de Arcos; see Appendix III). Because of the taxonomic problems related to the G. pseuste s complex we decided to use for the present analysis only the recordings obtained from populations situated nearby the type locality, south of the Jubones- Gir��n river valley in Loja Province (e.g. G. pseustes 1). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call of G. pseustes is a complex call, composed of one to five long pulsed notes and followed (or not) by one to six short, single-pulsed notes (Fig. 23 H���N). The long note had a mean duration of 0.878 s (SD = 0.188) and consisted on average of 32.72 (SD = 6.867) distinct pulses, partly fused, without silent intervals (amplitude modulation close but less than 100%). The amplitude of the long note increases gradually towards the end of the note after which it decreases a little by the end. The short notes had a mean duration of 0.080 s (SD = 0.046) and the inter-note interval is on average of 0.795 s (SD = 0.502). The mean dominant frequency of the call is 1359.0 Hz (SD = 37.981), with a mean 90% bandwidth of 1043.2���1483.4 Hz. The fundamental frequency is usually recognizable; when visible, 6 to 7 harmonics are distinguishable. Comparisons: The advertisement call of Gastrotheca pseustes is most similar to that of G. yacuri, but G. pseustes has a much longer call duration, longer long notes duration, longer inter-note interval a lower dominant frequency and a lower 90% bandwidth frequency compared with the call of G. yacuri (Table 5). Also, G. pseustes usually emits 4���6 short notes compared with the only two short notes that G. yacuri emits. The call of G. pseustes can be easily distinguished from that of G. lojana and G. testudinea by the amplitude modulation of the longer note, much longer call duration, lower short note rate, longer long notes duration, longer inter-note interval, larger number of pulses, higher pulse rate, a higher dominant frequency, and higher 90% bandwidth frequency. Also, G. pseustes emits a larger number of long and short notes compared with G. lojana and G. testudinea (Table 5)., Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 79-85, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Duellman, W. E. & Hillis, D. M. (1987) Marsupial frogs (Anura: Hylidae: Gastrotheca) of the Ecuadorian Andes: resolution of taxonomic problems and phylogenetic relationships. Herpetologica, 43, 141 - 173.","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp.","Blackburn, D. C. & Duellman, W. E. (2013) Brazilian marsupial frogs are diphyletic (Anura: Hemiphractidae: Gastrotheca). Molecular Phylogenetics and Evolution, 68 (3), 709 - 714. https: // doi. org / 10.1016 / j. ympev. 2013.04.021","Duellman, W. E., Barley, A. J. & Venegas, P. J. (2014) Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru. Zootaxa, 3768 (2), 159 - 177. https: // doi. org / 10.11646 / zootaxa. 3768.2.4","Yanez-Munoz, M. H., Sanchez L, J. C., Lopez, K., Rea S, E., Ramos, P. A. M., Oyagata, L. A. & Paul, C. (2014) Expansion of the distributional range of some species of amphibians and reptiles in southwestern Ecuador. Ampliaciones del rango de distribucion de algunas especies de anfibios y reptiles en el suroccidente de Ecuador. Avances en Ciencias e Ingenierias, 6 (1), B 2 - B 5.","Castroviejo-Fisher, S., Padial, J., De La Riva, I., Pombal Jr., J., Da Silva, H., Rojas-Runjaic, F., Medina-Mendez, E. & Frost, D. (2015) Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development. Zootaxa, 4004 (1), 1 - 75. https: // doi. org / 10.11646 / zootaxa. 4004.1.1","Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661"]}

Gastrotheca turnerorum sp. nov. urn:lsid:zoobank.org:act: B72F71A3-671F-4F89-91D5-2410C1D0B6DF Holotype. CJ 393 (Fig. 18), an adult female (collected as tadpole and raised in captivity), from Laguna Negra de Jimbura, Parque Nacional Yacuri, Amaluza, 3406 m (04�� 42' 44.24" S, 79�� 25' 42.38" W), Loja Province, Ecuador, one of a series collected on 12 June 2011 by Elicio E. Tapia, Sof��a Carvajal-Endara and Henry Grefa. Paratypes. (Total 9: 6 males, 2 females, 1 juvenile). Ecuador: Loja: CJ 394���5 (female, male), 415���7 (males, juvenile), 1386 (female), and KU 335390 (male), collected with the holotype. CJ 7823 (male, collected as tadpole and captive raised), from Parque Nacional Yacuri, Amaluza, 3331 m (04�� 43' 25.22" S, 79�� 26' 15.47" W), Loja Province, Ecuador, one of a series collected on 7 July 2016 by Dan Cog��lniceanu, Diana Sz��kely and Paul Sz��kely. Zamora Chinchipe: MUTPL 221, male from Reserva Tapichalaca, 3073 m (04�� 28' 55.58" S, 79�� 09' 29.76" W), collected on 9 December 2014 by Diego Armijos-Ojeda. Referred specimens. Ecuador: Zamora Chinchipe: QCAZ 9575 (captive raised tadpoles and juveniles) from Parque Nacional Podocarpus, Lagunas del Compadre, 3205 m (04�� 10' 29.24" S, 79�� 06' 59.54" W), on 1 December 1994 by Jenny Rudston; QCAZ 47299 (tadpole), QCAZ (sc 29154) (female) from Parque Nacional Podocarpus, Lagunas del Compadre, 3205 m (04�� 10' 29.24" S, 79�� 06' 59.54" W), on 20 October 2009 by Elicio E. Tapia. Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (54.0��� 58.2 mm SVL in females, n = 3; 50.0��� 55.6 mm SVL in males, n = 4), with tibia length 37��42% SVL, shorter than foot; (2) interorbital distance slightly greater than width of upper eyelid; (3) skin on dorsum areolate, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth; (7) Finger I slightly shorter than Finger II, width of discs much wider than digits; (8) fingers unwebbed; (9) foot webbing between external toes extending to antepenultimate subarticular tubercle on Toe IV, to penultimate subarticular tubercle on Toe V; (10) in life, dorsum uniform green without dark paravertebral marks; (11) head markings consisting of pale labial and canthal stripes formed by a series of small bronze dotes; (12) dorsolateral stripe present, also consisting in a series of small bronze dotes; (13) flanks brown and groin green both with or without a few cream spots; anterior surfaces of thighs dark brown, posterior surfaces of thighs bluish brown with numerous cream tubercles proximal to the vent; (14) venter dark brown with uniformly distributed cream spots; gular region dark brown; (15) brood pouch single, dorsal. Gastrotheca turnerorum most closely resembles three other species in southern Ecuador, G. pseustes, G. elicioi, and G. litonedis. Gastrotheca turnerorum differs from all of them by having a different skin texture on the dorsum, and a distinctive color pattern (compare Figs. 10J, 19���20 vs 10A���I, K, L). The dorsal skin is areolate in G. turnerorum, whereas it is weakly areolate or smooth in G. pseustes, finely granular in G. elicioi, and smooth in G. litonedis. In G. turnerorum the venter is dark brown with uniformly distributed cream spots, whereas it is cream with dark marks in G. pseustes and G. elicioi and pale brown-gray in G. litonedis. The labial stripe consists of a series of small bronze dots in G. turnerorum, and mostly uniform cream in G. pseustes, G. elicioi, and G. litonedis. Gastrotheca pseustes and G. elicioi have a dark brown canthal stripe, whereas G. turnerorum has a canthal stripe formed by a series of small bronze dots. Gastrotheca turnerorum is the sister species of G. aguaruna + G. yacuri, with a genetic distance of at least 2.85 % (in a DNA dataset of 438 bp, 16S gene). The Peruvian G. aguaruna differs from G. turnerorum by having a weakly granular to smooth dorsal skin (areolate in G. turnerorum), Fingers I and II equal in length (Finger I shorter in G. turnerorum), and a cream venter with dark fleck or spots (venter dark brown with uniformly distributed cream spots in G. turnerorum). Description of the holotype. An adult female (Fig. 18) that was collected as a tadpole and raised in the laboratory; body moderately robust; SVL 54.0 mm; head wider that long; snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis round in section; loreal region concave; lips rounded; top of head flat; interorbital distance 89% of width of upper eyelid; internarial area flat; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye by distance about equal to diameter of tympanum; tympanic annulus barely evident;, supratympanic fold moderately weak, extending from behind the tympanum to the insertion of the forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing five teeth. Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 34% of SVL); fingers unwebbed; discs large and rounded, width of disc of Finger III greater that diameter of tympanum; relative lengths of fingers IColoration in life. The dorsal surfaces of the head, body, and limbs are green; the inner and outer surfaces of the forearm, thighs, and shanks are brown (Fig. 18). Head markings are canthal and labial stripes that consist of a series of small bronze dots. The labial stripe continues to the insertion of the forelimb. The dorsolateral stripe also consists of a series of small bronze dots. The flanks are bronze-brown with some small white stippling; the groin is green. The anterior surfaces of the thighs are dark brown with some cream stippling; the posterior surfaces of the thighs are dark brown with cream tubercles lateral and below the vent. The ventral surfaces are dark brown with uniformly distributed cream spots. The iris is yellow-cream with brown reticulations and a brown horizontal bar across the eye. Coloration in preservative. The dorsum is uniform bluish-gray. The pale labial, canthal, and dorsolateral stripes formed by a series of small bluish gray dots. The labial stripe continues to the insertion of the forelimb. The flanks, groin, and anterior surfaces of thighs are uniform dull gray; the posterior surfaces of thighs are dark gray with numerous white tubercles lateral to, and below, the cloacal opening. The ventral surfaces are uniform dark bluish-gray. Measurements (in mm). SVL: 54.0, TIBL: 20.5, FL: 24.7, HL: 17.8, HW: 20.5, IOD: 4.9, EW: 5.6 IND: 3.4, ED: 5.6, EN: 4.3, TD: 2.3, FFL: 9.9, TFL: 18.5, TFD: 3.4. Variation. Morphometric variation of three females and three males is summarized in Table 3. Females are larger than males (55.9�� 2.1 mm; 52.7.�� 2.5 mm). The skin on the dorsum varies from weakly to coarsely areolate. All adults have a supratympanic fold, which usually extends from the posterior edge of the tympanum to a point above the insertion of the forelimb. Color variation in preservative. All preserved specimens have a uniformly bluish gray dorsum; labial, canthal, and dorsolateral stripes are formed by a series of small brown dots. The dorsal surfaces of hands and feet also have brown dots. The flanks, groin, and anterior surfaces of the thighs are uniform dull gray, but in some specimens the flanks are covered by small dark gray dots. The posterior surfaces of the thighs are dark gray with numerous white tubercles lateral to, and below the cloacal opening; a short, fragmented supracloacal stripe is present in some individuals. The ventral surfaces are uniform dark bluish-gray. Color variation in life. (Figs. 10J, 18���19). The dorsum is uniform green (e.g., CJ 1386) or brown (e.g., CJ 7823). In CJ 1386, the dorsal surfaces of limbs are green, whereas the inner and outer surfaces of the forearms, thighs, and shanks are brown. Dorsal surfaces of hands and feet have a pattern of brown dots over a lighter background. Head markings are like those of the holotype; a dorsolateral stripe is present in all specimens. The flanks are bronze-brown with or without white stippling; the axillae and groin are green, with a bluish suffusion in some specimens. The anterior surfaces of thighs are dark brown with or without cream stippling and, and the posterior surfaces of thighs are dark brown with cream tubercles lateral to, and below, the vent. The ventral surfaces are dark brown with uniformly distributed cream dots, but some specimens have larger cream spots, thereby resulting in a paler venter. Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 39, from a series (CJ 1959) obtained from a pond at Laguna Negra de Jimbura, Parque Nacional Yacuri, 3406 m, Loja Province, Ecuador, by Elicio E. Tapia, Sof��a Carvajal-Endara, and Henry Grefa on 12 June 2011 (Fig. 5F). Total length 61.3, body length 23.5 (38% of total length). Body robust, ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile; body width at level of spiracle 14.7, and height at same position 11.9; head width at level of eyes 12.9. Lateral-line system not evident. Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout���nostril distance 3.7. Eyes positioned and directed dorsally, eye length 2.4, eye width 2.2. Spiracle sinistral, located at midbody level; spiracular opening oriented posteriorly; distance from tip of snout to spiracle opening 15.9; spiracle end rounded, not free, attached to body wall, inner wall of spiracle not evident; tube length 3.4, tube transverse width 1.4. Vent tube dextral, the opening oriented posteriorly, tube length 4.0, tube transverse width 2.6. Tail length 38.8, caudal musculature robust in the two-thirds proximal to body, narrowing gradually until tail terminus; tail muscle height 5.4, tail muscle width 4.8; caudal fins thin at proximal half of tail, getting higher on distal half and raising near tail���body junction, dorsal fin height 3.5, ventral fin height 3.7; maximum height of tail 12.7; tail tip rounded, tail musculature not reaching fin terminus. Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 5.6. It is surrounded by an uniserial row of marginal papillae, interrupted medially in upper lip; lower lip papillae alternating in and out, giving the appearance of two series; upper lip with 18 papillae on right side and 17 papillae on left side; lower lip with 42 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, height 0.4, transverse width 3.3 (59% of oral disc width); lower jaw sheath V- shaped, open and finely serrated, width 2.4, height 0.7. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.7, A2: 4.0, P1 right row 1.9, P1 left row 2.1, P1 gap 0.1, P2: 3.6, P3: 3.5. (Fig. 6E). Color in preservative. Dorsum dull gray, with darker areas on flanks, above the eyes and on the throat; body contour and snout translucent. Caudal musculature and fins with scattered, white spots, densely arranged near tailbody junction; fins otherwise translucent. Venter dark gray; eyes lavender gray, oral apparatus translucent. Color in life. (CJ 1957, Stage 36). Fig. 5F. In dorsal and lateral views, body tan, speckled with black; areas around the snout are lighter. Venter cream with black markings, semi-translucent; guts visible as a darker area; throat translucent with small cream flecks; gills evident as a red hue. Caudal musculature reddish-pink in proximal half, decreasing its intensity towards distal half; myomeres and nerves barely visible; caudal musculature and fins with cream marks, clustered in dorsal line of caudal musculature, otherwise caudal fins translucent. Legs cream. Oral apparatus light cream. Iris copper-yellow, with small black reticulations. Variation. Variation of 26 meristic characters of tadpoles in Stages 35���39 (CJ 1959) are shown in Table 7. Total length varies between 61.3 (Stage 39) and 75.0 (Stage 35); tail length proportion varied from 63.2 to 67.0 until Stage 39; labial tooth row formula was 2/3(1). Number of marginal papillae varied among specimens and Gosner stages, variation in lower lip papillae is high (42���57). ......continued on the next page We documented changes in coloration during ontogenetic development of CJ 1958���9 (Fig. 20). At Stage 36, the dorsum and flanks are pale brown with dark brown areas on posterior body and flanks. By Stage 42, dorsal surfaces of body and limbs are uniform green; fine, cream, dorsolateral stripes are present, extending to midbody. Cream stripes border the outer dorsal margins of the limbs. At Stage 46, dorsal surfaces of body and limbs have more well-defined cream stripes, and a cream labial stripe; the canthus rostralis is green, and remaining flanks are mostly dark brown with an upper diffuse black-stripe. Dorsal surfaces of fingers and toes are brown except on finger and toe discs, which are yellowish-cream. The iris is copper-yellow, lacking the brown horizontal band that is observed in adults. Comparisons. Tadpoles of Gastrotheca turnerorum may occur in sympatry with those of G. elicioi, G. lojana, G. psychrophila, and G. pseustes, in the Loja-Abra de Zamora region. Gastrotheca turnerorum differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body; from G. lojana by having a more rounded tail terminus and having bold cream marks in a dorsal line of caudal musculature, and from G. pseustes by having a more rounded tail terminus (compare in Fig. 5). For G. psychrophila, see remarks under G. elicioi tadpole account. Distribution and ecology. Gastrotheca turnerorum only is known from three localities in the Cordillera Oriental of the Andes in southern Ecuador. These localities are in Parque Nacional Yacuri, Parque Nacional Podocarpus, and Reserva Tapichalaca in Zamora Chinchipe and Loja provinces. Its elevational range is 3073���3406 m in an area of extent of occurrence of about 450 km 2. This mostly nocturnal, semiarboreal species inhabits mainly paramos and a few forests in the Evergreen Shrub and Herbazal of Paramo, and the Evergreen Montane Forest of Catamayo-Alamor (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 895���1278 mm and the average annual temperature is 10.4��� 13.5 ��C (Fick & Hijmans 2017). At the Lagunas del Compadre in the Parque Nacional Podocarpus, a gravid female was collected on 1 December 1994. It deposited tadpoles on 12 December in the laboratory, some of them were raised, and others were preserved. On 20 September 2009, another female with eggs in her pouch was found at the same locality. It was sitting on moss at approximately 2 m from the R��o Sabanilla, which is about 1.5 km from Lagunas del Compadre. The holotype, from Laguna Negra de Jimbura in the Parque Nacional Yacuri (Fig. 11C), was found at the edge of a pond about 3 x 2 m at the edge of a lagoon, at 14:00 h. The water temperature was 16.4�� C, and water pH was 5.3. Eighteen tadpoles were collected at this pond. Another group of tadpoles was found at 18:00 h; at the same locality in another lagoon, the water temperature was 7.4 �� C. Strong winds blow across the lagoon, and only a small patch of forest remained at one border (Elicio E. Tapia field notes, 12 June 2011). A brooding female is depicted in Figure 10J. Conservation status. We suggest that Gastrotheca turnerorum should be considered as an Endangered species according to criteria B2ab(i,ii,iii) of the IUCN Red List. Although this species occurs at the Yacuri and Podocarpus National Parks, we suggest this conservation status because of its small known area of occurrence (159 km 2) that is vulnerable to improperly regulated tourism activities (Aguirre 2001), introduced species such as trout, climate change, and pathogens. Laguna Negra de Jimbura has suffered the synergistic effects of agriculture, cattle and sheep raising, fires, introduced species, pesticide use, and unregulated tourism activitites. Etymology. The specific name turnerorum is a latinized word that honors the Turner family. As unique as this marsupial frog is to the amphibian world, the Turner family stands out in their exceptional, unwavering and ever- growing commitment to conservation of the world natural resources. From Ted Turner���s pioneering efforts to protect very large land areas and restore them to their original and natural states of clean water and abundant fauna and flora to granddaughter Elizabeth���s awareness-raising book ���Our friends the frogs,��� there are multiple generations of active conservationists with a true appreciation for the value of all of our natural resources. In 1991, Laura Turner Seydel (with father Ted) co-founded the Captain Planet Foundation and continues to work diligently to inspire the same appreciation for conservation and sustainable living in the next generation of young people who will surely be making big decisions that will affect the health of our planet for themselves and their own grandchildren. Through the camera���s eye, Rhett Turner has managed to put conservation issues such as the pollinator perils and amphibian crisis in front of millions of people. Through the Turner Endangered Species Fund, there is hope for dozens of threatened species of plants and animals in the US and abroad that was not there before. Together, the Turner family has put their hearts, souls, and personal resources into helping make th, Published as part of Carvajal-Endara, Sof��a, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Sz��kely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, pp. 1-102 in Zootaxa 4562 (1) on pages 48-56, DOI: 10.11646/zootaxa.4562.1.1, http://zenodo.org/record/2627982, {"references":["Orton, G. L. (1953) The systematics of vertebrate larvae. Systematic Zoology, 2 (2), 63 - 75. https: // doi. org / 10.2307 / 2411661","Fick, S. E. & Hijmans, R. J. (2017) WorldClim 2: new 1 - km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37 (12), 4302 - 4315. https: // doi. org / 10.1002 / joc. 5086","Aguirre, Z. (2001) La influencia de las actividades turisticas en los paramos del sur del Ecuador. In: Mena-Vasconez, P., Medina, G. & Hofstede, R. G. M. (Eds.), Los Paramos del Ecuador. Particularidades, Problemas y Perspectivas. Abya Yala, Quito, pp 217 - 218.","Duellman, W. E. (2015) Marsupial Frogs: Gastrotheca and Allied Genera. JHU Press, Baltimore, MD, 408 pp."]}

Gastrotheca pseustes Duellman & Hillis 1987 Gastrotheca pseustes was described from 7.1 km by road north of San Lucas, 2940 m (03° 41' S, 79° 15' W), Loja Province, Ecuador, in the southern Andean Cordillera. The type locality is located about midway between San Lucas and Saraguro. Current assignment of specimens to this taxon is constrained because: (1) there is a high phenotypic similarity between G. pseustes and G. lateonota from the Cordillera de Huancabamba in Peru (compare both species in Fig. 10 with Duellman 2015: Fig. 12.21); (2). There is no genetic information for topotypic G. lateonota, thereby precluding comparisons with specimens from southern Ecuador; (3) according to Duellman (2015), variation in morphometrics, structural characters, and coloration within G. pseustes presents a geographic mosaic, except for the southernmost population, Saraguro. The frogs from Saraguro are somewhat more distinctive than the others in being larger in size, and in having broader heads and some consistent characteristics of color pattern, including dark flanks with pale spots and uniformly dark posterior surfaces of the thighs; and (4) new molecular phylogeographic data (not shown) suggest that there may be at least two species in what is now recognized as Gastrotheca pseustes. While awaiting ongoing analyses of molecular data and a future report with a more detailed taxonomic and phylogeographic revision of Gastrotheca pseustes, we provide additional data of specimens (adults, tadpoles and metamorphs) from the type locality and surrounding areas in Loja and El Oro provinces. The recognition of Gastrotheca lateonota in southern Ecuador (from El Oro and Loja provinces) by Blackburn & Duellman (2013), Duellman et al. (2014), Yáñez-Muñoz et al. (2014), and Duellman (2015) was based on specimens identified as that species by Carvajal-Endara and Coloma. Subsequent examination of the specimens from Chilla, El Oro Province, reveals that they do not differ from the holotype (KU 203443, Fig. 10G) and additional specimens (Figs. 10 D–F) of G. pseustes, from 3.7 km S Saraguro, 2800 m. Thus, we do not recognize G. lateonota to occur in Ecuador, and treat the specimens from Chilla as part of a series of populations we consider G. pseustes complex (G. pseustes 1, G. pseustes 2, Fig. 33), occurring from the Equator south to high elevations of Podocarpus National Park in southern Ecuador (see also map in Duellman 2015: Figure 12.47). Conceivably, molecular data from topotypic G. lateonota from 31.5 km [by road] E Canchaque, 2770 m, Cordillera de Huancabamba, Región de Piura, Peru, (Fig. 33), and additional populations from northern Peru would resolve by (1) placement of non-type populations in either G. pseustes or G. lateonota, or (2) placement of G. lateonota as a junior synonym of G. pseustes. Castroviejo-Fisher et al. (2015) included in a phylogenetic analysis a specimen UINHM 94580 from San Rafael, Azuay, Ecuador under the name Gastrotheca riobambae, a species that does not occur in southern Ecuador. Their analysis places it in the G. pseustes complex. We also found a locality named San Rafael in Azuay Province, which is located between Cuenca and Cumbe, in Parroquia Tarqui at about 2700 m asl (not in the lowlands as was stated by Castroviejo-Fisher et al., 2015). This locality is well within the altitudinal and latitudinal range of G. pseustes 2. Herein, we provide meristic data from specimens of Gastrotheca pseustes 1 and 2 (Table 3), images depicting intrapopulation color pattern variation of metamorphs (Fig. 34), of adults of G. pseustes 1 from the type locality (Fig. 10G) and from Chillacocha (Figs. 10 D–F). Data for additional specimens of G. pseustes 1 are: QCAZ 45121, 45124 – 5, adult males, and QCAZ 45123 adult female from Chillacocha, ~ 8 km SW Chilla, El Oro Province (03° 30' 39.1" S, 79° 36' 52.92" W; 3163 m) collected on 18 August 2009. The frogs were on leaves of Gunnera sp. (Gunneraceae) near a small stream and adjoining marshland about 10 x 5 m. The water temperature was 9.6° C. Two individuals—CJ 201 adult female and KU 335386 adult male—are from Chillacocha, ~ 8 km SW Chilla, El Oro Province (03° 30' 15.37" S, 79° 37' 01.7" W; 3250 m) collected on 10 June 2011. They were in amplexus on a leaf of Gunnera sp. (Gunneraceae) approximately 90 cm above the ground and a small stream. Tadpoles were collected from a puddle next to the road in the vicinity of Chilla, El Oro Province (03° 27' 29.41" S, 79° 34' 52.07" W; 2452 m) on 11 June 2011; one of these was raised to be an adult male (CJ 399). Tadpoles were collected from a puddle next to the road at Manu, Loja Province (03° 33' 17.03" S, 79° 22' 6.02" W; 2876 m) on 11 June 2011; one of these was raised to be an adult female (CJ 400). Tadpoles were collected from the entrance of Parque Nacional Podocarpus, Loja Province (04° 05' 25.51" S, 79° 12' 09.97" W; 2456 m) on 14 June 2011; one of these was raised to be an adult female (KU 335387). Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36, from a series (CJ 1949) obtained from a pond at Chillacocha, 8 km SW Chilla, 3250 m, El Oro Province, Ecuador, by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa on 10 June 2011 (Fig. 5E). Total length 58.2; body length 22.7 (39% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat concave in lateral profile, sloping from anterior margin of snout to belly; body width at level of spiracle 15.3, and height at same position 12.9; head width at level of eyes 11.7. Lateral-line system present but barely visible, supraorbital and infraorbital lines both originating at tip of snout, running parallel to the eye and making contact immediately behind the eye; angular line descending vertically from just posterior of eye to throat; it dorsally contacts with post infraorbital line; post-supraocular present in form of a few stitches, anterior oral line descending vertically from oral disc level and behind nares level to throat, making a curve that parallels infraorbital line, forming a circuit continuous with angular and loreal lines; ventral line surrounds dorsally the spiracle; dorsal body and middle body lines not visible. Nostril medium sized (in proportion to body length), ovoid, protruding, with a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 3.9; internarial distance 3.6. Eye positioned and directed dorsolaterally, eye length 2.9, eye width 2.3; interorbital distance 7.1. Spiracle sinistral, located at midbody, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 15.6; end of spiracle rounded, attached to body wall, inner wall of spiracle not evident; tube length 2.7, tube transverse width 3.0. Vent tube dextral, opening directed posteriorly, tube length 2.9, tube transverse width 2.8. Tail length 35.0, caudal musculature robust, narrowing gradually until tail terminus; caudal muscle height 4.5, caudal muscle width 4.3; caudal fins well developed and proportional, dorsal fin originating near tail-body junction, forming low hump; dorsal fin height 4.2, ventral fin height 3.8; maximum height of tail 12.3; tail terminus rounded, caudal musculature not reaching fin terminus. Oral disc small, ventral, anteriorly reaching level of tip of snout, not protruding laterally beyond body, not visible dorsally; transverse width 5.9, surrounded by an uniserial row of small, marginal papillae, interrupted medially in upper lip; lower lip papillae alternating in orientation in and out, giving appearance of two rows; upper lip with 17 papillae on right side and 16 papillae on left side; lower lip with 52 marginal papillae; upper jaw sheath medium-sized, forming a finely serrated, smooth arch, with lateral processes, height 0.56, transverse width 3.7 (63% of oral disc width); lower jaw sheath V- shaped, finely serrated, width 2.9, height 0.68. Labial tooth row formula 2/3(1), tooth rows lengths: A1: 4.2, A2: 4.5, P1 right row 1.9, P1 left row 2.0, P1 gap 0.3, P2: 4.1, P3: 3.7. (Fig. 6F). Color in preservative. Dorsum gray with darker gray areas on flanks, above eye and on throat; margin of snout paler than adjacent areas. Caudal musculature and fins with a profusion of medium-sized dots that are most dense on upper part of the musculature; fins otherwise translucent. Venter cream, speckled with white, guts not exposed; eyes lavender gray with irregular white markings, oral apparatus translucent. Color in life. In dorsal view, body tan with black flecks. Flanks reticulated with black and tan; areas around the eyes and snout paler tan. Venter cream with black markings; throat translucent with small cream flecks; reddish gills evident. Caudal muscles reddish-pink, more evident on proximal half; myomeres barely visible; caudal muscles and fins having medium-sized cream dots, most dense on upper side of caudal muscles and near tail-body junction, forming a nearly conitnuous stripe along dorsolateral caudal musculature; otherwise caudal fins translucent. Legs cream with black markings on toes. Oral apparatus pale cream. Iris copper-yellow, with black reticulations. ......continued on the next page Variation. Variation of 28 meristic characters of tadpoles in Stages 36–42 (CJ 1949) are shown in Table 10. Total length varies between 55.0 (Stage 40) and 58.2 (Stage 36) and tail length proportion varies from 60% to 70% until Stage 42. Number of marginal papillae varies among specimens and Gosner stages; number of lower lip papillae is high (43–52). We documented changes in coloration during ontogenetic development of one, mostly pale brown individual (CJ 1953) (Figs. 5E, 35). At Stage 41, the dorsum and flanks are yellowish-brown with a poorly defined pattern of brown-gray, elongated paravertebral marks and few blotches on dorsum of hind limbs. A diffuse brown-gray stripe borders the canthal and dorsolateral body, and is bordered above by a creamy-brown stripe. The iris has a reddishgold suffusion. By Stage 45, markings on the dorsum and flanks are better defined with brown markings darkest peripherally; scattered green ill-defined flecks are present on dorsum of head, limbs, and canthus rostralis; tip of fingers are yellowish-cream. At Stage 46, paravertebral green-brown markings on dorsum of body and green flecks on limbs are contrasting to surrounding light brown areas; flanks have a dark brown stripe. Color variation in 6 metamorphs (CJ 1949, 1953–56, QCAZ 45126) in Stage 46 is depicted in Figure 34. They vary from dark brown to green with either well-defined paravertebral marks to a uniformly colored dorsum. Comparisons. Tadpoles of Gastrotheca pseustes sensu stricto (in Loja and El Oro provinces) occur in sympatry with those of G. elicioi in the Saraguro region and with G. elicioi, G. lojana, G. psychrophila, and G. turnerorum in the Loja-Abra de Zamora region. Gastrotheca pseustes differs from G. elicioi by lacking a dorsal gray-pigmented fin that abruptly arises from the body; from G. lojana by having a reticulated pattern on flanks, and from G. turnerorum by having a less rounded tail terminus (compare in Fig. 5). For G. psychrophila, see remarks under G. elicioi tadpole account. Vocalization. Four individuals of Gastrotheca pseustes were recorded from three locations in Loja Province (2 individuals from Bosque Washapamba, one from Vía Urdaneta-Tutupali, and 1 from Cerro de Arcos; see Appendix III). Because of the taxonomic problems related to the G. pseuste s complex we decided to use for the present analysis only the recordings obtained from populations situated nearby the type locality, south of the Jubones- Girón river valley in Loja Province (e.g. G. pseustes 1). Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call of G. pseustes is a complex call, composed of one to five long pulsed notes and followed (or not) by one to six short, single-pulsed notes (Fig. 23 H–N). The long note had a mean duration of 0.878 s (SD = 0.188) and consisted on average of 32.72 (SD = 6.867) distinct pulses, partly fused, without silent intervals (amplitude modulation close but less than 100%). The amplitude of the long note increases gradually towards the end of the note after which it decreases a little by the end. The short notes had a mean duration of 0.080 s (SD = 0.046) and the inter-note interval is on average of 0.795 s (SD = 0.502). The mean dominant frequency of the call is 1359.0 Hz (SD = 37.981), with a mean 90% bandwidth of 1043.2–1483.4 Hz. The fundamental frequency is usually recognizable; when visible, 6 to 7 harmonics are distinguishable. Comparisons: The advertisement call of Gastrotheca pseustes is most similar to that of G. yacuri, but G. pseustes has a much longer call duration, longer long notes duration, longer inter-note interval a lower dominant frequency and a lower 90% bandwidth frequency compared with the call of G. yacuri (Table 5). Also, G. pseustes usually emits 4–6 short notes compared with the only two short notes that G. yacuri emits. The call of G. pseustes can be easily distinguished from that of G. lojana and G. testudinea by the amplitude modulation of the longer note, much longer call duration, lower short note rate, longer long notes duration, longer inter-note interval, larger number of pulses, higher pulse rate, a higher dominant frequency, and higher 90% bandwidth frequency. Also, G. pseustes emits a larger number of long and short notes compared with G. lojana and G. testudinea (Table 5).

Gastrotheca Fitzinger, 1843 (Hemiphractidae) is composed of 70 species classified into four species groups, among which Gastrotheca fissipes (Boulenger, 1882) and G. microdiscus (Andersson, 1910) species groups are distributed in the Brazilian Atlantic Forest (Castroviejo-Fisher et al. 2015; Duellman & Venegas 2016). The Gastrotheca fissipes group comprises six species: G. fissipes, G. flamma Juncá & Nunes, 2008, G. megacephala Izecksohn, Carvalho-e-Silva & Peixoto, 2009, G. prasina Teixeira Jr. et al. 2012, G. pulchra Caramaschi & Rodrigues, 2007, and G. recava Teixeira Jr. et al. 2012, and most of these species are distributed in the northern Atlantic Forest, the most threatened region of the Atlantic Forest Domain (Ribeiro et al. 2009).

Fritziana fissilis .— Brazil: state of Rio de Janeiro: Teresópolis (Serra dos Órgãos): MZUSP 20611 –20612; MZUSP 20614; Nova Friburgo: MNRJ 56942; MZUSP 30, 55691 – 55692., Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044

Fritziana goeldii . — Brazil: state of Rio de Janeiro: Teresópolis (Serra dos Órgãos): MZUSP 103959 – 103965; Petrópolis: MNRJ 76201; Rio de Janeiro: CFBH 26981; USNM 318126; Rio de Janeiro (Tijuca): MZUSP 103426 - 103448; Guapimirim: MNRJ 24631; Cachoeira de Macacu: MNRJ 46537, 53758; Duque de Caxias: MNRJ 54715. State of São Paulo: Ubatuba: CFBH 9540, 10909 – 10910., Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044

Fritziana mitus sp. nov. (Figs. 1–3; Table 1) Flectonotus fissilis: Del Pino 1980:11, in part; Dixo and Verdade 2006:4; Condez et al. 2009:161. Fritziana fissilis: Duellman and Gray 1983:359, in part; Heyer et al. 1990:356; Duellman et al. 2011:31; Blackburn and Duellman 2013:712; Folly et al. 2014:399, in part. Fritziana aff. fissilis: Wachlevski et al. 2014:100; Franz and Mello 2015:4; Ceron et al. 2017:296; Comitti 2017:94. Fritziana ulei: Folly et al. 2014:399, in part; Menegucci et al. 2017:1. Holotype. — CFBH 39949, an adult female collected in an Atlantic Forest area in the Parque Estadual da Serra do Tabuleiro (27 º 44 ' 30 '' S, 48 º 48 ' 28 '' W; 420 m above sea level; datum = WGS84), municipality of Santo Amaro da Imperatriz, Santa Catarina State, Brazil, by Milena Wachlevski, on 14 April 2010 (Figs. 1 and 2). The holotype was collected by active search (at 2130 h) in a bromeliad (Vriesea sp.) at a height of 60 cm in relation to the soil. Paratopotypes. — Ten specimens consisting of six adult females without eggs on the dorsum (CFBH 39932 - 39935, CHUFSC 2625, and MNRJ 90352), two adult females incubating eggs on the dorsum (CFBH 39936 and MNRJ 90351), and two adult males (CFBH 39937 and MNRJ 90350). These specimens were collected in the municipality of Santo Amaro da Imperatriz, Santa Catarina state, Brazil, between July 2009 and January 2013. Diagnosis. —Based on 26 males and 24 females collected throughout the known distribution area of the new species (Appendix I), the new species is characterized by the following: (1) TD equal to 89% of the DTIV (females, 55– 115% [87 ± 15%, n = 24]; males, 59–135% [91 ± 18%, n = 26]); (2) HW equal to 93% of the HL (females, 84–109% [95 ± 6%, n = 24]; males, 78–102% [91 ± 6%, n = 26]); (3) a mucronate snout tip; (4) distal subarticular tubercles which are simple and round on the hands and feet; (5) the most common color pattern characterized by an interorbital triangle followed posteriorly by two parallel lines on the dorsum; (6) a black stripe on the wrist; (7) an egg-brooding pouch on the back of the females with the skin covering the eggs, and with a medial opening (the pouch in the reproductive period is evident even when empty); (8) tadpoles deposited in bromeliads; and (9) a change in the mitochondrial tRNA gene order (instead of having the order proline–phenylalanine–12 S, the new species has the order phenylalanine–proline–12 S). Comparisons with other species. —On average, Fritziana mitus sp. nov. is small when compared to other species within the genus. Therefore, large specimens of F. mitus sp. nov. might overlap in size with small specimens of F. ohausi, F. goeldii, F. tonimi, and F. izecksohni (SVL: F. mitus sp. nov.: males = 16.16–23.67 mm [19.97 ± 1.65 mm, n = 26], females = 17.43–28.35 mm [24.28 ± 2.47 mm, n = 24]; F. tonimi: males = 21.4–26.62 mm [23.67 ± 1.71 mm, n = 14], females = 27.04–31.20 mm [28.72 ± 1.90 mm, n = 4]; F. goeldii: males = 19.52–30.16 mm [27.12 ± 2.69 mm, n = 21], females = 30.41–37.03 mm [34.16 ± 3.47 mm, n = 7]; F. ohausi: males = 18.74–31.57 mm [26.80 ± 3.11 mm, n = 43], female = 27.6 mm [n = 1]; F. fissilis: males = 25.51– 29.97 mm [n = 2], females = 28.29–31.68 mm [30.13 ± 1.63 mm, n = 5]; F. ulei: males = 21.68–27.59 mm [23.24 ± 1.56 mm, n = 14], females = 23.07–25.07 mm [n = 2]; F. izecksohni: males = 21.5–29.4 mm [25.9 ± 2.8 mm, n = 5], female = 33.5 mm [n = 1]). The TD:DTIV ratio is larger in Fritziana mitus sp. nov. than in the other species except in F. goeldii (TD / DTIV = 0.89 in F. mitus sp. nov., females = 0.55–1.15 [0.87 ± 0.15, n = 24], males = 0.59–1.35 [0.91 ± 0.18, n = 26]; 0.79 in F. fissilis, females = 0.70–0.88 [0.77 ± 0.07, n = 5], males = 0.73–0.95 [n = 2]; 0.99 in F. goeldii, females = 0.92–1.31 [1.09 ± 0.12, n = 7], males = 0.61–1.38 [0.95 ± 0.16, n = 24]; 0.84 in F. tonimi, females = 0.72–1.18 [0.90 ± 0.22, n = 4], males = 0.64–0.98 [0.83 ± 0.11, n = 14]; 0.70 in F. ohausi, female = 0.78 [n = 1], males = 0.42–0.96 [0.70 ± 0.14, n = 43]; and 0.70 in F. ulei, females = 0.75–0.78 [n = 2], males = 0.42–0.96 [0.69 ± 0.13, n = 14]). Fritziana mitus sp. nov. has a proportionately wider head than F. ohausi and F. izecksohni, and a longer head than the other species (mean HW / HL for males = 0.93 in F. mitus sp. nov., females = 0.84–1.09 [0.95 ± 0.06, n = 24], males = 0.78–1.02 [0.91 ± 0.06, n = 26]; 0.83 in F. izecksohni; 0.86 in F. ohausi males = 0.79–0.96 [0.86 ± 0.04, n = 43]; 0.94 in F. goeldii, females = 0.90–1.01 [0.96 ± 0.04, n = 7], males = 0.89–1.01 [0.89 ± 0.03, n = 24]; 0.91 in F. tonimi, females = 0.89–0.94 [0.93 ± 0.02, n = 4], males = 0.84–0.98 [0.90 ± 0.04, n = 14]; 0.95 in F. ulei, females = 0.94–0.99 [n = 2], males = 0.91–1.99 [0.95 ± 0.03, n = 14]; and 0.93 in F. fissilis, females = 0.87– 1.00 [0.93 ± 0.05, n = 5], males = 0.88–0.97 [n = 2]). Fritziana mitus sp. nov. differs from F. fissilis and F. ohausi by having a mucronate snout tip (the tip of the snout is not mucronate in F. fissilis and in F. ohausi), and differs from F. fissilis and F. izecksohni by having round subarticular tubercles on all the fingers and toes (subarticular tubercles are bifid in F. fissilis and F. izecksohni). Although Fritziana mitus sp. nov. can exhibit a uniform color pattern (Fig. 3 F) or a bronze blotch covering the entire dorsal surface of the head (similar to F. ulei, Folly et al. 2014); the most common pattern (Fig. 3 A) is characterized by an interorbital triangle (also present in F. goeldii, F. ohausi, and F. tonimi) followed posteriorly by two parallel lines on the dorsum (a bilobated stain in F. goeldii and F. tonimi; an inverted V in F. ohausi). The type series of F. fissilis is discolored, but the dorsal pattern described in the original description seems to be similar to the pattern of the new species. Folly et al. (2014) noted the presence of an hourglass-shaped interorbital mark in other specimens identified by them as F. fissilis. Fritziana izecksohni exhibits a triangular mark covering the dorsum to the sacral region, but few specimens are known, and the occurrence of other patterns is possible. Fritziana tonimi can also have two long lines from the eye to the cloacal region forming a large V. The black stripe on the wrist is present in only the new species. Fritziana mitus sp. nov. differs from F. goeldii, F. ohausi, and F. tonimi, by having a closed dorsal pouch in females during egg incubation (as opposed to an opened pouch in F. goeldii, F. ohausi, and F. tonimi). Although the holotype of F. izecksohni shows dorsal folds on its back, indicating a closed dorsal pouch, no recorded observation of a female transporting eggs has occurred; therefore, its pouch morphology remains unknown. The new species further differs from F. ohausi on account of its reproductive behavior of releasing tadpoles into bromeliads (tadpoles are released into bamboo by F. ohausi). Although Folly et al. (2018) suggested that tadpoles of F. izecksohni develop in bromeliads, no clutch or tadpoles have been observed, so the microhabitat for this life-history stage remains unknown for F. izecksohni. The new species differs from Fritziana fissilis, F. goeldii, F. ohausi, F. tonimi, and F. ulei by possessing a unique order of mitochondrial genes. Instead of the most common neobatrachian gene order (proline–phenylalanine–12 S; Zhang et al. 2013), the new species has an order of phenylalanine–proline–12S (Walker et al. 2018: Fig. 4). The gene order in F. izecksohni remains unknown. Description of holotype. —Adult female without eggs on dorsum; head wider than long; HL 33% of SVL; snout rounded with tip slightly mucronate in dorsal view, protruding in lateral view. Nostril elliptical and protuberant, laterally oriented; IND 17% of the HW. Canthus rostralis distinct and straight; loreal region straight. Tympanum slightly evident; tympanic annulus hidden beneath skin; TD 35% of ED; supratympanic fold slightly developed. Postrictal tubercles absent. Eye oriented anterolaterally, its diameter 35% of HW; IOD 33% of HW; upper eyelid and interorbital region with small tubercles. Tongue ovoid, free on its posterior portion, and lacking a posterior notch; vomerine teeth in two oblique series that are next to each other; choanae large and rounded, well separated from each other. Dorsum smooth, belly areolate; ventral surfaces of thigh and pericloacal area granulated; gular region smooth. Flank homogeneous in color. Cloacal opening directed posteriorly at the upper level of thighs. Row of tubercles present on forearms; adhesive discs of fingers developed, expanded, and not emarginated; interdigital webbing absent on hand; relative length of fingers: III < II < V < IV; subarticular tubercles rounded; thenar tubercle developed and elliptic; two palmar tubercles developed, elliptical, and parallel to each other. FL 41% of SVL; THL 89% of the TBL; very small calcar present; tubercle on tibia absent; row of tubercles present on the tarsus; tibia slightly longer than thigh and together approximately the same length as SVL; adhesive discs of toes moderate in size, expanded, and not emarginated; interdigital webbing absent between Toes I and II and vestigial between others; relative length of toes: I < II < III < V < IV; subarticular tubercles rounded, few supernumerary tubercles; inner metatarsal tubercle elliptic and well developed; outer metatarsal tubercle slightly evident and rounded. Coloration of holotype. —In life, dorsal coloration of holotype brownish-yellow with a brownish interorbital fleck on the head. Dorsolateral stripe brownish with creamcolored outer edge extending from posterior margin of eye to midbody. Dorsal surface of discs on fingers dark. A dark brown band present on the wrist. In lateral view, maxilla with irregular dark spots. Venter with a uniform cream color. Iris pale bronze; a red horizontal stripe across eye (Fig. 1). In preservative, the specimen has the same general color patterns, but the colors are paler and the dark surface of the finger discs is not visible. Measurements of holotype (in mm). —SVL 24.7, HW 8.42, HL 8.13, AL 6.9, FAL 5.13, HAL 6.81, THL 11.62, TBL 13.01, TL 7.26, FL 10.1, IND 1.42, IOD 2.78, ED 2.96, END 2.47, TD 1.05, DFIV 1.09, and DTIV 1.04. Morphological variation (Table 1). —The snout is usually round in the lateral view (72.5% of the specimens), but sometimes truncate (20.7%) or protruding (6.8%). In the dorsal view, it is round (34.5%) or truncate (31%), but can also be pointed (17.25%) or subovoid (17.25%). The snout tip is slightly mucronate in the majority of specimens (86.25%), but not mucronate in others (13.75%). The dorsal skin is smooth with tubercles on the upper eyelid present in the minority of preserved specimens (17.3%), but they were apparent in live specimens even though they were not visible in the same specimens when preserved. Although rare, tubercles might also be present on the snout and between the eyes. The belly is grainy and the throat might be slightly grainy or smooth, with the rest of the ventral surface smooth. A dorsolateral stripe is usually present (65.6%) and when present it begins on the snout, passes over the eye, and extends posteriorly until the tympanum, the middle of the body, or the cloacal region. A characteristic pattern is formed by an interorbital bar and two stripes; the two stripes originate on the posterior corner of the eyes and converge in the middle of the head where they extend parallel to the middle of the body (Fig. 3 A). A small stripe might be present anteriorly and perpendicularly to the interorbital bar. Some specimens are uniformly beige with no distinct pattern (34.5%). In life, the dorsal coloration can be greenish, yellowish, bronze, or light brown. There might also be a bronze mark on the head, but this color is not visible in preserved specimens (Fig. 1 B). Specimens mostly have only one stripe on the wrist, like a bracelet, and other stripes or dots on the forelimb are rare (10.35%). The fingertips might be blackish, but this is normally only observed in live specimens. The hind limbs usually lack any type of pattern, but on rare occasions they might have dots or stripes (20.7%). When large dots are present on the legs they can be brown, bronze, or cream (Fig. 1 B). Also, a row of tubercles might be present on the forearms (41.4%) or more often, on the hind limbs (86.25%). Sexual dimorphism. —During the reproductive period, females exhibit a closed dorsal pouch when transporting eggs (Fig. 1 B). The eggs are normally distributed as a layer, without any regular pattern common to all females. Females without eggs might possess, on the dorsum, a skin crest that corresponds to the empty pouch (Fig. 3 A, E). Females with eggs removed from their dorsum after preservation possess individual skin chambers on the dorsum. Males possess nuptial pads on the base of Finger II, which can be pale beige or brown, and vocal slits on the floor of the mouth lateral to the tongue. The vocal sac is indistinct in preserved males. External morphology of tadpoles. —Even when belonging to the same egg clutch and being euthanized simultaneously, the tadpoles show different stages of development. In Stage 35, tadpoles have an elongated body, elliptical in both the dorsal and lateral views (Fig. 4). Their BL is 39.8% of the total length (Table 2). The body is wider than high; BH is 85.7% of its width. The BH is 93.7% of the TAH. The snout is rounded in dorsal view and truncate in lateral view. The eye is dorsally located, anterolaterally directed; with an ED equal to 16.9% of the BL; the nostril is closer to the tip of snout than to the eye; the distance between the eye and nostril is 63.6% of the distance between the eye and the tip of snout. The nostril is elliptical in the lateral view and the opening of the nostril is not visible dorsally. The nostril is anteriorly located and directed; the IND is 56.2% of the IOD; the distance between the nostril and the tip of snout is 8.4% of the BL. The spiracle is sinistral, ventral, short, and with the inner wall connected to the body. The SSD is 45.7% of the BL. The vent tube is as long as it is wide, dextral, with a wide opening, and attached to the ventral fin. The TAL is 69.6% of the total length and tail musculature does not reach the tail tip, which is rounded. In the lateral view, the dorsal fin originates on the tail musculature; the dorsal fin is ascendant in the first third of the tail, then becomes parallel to the tail musculature. The ventral fin is parallel to the tail musculature (Fig. 4). The oral disc is ventral and the ODW is 28.6% of the BW, bordered by a single row of papillae with a large dorsal gap (Fig. 5). Submarginal rows of papillae and labial teeth are absent. Jaw sheaths are developed and completely serrated; the upper is U-shaped and the lower is V-shaped (Fig. 5). Tadpole color. —Tadpoles preserved in 5% formalin are light brown with very small dark brown blotches; the flanks are lighter than the dorsum. Laterally, the tail is light cream, adorned with small dark brown blotches concentrated in the tail musculature and in the final third of the tail. Ventrally, the body is light brown with a translucent ventral region; the yolk is visible through the skin. The spiracle, the oral disc, and the cloacal tube are translucent beige. Molecular analyses. — Walker et al. (2018) reported the occurrence of a different gene order for mitochondrial tRNA in Fritziana mitus sp. nov. (named by them as F. CS 3) that is seemingly exclusive to this species until now. If the 12S gene is amplified using the primer MVZ59 (Graybeal 1997), the specimens show a fragment with about 200 more base pairs than in other species. We associate this with the proline tRNA gene, indicating a change in the gene order from the common neobatrachian order of proline–phenylalanine–12S, to an order of phenylalanine–proline–12S in the new species (Walker et al. 2018). All the other species of Fritziana possess the common neobatrachian size for this fragment of tRNA (Walker et al. 2018); however, this information is missing for F. izecksohni, whose gene order is unknown. Complete Bayesian analyses were made for Fritziana by Walker et al. (2018) and by our group (with only part of the 16 S fragment, including Fritziana izecksohni and outgroups; Fig. 6). Both analyses show that F. mitus clusters together, and is a sister group to only one sample from the state of Minas Gerais (F. CS 2 in Walker et al. 2018). Morphologically, this specimen is similar to F. mitus, but differs from the new species by the presence of an additional fragment in its tRNA (Walker et al. 2018: Fig. 4). The few specimens collected of F. CS2 preclude proper comparisons. Fritziana CS 2 (Walker et al. 2018) + F. mitus sp. nov. form the sister group of another new species, informally named by Walker et al. (2018) as F. CS1. The two samples of Fritziana izecksohni used by Folly et al. (2018) might have been contaminated because they cluster with Euparkerella and Cophomantini (Hylidae) in the present analyses (Fig. 6). This also explains the high inter- and intraspecific divergence and high length of branches found fo, Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044, {"references":["Del Pino, E. M. 1980. Morphology of the pouch and incubatory integument in marsupial frogs (Hylidae). Copeia 1980: 10 - 17.","Dixo, M., and V. Verdade. 2006. Herpetofauna de serrapilheira da Reserva Florestal do Morro Grande, Cotia (SP). Biota Neotropica 6: 1 - 20.","Condez, T. H., R. J. Sawaya, and M. Dixo. 2009. Herpetofauna dos remanescentes de Mata Atlantica da regiao de Tapirai e Piedade, SP, sudeste do Brasil. Biota Neotropica 9: 157 - 185.","Duellman, W. E., and P. Gray. 1983. Developmental biology and systematics of the egg-brooding hylid frogs, genera Flectonotus and Fritziana. Herpetologica 39: 333 - 359.","Heyer, W. R., A. S. Rand, C. A. Cruz, O. L. Peixoto, and C. E. Nelson. 1990. Frogs of Boraceia. Arquivos de Zoologia 31: 1 - 410.","Duellman, W. E., K. H. Jungfer, and D. C. Blackburn. 2011. The phylogenetic relationship of geographically separated '' Flectonotus '' (Anura: Hemiphractidae), as revealed by molecular, behavioral, and morphological data. Phyllomedusa 10: 29 - 43.","Blackburn, D. C., and W. E. Duellman. 2013. Brazilian marsupial frogs are diphyletic (Anura: Hemiphractidae: Gastrotheca). Molecular Phylogenetics and Evolution 68: 709 - 714.","Folly, M., F. Hepp, S. P. Carvalho-e-Silva, and W. E. Duellman. 2014. Taxonomic status and redescription of Flectonotus ulei (Anura: Hemiphractidae), with a key for the species of Fritziana. Zoologia 31: 393 - 399.","Wachlevski, M., L. K. Erdtmann, and P. C. A. Garcia. 2014. Anfibios anuros em uma area de Mata Atlantica da Serra do Tabuleiro, Santa Catarina. Biotemas 27: 97 - 107.","Franz, I., and M. H. Mello. 2015. Fritziana aff. fissilis (Miranda-Ribeiro, 1920) (Anura, Hemiphractidae): The first hemiphractid for the state of Rio Grande do Sul, southern Brazil. Check List 11: 1594.","Ceron, K., E. M. Lucas, and J. J. Zocche. 2017. Anurans of Parque Estadual da Serra Furada, Santa Catarina, Brazil. Herpetology Notes 10: 287 - 296.","Comitti, E. J. 2017. Herpetofauna da bacia do Rio Cachoeira, municipio de Joinville, Santa Catarina, Sul do Brasil. Acta Biologica Catarinense 4: 90 - 105.","Menegucci, R. C., R. Gaiga, and I. F. Machado. 2017. Fritziana ulei (Miranda-Ribeiro, 1926): Geographic extension, with comments on the natural history of this species. Check List 13: 2061.","Folly, M., F. Hepp, and S. P. Carvalho-e-Silva. 2018. A new bromeligenous species of Fritziana Mello-Leitao, 1937 (Amphibia: Anura: Hemiphractidae) from high elevations in the Serra dos Orgaos, Rio de Janeiro, Brazil. Herpetologica 74: 58 - 72.","Zhang, P., D. Liang, R. L. Mao, D. M. Hillis, D. B. Wake, and D. C. Cannatella. 2013. Efficient sequencing of anuran mtDNAs and a mitogenomic exploration of the phylogeny and evolution of frogs. Molecular Biology and Evolution 30: 1899 - 1915.","Walker, M., M. L. Lyra, and C. F. B. Haddad. 2018. Phylogenetic relationships and cryptic species diversity in the Brazilian egg-brooding tree frog, genus Fritziana Mello-Leitao 1937 (Anura: Hemiphractidae). Molecular Phylogenetics and Evolution 123: 59 - 72.","Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica 16: 183 - 190.","Graybeal, A. 1997. Phylogenetic relationships of bufonid frogs and tests of alternate macroevolutionary hypotheses characterizing their radiation. Zoological Journal of the Linnean Society 119: 297 - 338.","Carvalho-e-Silva, S. P., and A. M. Telles. 2010. Fritziana fissilis. The IUCN Red List of Threatened Species 2010. International Union for Conservation of Nature, Switzerland. Available at http: // dx. doi. org / 10. 2305 / IUCN. UK. 2010 - 2. RLTS. T 55315 A 11290233. en. Archived by Web- Cite at http: // www. webcitation. org / 6 uaUPaxx 3 on 29 October 2017."]}

Fritziana mitus .— Brazil: state of São Paulo: between Apia´&imath; e Iporanga (PETAR): CFBH 25632 – 25634; Biritiba Mirim: CFBH 12240; Caraguatatuba: CFBH 8273; Cunha: CFBH 12194; Iporanga: CFBH 6301; Itanhaém: CFBH 8333, 11177, 12205, 22113; Juquitiba: MZUSP 134762, 134764 – 134767; Piedade: CFBH 23316; Pilar do Sul: CFBH 8290, 8614, 10008; Salesópolis (Boracéia): MZUSP 137443; CFBH 1557, 1558; Santo André (Paranapiacaba): CFBH 28980; São Paulo: CFBH 13572; Ilhabela: CFBH 17496; Sete Barras: MZUSP 136570; Tapiraí: CFBH 10315; Ubatuba: CFBH 4542, 19955. State of Paraná: Antonina: CFBH 11106; Garuva: UFMG 18553; Guaraqueçaba: MZUSP 137966 – 137967; Guaratuba: CFBH 5482; Morretes: DZSJPR 5251; Tijucas do Sul: UFMG 18554. State of Santa Catarina: Brusque: MZUSP 9806 – 9809; Florianópolis: MCP-PUCRS 8284; Santo Amaro da Imperatriz: CFBH 39932 – 39937, 39949; CHUFSC 2625; MNRJ 90350 - 90352; São Francisco do Sul: UFMG 17384; Siderópolis: CFBH 24277, 25723. State of Rio Grande do Sul: Osório: UFMG 17436.

Fritziana ulei .— Brazil: state of Rio de Janeiro: Teresópolis (Serra dos Órgãos): MZUSP 103966 – 103973; 103976 –103978; 103980 – 103985; Cachoeira de Macacu: MNRJ 44622, 56922., Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044

Fritziana tonimi : Brazil: state of Espírito Santo: Santa Teresa: CFBH 24809; CFBH 30710 – 30712; CFBH 30920; MNRJ 28376, 38397 – 38398, 40700, 56060 – 56062; Domingos Martins: MNRJ 46719 – 46720., Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044

Fritziana ohausi. — Brazil: state of Rio de Janeiro: Teresópolis (Serra dos Órgãos): MZUSP 77702 – 77704; CFBH 32572 – 32573. State of São Paulo: São Luiz do Paraitinga (Parque Estadual da Serra do Mar–Núcleo Santa Virgínia): CFBH 7611, 14817 – 14830, 16276 – 16282; Pilar do Sul: CFBH 7552, 7620; Santos: CFBH 15950, 23920; Tapiraí: CFBH 16545; Apiaí and Iporanga: CFBH 25601; São José do Barreiro (Serra da Bocaina): MZUSP 31241, 34607 – 34611, 57731 – 57732; Santo André (Parque Estadual Municipal Nascentes de Paranapiacaba): CFBH 28974, 28982, MZUSP 77000 – 77001; Ribeirão Grande (Estação Ecológica de Xitué): MZUSP 139225; Salesópolis (Boracéia): MZUSP 137418 – 137419., Published as part of Walker, Marina, Wachlevski, Milena, Nogueira-Costa, Paulo, Garcia, Paulo C. A. & Haddad, Célio F. B., 2018, A New Species of Fritziana Mello-Leitão 1937 (Amphibia: Anura: Hemiphractidae) from the Atlantic Forest, Brazil, pp. 329-341 in Herpetologica 74 (4) on pages 329-341, DOI: 10.1655/0018-0831.329, http://zenodo.org/record/7803044

Hemiphractus elioti new species (FIgS. 2���4, 6) Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from western Panama. Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from western Panama]; Duellman 2001 [in part; for reference to specimens from western Panama]; Crawford et al. 2012 [in part, for samples from R��o Blanco and El Cop��, Prov. Cocl��, and Altos de Maria, Prov. Panam��, Panam��]; Castroviejo-Fisher et al. 2015 [in part; for sample from El Cop�� Prov. Cocl��, Panam��]; K��hler 2011 [fig. 467; in part, for map showing isolated populations in western Panama]. Holotype. UTACV A-64827, A SMALL, ADULT MALE, OBTAINED AT PANAM��: PROV. PANAM��: R��O MARIA, ALTOS DE MARIA, N 0 8 o 38.585���, W 0 80o 04.357���, 875 M, ON 20 MAY 2005, BY EDgARDO GRIFFITh AND HEIDI ROSS. Paratypes. FEMALES: UMMZ 152622, 152623, PANAMA: COCL��: 6 MI [9.7 kM] NNW EL COP��, 600���800 M; AMNH A124114, PANAMA: ChIRIqUI: CONTINENTAL DIVIDE ABOVE QUEBRADA DE ARENA, 1200���1300 M. MALES: AMNH A124113, A124115, PANAMA: ChIRIqUI: CONTINENTAL DIVIDE ABOVE QUEBRADA DE ARENA, 1200���1300 M; UMMZ 153118, PANAMA COCL��: 6 MI [9.7 kM] NNW EL COP��, ON CONTINENTAL DIVIDE AT SAwMILL, 600���800 M, USNM 572599, PANAMA: COCL��: N OF EL COP��. Diagnosis. Hemiphractus elioti MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. fasciatus, H. kaylockae, H. panamensis, AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION. Hemiphractus elioti MAY BE DISTINgUIShED FROM H. scutatus, AN AMAzONIAN SPECIES NOT OCCURRINg IN PANAMA, BY ITS SMALLER SIzE (MAXIMUM SVL IN MALES 52.5 MM VS. 57.4 MM IN H. scutatus; FEMALES 64.7 MM VS. 80.5 MM IN H. scutatus; DATA FOR H. scutatus FROM TRUEB 1974), AND BY hAVINg SMALL EXPANDED PADS ON ThE FINgERS AND TOES (ABSENT IN H. scutatus). Hemiphractus elioti DIFFERS FROM H. fasciatus, H. kaylockae, AND H. panamensis BY hAVINg NEOPALATINE ODONTOIDS IN CONTACT wITh VOMERINE TEETh AT ThE CENTER OF ThE PALATAL REgION (NARROwLY SEPARATED IN H. kaylockae VS. wIDELY SEPARATED IN BOTh H. panamensis AND H. fasciatus; CONDITION UNkNOwN IN H. scutatus; FIg. 6). Hemiphractus elioti FURThER DIFFERS FROM H. panamensis AND H. fasciatus BY hAVINg APPROXIMATELY SEVEN VOMERINE ODONTOIDS (VS. APPROXIMATELY TwO; CONDITION UNkNOwN IN H. scutatus; FIg. 6). Hemiphractus elioti MAY BE DISTINgUIShED FROM H. kaylockae BY hAVINg STRAIghT MARgIN OF ThE LATERAL MARgINS OF ThE PARAOCCIPITAL hORNS (wITh DISTINCT INDENTATION, FORMINg AN ANgULAR MARgIN IN H. kaylockae; FIg. 2); SUPRAORBITAL RIDgES ABSENT (EVIDENT, DISTINCT IN H. fasciatus, wEAkLY DEVELOPED IN H. panamensis; FIgS. 2, 9); CANThAL RIDgES USUALLY ABSENT OR INDISTINCT (EVIDENT, DISTINCT IN H. fasciatus, H. kaylockae, AND SLIghTLY DEVELOPED IN H. panamensis; FIgS. 2, 9). Hemiphractus elioti hAS A SUBTEMPORAL FENESTRA VERY SMALL IN MALES, APPROXIMATELY ONE-hALF OF ORBIT IN FEMALES, whEREAS ThE FENESTRAE ARE APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES OF H. kaylockae (FIg. 2), APPROXIMATELY �� DIAMETER OF ThE ORBIT OF FEMALES IN H. fasciatus (FIg. 9; MALES UNkNOwN), VERY LARgE, APPROXIMATELY 1.5 TIMES SIzE OF ORBIT, IN FEMALES OF H. panamensis (FIg. 9; MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgALS EXTEND TO LEVEL OF ThE LATERAL TIPS OF, OR VERY SLIghTLY BEYOND, ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (EXTENDINg FAR BEYOND ThE LATERAL TIPS OF ThE PARAOCCIPITAL hORNS IN H. panamensis; FIgS. 2, 9). Description of the holotype. SMALL, ADULT MALE, SNOUT ��� VENT LENgTh (SVL) 40.5 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 22.3 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 19.6 MM; hEAD LENgTh 20.3 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 20.2 MM; SNOUT ��� PARAOCCIPITAL PROCESS LENgTh 20.2 MM; TYMPANUM OVOID, 5.4 MM hIgh X 3.8 MM wIDE; EYE DIAMETER 5.5 MM; INTERORBITAL DISTANCE 6.9 MM; EYE ��� TYMPANUM DISTANCE 3.1 MM; BODY MODERATE; hEAD LARgE, wIDER ThAN BODY, gREATEST wIDTh AT LEVEL OF PARS ARTICULARIS, BEARINg PROMINENT PARAOCCIPITAL PROCESSES; PARAOCCIPITAL PROCESSES PROjECTINg POSTEROLATERALLY IN DORSAL VIEw, EXTENDINg POSTERIOR TO LEVEL OF PARS ARTICULARIS; SkIN CO-OSSIFIED OVER DORSAL SURFACE OF SkULL; OCCIPITAL AND PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL, zYgOMATIC RIDgES ABSENT; PREORBITAL, CANThAL RIDgES BARELY EVIDENT, LESS PROMINENT ThAN OCCIPITAL AND PARAOCCIPITAL RIDgES; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, TRIANgULAR IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT LESS ThAN 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY, SLIghTLY LATERALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 34 % hEAD wIDTh; EYE DIAMETER 27 % hEAD wIDTh; TYMPANUM VERTICALLY OVOID, ORIENTED POSTERIORLY AND SLIghTLY LATERALLY, VISIBLE BELOw PARAOCCIPITAL PROCESSES IN DORSAL VIEw, gREATEST DIAMETER 1.5 TIMES EYE DIAMETER; wIDTh OF TYMPANUM 1.2 TIMES EYE ��� TYMPANUM DISTANCE; UPPER EYELID BEARINg SINgLE ENLARgED POINTED PROjECTION AND SCATTERED SMALLER TUBERCLES; INTERORBITAL, POSTORBITAL TUBERCLES ABSENT; SkIN OVER POSTERIOR SURFACE OF PARS ARTICULARIS BEARINg ONE SMALL, FLEShY TUBERCLE. TONgUE LARgE, OVOID; VOCAL SLITS ABSENT; MANDIBLE BEARINg SERRATIONS ALONg ENTIRE LENgTh OF LOwER jAw, wITh TwO PROMINENT, TUSk- LIkE ODONTOIDS PROjECTINg FROM MENTOMECkELIAN BONES. AXILLARY MEMBRANE ABSENT; ThORACIC FOLD ABSENT; DERMAL FOLDS ON wRISTS ABSENT; FINgERS LONg, SLENDER, BEARINg ENLARgED DISTAL PADS; FINgER PADS 2.6 TIMES DIAMETER OUTER METACARPAL TUBERCLE LOw, TRIPARTITE, DIFFUSE; INNER METACARPAL TUBERCLE OVOID, RAISED; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FLANgES ABSENT ON FINgERS; FOREARM wITh SCATTERED TUBERCLES, LOw, ROUND, BEARINg wEAkLY DEVELOPED LATERAL FRINgE; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION EXTENDINg TO SNOUT whEN hINDLIMB EXTENDED; TARSAL FOLD PRESENT, wEAkLY DEVELOPED; TIBIA LENgTh 55 % SVL; FOOT LENgTh 48 % SVL; OUTER METATARSAL TUBERCLE wEAkLY DEVELOPED, DIFFUSE; INNER METATARSAL TUBERCLE PRESENT, RAISED, OVOID; TOES II���IV wITh PROMINENT, LATERAL FRINgE; FRINgE OF TOE V EXTENDINg ONTO OUTER SURFACE OF BASE OF TARSUS; DORSAL SURFACE OF hIND LIMB RELATIVELY SMOOTh, BEARINg IRREgULAR SETS OF TUBERCLES CONCENTRATED ON ThREE TRANSVERSE BANDS OF DARk-BROwN COLORATION ON ShANk; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; TOE PADS 2 X wIDTh OF PENULTIMATE PhALANgES OF DIgITS; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh. DORSUM AND FLANkS SMOOTh; VENTRAL SURFACES AND ThROAT AND ChIN IRREgULARLY gRANULAR; SACRAL DIAPOPhYSES PROjECTINg LATERALLY BEYOND TRANSVERSE PROCESSES OF ALL VERTEBRAE; LATERAL TERMINI OF TRANSVERSE PROCESSES OF PRESACRAL VERTEBRAE VI���VIII NOT CO-OSSIFIED wITh SkIN (NEARLY TO LEVEL OF SACRAL DIAPOPhYSES) AND VISIBLE EXTERNALLY; SIX NEURAL SPINES PROjECTINg DORSALLY, CO-OSSIFIED wITh SkIN, VISIBLE EXTERNALLY. PUPIL hORIzONTAL. IN ALCOhOL, DORSUM BEIgE wITh MINUTE BLACk PUNCTATIONS (FIg. 4). ThIgh AND ShANk BEARINg ThREE INDISTINCT TRANSVERSE BANDS OF IRREgULAR, BROwN MOTTLINg ON DORSAL SURFACES, CORRESPONDINg wITh ThREE BANDS OF TUBERCLES ON ShANk ONLY; ULTIMATE AND PENULTIMATE PhALANgES OF TOES IV���V DARk BROwN; FOREARM BEARINg BROAD, BROwN, TRANSVERSE BAND ON DORSAL SURFACE; DORSAL SURFACE OF MANUS MORE PALE ThAN FOREARM, ULTIMATE AND PENULTIMATE PhALANgES OF FINgERS V���IV DARk BROwN. UPPER AND LOwER LIPS wITh NARROw, TRANSVERSE, DARk BROwN BANDS, NOT EXTENDINg ONTO PROXIMAL SURFACES. FLANkS BEIgE wITh DIFFUSE BROwN PUNCTATIONS; VENTER BEIgE wITh SCATTERED DIFFUSE, FINE BROwN MARkINgS; ThROAT BEIgE wITh FINE BROwN MOTTLINg. VENTRAL SURFACE OF FOREARM BEIgE wITh FINE, BROwN MOTTLINg; VENTRAL SURFACE OF hAND BEIgE. VENTRAL SURFACES OF hIND LIMB AND FOOT BEIgE wITh FINE, PALE BROwN MOTTLINg; TOES I AND II ARE UNIFORMLY BEIgE, TOES III���V UNIFORMLY BROwN. Variation. CONSISTENT SEXUAL DIMORPhISM IN COLORATION OR SkIN TEXTURE wAS NOT EVIDENT. GRANULATION OF ThE ThROAT IN Hemiphractus elioti IS VARIABLE AMONg SPECIMENS EXAMINED, VARYINg FROM SMOOThLY gRANULAR wITh NO TUBERCULATION (E.g., USNM 153118, 297543, 572600, 572601) TO SMOOThLY gRANULAR wITh DISCRETE TUBERCLES PRESENT (E.g., AMNH 107955, 124113, 124115, UMMZ 167358, 297542, 572599). TEXTURE OF ThE VENTER RANgES FROM SMOOTh (E.g., UMMZ 152622,152623) TO UNIFORMLY gRANULAR (E.g., UMMZ 297543); SOME SPECIMENS (E.g., AMNH 107955, 124113, 124114, UMMZ 153118, USNM 297541, 297542, 572599) wITh gRANULAR VENTERS ALSO hAD VARIABLE NUMBERS OF DISCRETE TUBERCLES PRESENT. ThROAT COLORATION IS hIghLY VARIABLE, INCLUDINg SPECIMENS wITh UNIFORM TAN (E.g., AMNH 107955, USNM 297542, 297543) TO UNIFORM DARk BROwN (E.g., AMNH 124113���15, UMMZ 152622, 152623, USNM 572599); EIThER PATTERN MAY INCLUDE A FAINT TO PROMINENT PALE-COLORED STRIPE EXTENDINg FROM ThE SYMPhYSIS OF ThE LOwER jAw ONTO ThE PECTORAL REgION, A SPECkLED PATTERN OF whITE AND BROwN (E.g., USNM 572600), OR A gENERALLY whITE gROUND COLORATION, wITh DISTINCT MARgINAL gRAY MARBLINg (E.g., USNM 572601). VENTRAL COLORATION IN MOST SPECIMENS TYPICALLY IS A CONTINUATION OF ThE gULAR COLORATION OF DARk BROwN OR TAN EXTENDINg TO APPROXIMATELY MID-ABDOMEN whERE IT ABRUPTLY ChANgES TO A TAN COLORATION (E.g., AMNH 107955, 124113���15, UMMZ 153118, 152622, 152623, USNM 572599). VENTRAL COLORATION OF SOME SPECIMENS (E.g., USNM 297541���43) IS UNIFORMLY PALE TAN, OR whITE (E.g., UMMZ 167358, 572600, USNM 572601); ONE SPECIMEN (USNM 572600) IS UNIqUE IN hAVINg A SPECkLED VENTRAL PATTERN OF BROwN AND whITE. ThE DORSUM IS UNIFORMLY SMOOThLY gRANULAR IN ALL SPECIMENS AND OVERALL A UNIFORM ShADE OF BROwN. MOST SPECIMENS ShOw A SINgLE DARk BAND ON ThE FOREARM (ABSENT OR BARELY DISCERNIBLE IN AMNH 107955, UMMZ 152623, USNM 297542, 297543, 572599, 572600); TwO OR ThREE DARk TRANSVERSE BANDS ON ThE FEMORAL AND TIBIAL REgIONS (ABSENT OR BARELY DISCERNIBLE ON AMNH 10795, UMMZ 152623, 297541���43, 572599���601), AND A DARk BAND OR ChEVRON PRESENT ON ThE SACRAL REgION (E.g., AMNH 124113���15, UMMZ 152623); IN ADDITION, SPECIMEN AMNH 124113 hAS A DARk IRREgULAR VERTEBRAL STRIPE EXTENDINg FROM ThE SACRAL BAND TO ThE OCCIPUT. ThE POSTERIOR SURFACE OF ThE ThIgh OF USNM 572599 BEARS A PATTERN OF TAN SPOTS OVER A BROwN BACkgROUND COLORATION, AND UMMZ 152623 hAS A DISTINCT whITE STRIPE ALONg ThIS SURFACE. Etymology. EDgARDO GRIFFITh AND HEIDI ROSS hAVE gREATLY CONTRIBUTED TO ThE kNOwLEDgE BASE OF ThE NATURAL hISTORY, REPRODUCTIVE BIOLOgY, AND CONSERVATION OF PANAMANIAN AMPhIBIANS, INCLUDINg Hemiphractus. WE ARE PROUD TO ACkNOwLEDgE ThE EFFORTS OF OUR FRIENDS AND COLLEAgUES BY NAMINg ThIS NEw SPECIES OF Hemiphractus IN hONOR OF ThEIR SON, ELIOT. ThE SPECIFIC EPIThET IS ThE SINgULAR gENITIVE CASE OF ThE NAME ELIOT. Distribution and ecology. SPECIMENS OF Hemiphractus elioti hAVE BEEN COLLECTED FROM VARIOUS LOCALITIES IN ThE CORDILLERA DE TALAMANCA, INCLUDINg SITES IN ThE PROVINCES OF BOCAS DEL TORO, ChIRIqUI, COCL��, AND PANAM�� IN wESTERN PANAMA (FIg. 5). HABITAT IS PRE-MONTANE MOIST PRIMARY FOREST RANgINg IN ELEVATION FROM 600���1600 M. LIVINg SPECIMENS hAVE BEEN OBSERVED FROM gROUND LEVEL TO Hemidactylus SPP.), NEwBORN MICE (Mus musculus), AND EARThwORMS (H. ROSS AND E. GRIFFITh PERS. COMM., RLH PERS. OBS.). WhILE DISSECTINg SPECIMENS TO DETERMINE SEX, UNIDENTIFIED ANURANS wERE EVIDENT IN ThE STOMAChS OF SPECIMENS. NUMEROUS UNIDENTIFIED NEMATODE wORMS wERE ALSO OBSERVED IN ThE COELOM AND IMBEDDED IN ThE PERITONEUM AND TISSUES OF OThER ABDOMINAL ORgANS. MYERS (1966) DESCRIBED A DISTINCTIVE MOUTh gAPINg BEhAVIOR, YELLOw ORAL COLORATION, AND DEFENSIVE BITINg IN SPECIMENS REFERABLE TO H. panamensis (FIg. 3B) AND H. kaylockae (FIg. 8C). SIMILAR BEhAVIORS hAVE BEEN OBSERVED IN H. elioti. REPEATED OBSERVATIONS OF CAPTIVE REPRODUCTION OF ThIS SPECIES AT ThE EL VALLE AMPhIBIAN CONSERVATION CENTER hAVE ShOwN ThAT FEMALES CARRY EggS AND YOUNg ON ThEIR BACkS, AS IS kNOwN TO OCCUR IN ALL DESCRIBED SPECIES OF Hemiphractus., Published as part of Hill, Robert L., Martin, Kathryn G., Stanley, Edward & Mendelson Iii, Joseph R., 2018, A taxonomic review of the genus Hemiphractus (Anura: Hemiphractidae) in Panama: Description of Two New Species, Resurrection of Hemiphractus panamensis (Stejneger, 1917), and Discussion of Hemiphractus fasciatus Peters, 1862, pp. 495-512 in Zootaxa 4429 (3) on pages 499-503, DOI: 10.11646/zootaxa.4429.3.3, http://zenodo.org/record/1285044, {"references":["Duellman, W. (1970) The hylid frogs of Middle America. Vol. 1. Monograph of the Museum of Natural History, University of Kansas, 1, 1 - 427.","Trueb, L. (1974) Systematic relationships of Neotropical horned frogs, genus Hemiphractus (Anura: Hylidae). Occasional Papers of the Museum of Natural History, University of Kansas, 29, 1 - 60. https: // doi. org / 10.5962 / bhl. part. 11750","Duellman, W. (2001) The Hylid Frogs of Middle America. Vol. 2. Society for the Study of Amphibians and Reptiles Contributions in Herpetology, Ithaca, New York, 1170 pp.","Crawford, A., Cruz, C., Griffith, E., Ross, H., Ibanez, R., Lips, K. & Crump, P. (2012) DNA barcoding applied to ex situ tropical amphibian conservation programme reveals cryptic diversity in captive populations. Molecular Ecology Resources, 13, 1005 - 1018.","Castroviejo-Fisher, S., Padial, J., de La Riva, I., Pombal Junior, J., da Silva, H., Rojas-Runjaic, F. J. M., Medina-Mendez, E. & Frost, D. (2015) Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development. Zootaxa, 4004 (1), 1 - 75. https: // doi. org / 10.11646 / zootaxa. 4004.1.1","Myers, C. (1966) The distribution and behavior of a tropical horned frog Cerathyla panamensis Stejneger. Herpetologica, 22, 68 - 71."]}

Hemiphractus kaylockae new species (FIgS. 2, 4, 6, 8) Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from eastern Prov. Darién, Panamá. Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Duellman 2001 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Crawford et al. 2012 and Castroviejo-Fisher et al. 2015 [in part, for samples from Cana, Prov. Darién, Panamá]; Köhler 2011 [in part; for map showing isolated populations in extreme eastern Panama]. Holotype. KU 107409, ADULT FEMALE, OBTAINED AT PANAMá: PROV. DARIéN: N RIDgE OF CERRO CITURO, SIERRA DE PIRRE, N 0 7 o 56’, W 77o 42’ (APPROX.), 900–1000 M ON 17 JANUARY 1966, BY ChARLES W. MYERS. Paratypes, MALES: KU 107411, 107414 PANAMA: DARIEN: N RIDE CERRO CITURO, SERRANIA DE PIRRE, 1000 M; KU 107419 PANAMA: DARIEN: N RIDE CERRO CITURO, SERRANIA DE PIRRE, 900 M; KU 154992, PANAMA: DARIEN: S SLOPE CERRO CITURO, SERRANIA DE PIRRE, 1100 M. FEMALES: AMNH A92668, PANAMA: DARIEN: N RIDgE CERRO CITURO, SERRANIA DE PIRRE, 1100 M; KU 93506, PANAMA: DARIEN: SE SLOPE CERRO PIRRE, 1440 M; KU 107413, PANAMA: DARIEN: N RIDgE CERRO CITURO, SERRANIA DE PIRRE, 1000 M; KU 93507, PANAMA: DARIEN, SE SLOPE CERRO PIRRE, 1440 M. Diagnosis. Hemiphractus kaylockae MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. elioti, H. fasciatus, H. panamensis AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION (FIg. 2). Hemiphractus kaylockae MAY BE DISTINgUIShED FROM H. scutatus, AN AMAzONIAN SPECIES NOT OCCURRINg IN PANAMA, BY ITS SMALLER SIzE (MAXIMUM SVL IN MALES 52.5 MM VS. 57.4 MM IN H. scutatus; FEMALES 64.7 MM VS. 80.5 MM IN H. scutatus; DATA FOR H. scutatus FROM TRUEB, 1974), AND BY hAVINg SMALL EXPANDED PADS ON ThE FINgERS AND TOES (ABSENT IN H. scutatus). Hemiphractus kaylockae DIFFERS FROM H. elioti BY hAVINg NEOPALATINE AND VOMERINE ODONTOIDS NARROwLY SEPARATED (IN CONTACT IN H. elioti; CONDITION UNkNOwN IN H. scutatus), whEREAS ThESE ROwS OF ODONTOIDS ARE wIDELY SEPARATED IN H. panamensis AND H. fasciatus (FIg. 6). Hemiphractus kaylockae FURThER DIFFERS FROM H. panamensis AND H. fasciatus BY hAVINg APPROXIMATELY SEVEN VOMERINE ODONTOIDS (VS. APPROXIMATELY TwO IN BOTh H. panamensis AND H. fasciatus). Hemiphractus kaylockae MAY BE DISTINgUIShED FROM H. elioti, H. fasciatus, AND H. panamensis BY hAVINg A DISTINCT INDENTATION ON ThE paraoccipital hORMS, FORMINg AN ANgULAR MARgIN OF ThE PARAOCCIPITAL hORNS (FIgS. 2, 9); SUPRAORBITAL RIDgES ABSENT (EVIDENT, DISTINCT IN H. fasciatus AND wEAkLY DEVELOPED IN H. panamensis; FIgS. 2, 9); CANThAL RIDgES EVIDENT, DISTINCT (ABSENT OR wEAkLY DEVELOPED IN H. elioti; FIg. 2). Hemiphractus kaylockae hAS A SUBTEMPORAL FENESTRA APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES, APPROXIMATELY ONE-hALF DIAMETER OF ORBIT IN FEMALES IN H. elioti (VERY SMALL IN MALES; FIg. 2), APPROXIMATELY ONEhALF DIAMETER OF ThE ORBIT IN FEMALES OF H. fasciatus (FIg. 9; MALES UNkNOwN), VERY LARgE, APPROXIMATELY 1.5 X SIzE OF ORBIT, IN FEMALES OF H. panamensis (FIg. 9; MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgALS EXTEND TO LEVEL OF LATERAL TIPS OF, OR SLIghTLY LESS ThAN, BEYOND ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (EXTENDINg FAR BEYOND ThE PROFILE OF ThE PARAOCCIPITAL hORNS IN H. panamensis; FIgS. 2, 9). Description of the holotype. AN ADULT FEMALE: SVL 54.2 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 28.4 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 25.3 MM; hEAD LENgTh 24.3 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 26.5 MM; SNOUT-PARAOCCIPITAL PROCESS LENgTh 25.1 MM; TYMPANUM OVOID, 6.6 MM hIgh X 3.6 MM wIDE; EYE DIAMETER 6.5 MM; INTERORBITAL DISTANCE 10.2 MM; EYE – TYMPANUM DISTANCE 5.4 MM; BODY MODERATE; hEAD LARgE, wIDER ThAN BODY, gREATEST wIDTh AT LEVEL OF PARS ARTICULARIS, BEARINg PROMINENT PARAOCCIPITAL PROCESSES; PARAOCCIPITAL PROCESSES PROjECTINg POSTEROLATERALLY IN DORSAL VIEw, EXTENDINg POSTERIOR TO LEVEL OF PARS ARTICULARIS; SkIN CO-OSSIFIED OVER DORSAL SURFACE OF SkULL; OCCIPITAL RIDgE PRESENT, wELL-DEVELOPED; PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL, zYgOMATIC, PREORBITAL, AND CANThAL RIDgES ABSENT; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, LATERALLY FLATTENED IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT LESS ThAN 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 38 % OF hEAD wIDTh; EYE DIAMETER 25 % hEAD wIDTh; TYMPANUM VERTICALLY OVOID, ORIENTED POSTERIORLY AND SLIghTLY LATERALLY, VISIBLE BELOw PARAOCCIPITAL PROCESSES IN DORSAL VIEw, gREATEST DIAMETER EqUIVALENT TO EYE DIAMETER; wIDTh OF TYMPANUM 1.6 TIMES EYE – TYMPANUM DISTANCE; UPPER EYELID wITh ONE ENLARgED POINTED PROjECTION AND SCATTERED SMALLER TUBERCLES; INTERORBITAL TUBERCLE ABSENT; POSTORBITAL TUBERCLE ABSENT; SkIN OVER POSTERIOR SURFACE OF PARS ARTICULARIS BEARINg ONE SMALL, FLEShY TUBERCLE. TONgUE LARgE, OVOID; VOCAL SLITS ABSENT; MANDIBLE BEARINg SERRATIONS ALONg ENTIRE LENgTh OF LOwER jAw, wITh TwO PROMINENT, TUSk-LIkE ODONTOIDS PROjECTINg FROM MENTOMECkELIAN BONES. AXILLARY MEMBRANE ABSENT; ThORACIC FOLD ABSENT; DERMAL FOLDS ON wRISTS ABSENT; FINgERS LONg, SLENDER, BEARINg ENLARgED DISTAL PADS; FINgER PADS ABOUT 1.8 TIMES wIDTh OF DIgITS AT PENULTIMATE PhALANgE; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FRINgE ON FINgERS ABSENT; OUTER METACARPAL TUBERCLE LOw, OVOID; INNER METACARPAL TUBERCLE OVOID, DIFFUSE; FOREARM wITh SCATTERED TUBERCLES, LARgE, POINTED, VARIOUSLY FUSED TO PRODUCE AN EVIDENT LATERAL FRINgE; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION EXTENDINg TO SNOUT whEN hIND LIMB EXTENDED; TARSAL FOLD PRESENT, wEAkLY DEVELOPED; TIBIA LENgTh 52 % SVL; FOOT LENgTh 47 % SVL; OUTER METATARSAL TUBERCLE ABSENT; INNER METATARSAL TUBERCLE PRESENT, LARgE, OVOID; TOES IV AND V wITh LATERAL FRINgE, PROMINENT ON TOE V; FRINgE OF TOE V EXTENDINg ONTO OUTER SURFACE OF BASE OF TARSUS; DORSAL SURFACE OF hIND LIMB RELATIVELY SMOOTh, BEARINg IRREgULAR SETS OF TUBERCLES ALONg ThREE TRANSVERSE BANDS ON ShANk; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; TOE PADS ABOUT 1.5 TIMES wIDTh OF PENULTIMATE PhALANgE; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh. DORSUM SMOOTh wITh SCATTERED, LOw, DIFFUSE TUBERCLES; FLANkS SMOOTh; VENTRAL SURFACES AND ThROAT SMOOTh wITh SCATTERED LOw, DIFFUSE TUBERCLES; SACRAL DIAPOPhYSES PROjECTINg LATERALLY, SLIghTLY NARROwER ThAN POSTERIORMOST TRANSVERSE PROCESSES OF ALL VERTEBRAE; LATERAL TERMINI OF TRANSVERSE PROCESSES OF PRESACRAL VERTEBRAE VI– VIII NOT CO-OSSIFIED wITh SkIN; FIVE NEURAL SPINES PROjECTINg DORSALLY, CO-OSSIFIED wITh SkIN, VISIBLE EXTERNALLY. PUPIL hORIzONTAL. ALL DORSAL SURFACES UNIFORMLY BROwN (FIg. 4). ThIgh BEARINg TwO, ShANk BEARINg ThREE INDISTINCT TRANSVERSE BANDS OF IRREgULAR, BROwN MOTTLINg ON DORSAL SURFACES, CORRESPONDINg wITh ThREE BANDS OF TUBERCLES ON ThESE BANDS; TOES I AND II ARE UNIFORMLY BEIgE, TOES III–V UNIFORMLY BROwN; FOREARM DARk BROwN, BEARINg BROAD, BROwN, TRANSVERSE BAND ON DORSAL SURFACE; DORSAL SURFACE OF MANUS MORE PALE ThAN FOREARM, ULTIMATE AND PENULTIMATE PhALANgES OF FINgERS I AND II BEIgE, FINgERS III AND IV DARk BROwN. UPPER AND LOwER LIPS wITh NARROw, TRANSVERSE, DARk BROwN BANDS, NOT EXTENDINg ONTO PROXIMAL SURFACES. FLANkS BEIgE wITh DIFFUSE BROwN PUNCTATIONS; VENTER BEIgE wITh SCATTERED DIFFUSE, FINE BROwN MARkINgS; ThROAT DARk BROwN, EXTENDINg ONTO PECTORAL REgION. VENTRAL SURFACES OF FOREARMS BEIgE wITh FINE, BROwN MOTTLINg; VENTRAL SURFACES OF hANDS BEIgE. VENTRAL SURFACES OF hIND LIMB AND FOOT BEIgE wITh FINE, PALE BROwN MOTTLINg. Variation. CONSISTENT SEXUAL DIMORPhISM IN COLORATION OR SkIN TEXTURE wAS NOT EVIDENT (FIgS. 4, 8). GRANULATION ON ThE ThROAT IN Hemiphractus kaylockae IS VARIABLE AMONg SPECIMENS EXAMINED, VARYINg FROM SMOOTh (E.g., KU 93506, 93507, 107411, 107417, 154992) TO SMOOTh wITh TUBERCLES (E.g., AMNH 92668, KU 107419), SMOOThLY gRANULAR wITh NO TUBERCLES (E.g., KU 107412, 107413), TO SMOOThLY gRANULAR wITh TUBERCLES PRESENT (E.g., KU 107408, 107414, 107422, 107423, 154991, UMMZ 152624). GRANULATION OF ThE VENTER RANgES FROM SMOOTh TO UNIFORMLY gRANULAR (E.g., KU, 107411, 107422, 154992, UMMZ 152624); SOME SPECIMENS wITh SMOOTh OR UNIFORMLY gRANULAR VENTERS ALSO hAD VARIABLE NUMBERS OF DISCRETE TUBERCLES PRESENT (E.g., AMNH 92668, KU 93506, 93507, 107412, 107413, 107414, 107417, 107419, 107423, 154991). ThROAT COLORATION IN MOST SPECIMENS IS TAN (E.g. KU 107417) TO BROwN OR DARk BROwN (E.g., AMNH 92668, KU 93506, 107408, 107411, 107412, 107419, 107422, 154992) wITh SOME ShOwINg PROMINENT whITE MOTTLINg, STRIPINg, AND/OR PROMINENT LABIAL BARRINg (E.g. KU 93506, 107412, 107422, 154992) AND KU 93507, 107413, 107414, 107417, 107423, 154991, UMMZ 152624 DISPLAYINg STIPPLINg OR SPECkLINg OF BROwN AND whITE. VENTRAL COLORATION IN MOST SPECIMENS TYPICALLY IS A CONTINUATION OF ThE gULAR COLORATION OF DARk BROwN OR TAN EXTENDINg TO APPROXIMATELY MID-ABDOMEN whERE IT ABRUPTLY ChANgES TO A BEIgE COLORATION ThAT TYPICALLY CONTAINS VARIABLE AMOUNTS OF DARkER BROwN STIPPLINg (E.g., AMNH 92668, KU 93507, 107408, 107411, 107414, 107417, 107419, 107423, 154992); KU 93506 AND KU 107413 ShOw A SIMILAR PATTERN hOwEVER ALSO DISPLAY whITE MOTTLINg IN ThE PECTORAL REgION whILE KU 107422 DISPLAYS whITE SPOTTINg POSTERIORLY. ThE DORSUM OF H. kaylockae IS SMOOTh OR UNIFORMLY SMOOThLY gRANULAR IN ALL SPECIMENS wITh KU 93506, 107423, 154991, 154992, AND UMMZ 152624 hAVINg SOME TUBERCLES. DORSAL COLORATION IN MOST IS AN OVERALL TAN OR BEIgE ThAT MAY OR MAY NOT INCLUDE SOME LEVEL OF DARkER BROwN STIPPLINg OR MOTTLINg (E.g. KU 107408, 107412, 107413, 107417, 107419, 107422, 107423, 154991, 154992) whILE SOME SPECIMENS DISPLAY AN OVERALL UNIFORM BROwN COLOR (E.g. AMNH 92668, KU 93506, 93507, 107411, 107414). SOME SPECIMENS ShOw A PROMINENT DARk BAND ON ThE FOREARM (E.g. KU 107408, 107412, 107422, 107423, 154992). HOwEVER, ThIS BAND IS DIFFUSE OR wEAkLY DEVELOPED IN ALL OThER SPECIMENS. TwO OR ThREE DARk TRANSVERSE BANDS ARE PRESENT ON ThE FEMORAL AND TIBIAL REgIONS (ABSENT OR BARELY DISCERNIBLE ON AMNH 92668, KU 93506, 107411, 107419, 154991), AND A DIFFUSE OR VERY EVIDENT DARk BAND OR ChEVRON PRESENT ON ThE SACRAL REgION (E.g., KU 107408 [wEAkLY DEVELOPED], 107412, 107414, 107417, 107422, 154992, UMMZ 152624); IN SPECIMENS KU 107422 AND 154992 ThERE ARE TwO ChEVRONS PRESENT; IN ADDITION, SPECIMENS KU 107414, 107417, 107422, AND 154992 hAVE A DARk IRREgULAR VERTEBRAL STRIPE EXTENDINg FROM ThE SACRAL BAND CONNECTINg TO A DARk BLOTCh AT ThE OCCIPUT. ThE POSTERIOR SURFACE OF ThE ThIgh OF KU 107412–14 AND UMMZ 152624 BEARS A PATTERN OF TAN OR BROwN MOTTLED wITh whITE OR BEIgE whILE ALL OThER SPECIMENS BEAR A DIFFUSE (E.g., KU 93506, 93507, 107411, 107422, 107423, 154991, 154992) OR DISTINCT whITE STRIPE ALONg ThIS SURFACE (E.g., AMNH 92668, KU 107408, 107417, 107419). Etymology. ThIS SPECIES IS NAMED IN hONOR AND MEMORY OF JULIA KAYLOCk. ShE wAS A COLLEAgUE AND FRIEND, BELOVED BY MANY, whO wAS PASSIONATE ABOUT ThE CONSERVATION OF ThE wORLD’S BIODIVERSITY, ESPECIALLY ThAT OF AMPhIBIANS. ShE SPENT ThE LATTER PART OF hER LIFE wORkINg TIRELESSLY TO IMPROVE AND DOCUMENT hUSBANDRY TEChNIqUES FOR ThREATENED PANAMANIAN AMPhIBIANS (E.g., HILL et al. 2012) MAINTAINED IN ex situ FACILITIES. ShE LOST A LIFELONg BATTLE wITh TYPE I DIABETES AT ThE AgE OF 28 ON 12 JUNE 2009. WE gRACIOUSLY ACkNOwLEDgE hER PASSION FOR AMPhIBIAN CONSERVATION IN ThE NAMINg OF ThIS SPECIES OF Hemiphractus. ThE SPECIFIC EPIThET IS ThE SINgULAR gENITIVE CASE OF ThE NAME KAYLOCk. Distribution and ecology. SPECIMENS OF Hemiphractus kaylockae hAVE BEEN COLLECTED ONLY IN ONE REgION OF FAR EASTERN PANAMA, IN ThE SERRANíA DE PIRRE, PROV. DARIEN (FIg. 5 ). ThIS REgION REMAINS POORLY DOCUMENTED BIOLOgICALLY, AND ThIS SPECIES IS EXPECTED TO OCCUR SOMEwhAT MORE BROADLY—AT LEAST UNTIL ThE RECENT INVASION OF ThE AMPhIBIAN PAThOgEN Batrachochytrium dendrobatidis. ThIS FROg hAS BEEN FOUND ONLY IN MOIST, LOwER MONTANE FORESTS AT ELEVATIONS FROM 900 TO 1550 M (MYERS 1966). SPECIMENS COLLECTED AT NIghT AT CERRO PIRRE (PROV. DARIEN) BY MYERS (1966: 70) wERE OBSERVED “…ON LOw PERChES (LOg, STICkS, DEAD PALM FRONDS) ONLY ONE-hALF TO 2 FEET ABOVE gROUND, AND TwO wERE ON PALM FRONDS 3 TO 4 FEET ABOVE ThE FOREST FLOOR. MYERS (1966) ALSO PROVIDED IN ThIS ACCOUNT A DESCRIPTION OF A MOUTh gAPINg BEhAVIOR IN whICh ThE YELLOwISh ORANgE TONgUE IS DISPLAYED FOLLOwED BY A PAINFUL BITE wITh CONTINUED AgITATION AS wELL AS A “ LOUD CATLIkE SqUALL EMITTED whEN REMOVINg AN INDIVIDUAL FROM A BAg (FIg. 8C). IT IS LIkELY ThAT FEMALES OF ThIS SPECIES CARRY EggS ON ThE BACk AS hAS BEEN DOCUMENTED FOR ALL OThER DESCRIBED SPECIES OF Hemiphractus. AS ALL DOCUMENTED OBSERVATIONS hAVE OCCURRED AT NIghT, IT IS INFERRED ThAT ThIS SPECIES IS MOST LIkELY NOCTURNAL.

Hemiphractus panamensis (FIgS. 3, 4, 6, 7, 9) Cerathyla panamensis Stejneger 1917. Holotype: USNM 55320. Type locality: Signal Loma (Loma Peak) on the north coast of Panama, three miles [4.8 km] south of Santa Isabel, Prov. Col��n, Panama. Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from Chagres Highlands, Panama]; Duellman, 2001 [in part; for reference to specimens from Chagres Highlands, Panama]; Crawford et al. 2012 [in part, for samples from Cerro Brewster, Prov. Panama, Panama]; Castroviejo-Fisher et al. 2015 [in part, for samples from Cerro Bruja and Cerro Brewster, Prov. Panama, Panama]; K��hler, 2011 [figs. 472, 473; in part; for map showing isolated populations in the Chagres Highlands, Panama]. Re-description of the Holotype. MEDIUM-SIzED ADULT FEMALE, SVL 59.3 MM (NOT INCLUDINg PROBOSCIS); TIBIA LENgTh 31.2 MM (NOT INCLUDINg CALCAR); FOOT LENgTh 27.3 MM; hEAD LENgTh 26.6 MM (NOT INCLUDINg PROBOSCIS); hEAD wIDTh 31.4 MM; SNOUT���PARAOCCIPITAL PROCESS LENgTh 25.8 MM; TYMPANUM OVOID, 5.2 MM hIgh X 2.6 MM wIDE; EYE DIAMETER 7.4 MM; INTERORBITAL DISTANCE 12.0 MM; EYE���TYMPANUM DISTANCE 3.3 MM; BODY MODERATE; hEAD LARgE, wIDER ThAN BODY, gREATEST wIDTh AT LEVEL OF PARS ARTICULARIS, BEARINg PROMINENT PARAOCCIPITAL PROCESSES; PARAOCCIPITAL PROCESSES PROjECTINg POSTEROLATERALLY IN DORSAL VIEw, EXTENDINg POSTERIOR TO LEVEL OF PARS ARTICULARIS; SkIN CO-OSSIFIED OVER DORSAL SURFACE OF SkULL; OCCIPITAL RIDgES INDISTINCT, PARAOCCIPITAL RIDgES PRESENT, wEAkLY DEVELOPED; SUPRAORBITAL AND CANThAL RIDgES SLIghTLY DEVELOPED; zYgOMATIC AND PREORBITAL RIDgES ABSENT; CANThUS ROSTRALIS INDISTINCT; LOREAL REgION FLAT; PROBOSCIS ShORT, FLEShY, POINTED, TRIANgULAR IN CROSS SECTION; SNOUT ANgLINg POSTERIORLY AT APPROXIMATELY 45o FROM hORIzONTAL; NOSTRILS OVOID, SLIghTLY PROTUBERANT, PROjECTINg DORSALLY, SLIghTLY LATERALLY; LIPS NOT FLARED. TOP OF hEAD RELATIVELY FLAT; INTERORBITAL REgION 38 % hEAD wIDTh; EYE DIAMETER 24 % hEAD wIDTh; gREATEST DIAMETER 70 % TIMES EYE DIAMETER; wIDTh OF TYMPANUM 84 % TIMES EYE���TYMPANUM DISTANCE; UPPER EYELID APPEARINg TO BEAR ONLY A SINgLE ENLARgED POINTED PROjECTION, SCATTERED SMALLER TUBERCLES ABSENT; INTERORBITAL, POSTORBITAL TUBERCLES ABSENT; SkIN OVER POSTERIOR SURFACE OF PARS ARTICULARIS BEARINg ONE LARgE, FLEShY TUBERCLE. TONgUE LARgE, OVOID; MANDIBLE BEARINg SERRATIONS ALONg ENTIRE LENgTh OF LOwER jAw, wITh TwO PROMINENT, TUSk-LIkE ODONTOIDS PROjECTINg FROM MENTOMECkELIAN BONES. AXILLARY MEMBRANE ABSENT; ThORACIC FOLD ABSENT; DERMAL FOLDS ON wRISTS ABSENT; FINgERS LONg, SLENDER, BEARINg SLIghTLY ENLARgED DISTAL PADS; INNER OUTER METACARPAL TUBERCLES PRESENT, BUT CONDITION OF SPECIMEN DOES NOw ALLOw FOR ACCURATE DESCRIPTION; RELATIVE LENgThS OF FINgERS: III> IV> II> I; LATERAL FLANgES APPEAR TO BE ABSENT ON FINgERS; CONDITION OF SPECIMEN DOES NOT PERMIT DESCRIPTION, OR DETERMINATION OF PRESENCE/ ABSENCES OF TUBERCLES ON FOREARMS; wEBBINg ON hANDS ABSENT. HEELS OF ADPRESSED hIND LIMBS OVERLAP; TIBIOTARSAL ARTICULATION NOT EXTENDINg TO SNOUT whEN hINDLIMB EXTENDED; CONDITION OF SPECIMEN DOES NOT PERMIT DESCRIPTION OF TARSAL FOLDS; TIBIA LENgTh 53 % SVL; FOOT LENgTh 46 % SVL; INNER AND OUTER METATARSAL TUBERCLES PRESENT, BUT CONDITION IS UNCLEAR; PRESENCE OF CONDITION OF LATERAL FRINgE ON TOES UNCLEAR; CALCAR PRESENT, SMALL; RELATIVE LENgThS OF TOES: IV> V> III> II> I; wEBBINg BASAL. CLOACAL OPENINg DIRECTED POSTEROVENTRALLY AT MIDLEVEL OF ThIgh. TEXTURE AND COLORATION OF ThE SkIN, AS wELL AS ThE FINER DETAILS OF ThE hANDS AND FEET ARE UNCLEAR, AS ThE TYPE SPECIMEN IS qUITE DESICCATED. SIMILARLY, ThE DESICCATED NATURE OF ThE hOLOTYPE MAkES IT UNCLEAR ThE LEVEL TO whICh ThE TRANSVERSE PROCESSES OF ThE VERTEBRAE ARE CO-OSSIFIED wITh ThE SkIN. ThE PUPIL IS NOT VISIBLE. IN ALL MAjOR MORPhOLOgICAL FEATURES AND MEASUREMENTS, STEjNEgER���S (1917) DESCRIPTION CLOSELY MATChES ThE CURRENT CONDITION OF ThE hOLOTYPE (USNM 55330). EXCEPTIONS INCLUDE COLORATION NOTES AND TEXTURAL DETAILS OF ThE SkIN, VARIOUS TUBERCLES, AND DIgITAL PADS (STEjNEgER 1917: 32), ALL OF whICh ARE NOw FADED OR PhYSICALLY DISTORTED���EVIDENTLY AS A RESULT OF DESICCATION AT SOME POINT IN ITS hISTORY. LIkEwISE, ThE INDENTATIONS OF EggS ON ThE DORSAL SURFACE OF ThIS FEMALE ThAT STEjNEgER (1917: 33) DESCRIBED NO LONgER ARE EVIDENT. Diagnosis. Hemiphractus panamensis MAY BE DISTINgUIShED FROM ALL SPECIES OF Hemiphractus EXCEPT H. elioti, H. fasciatus, H. kaylockae, AND H. scutatus BY NOT hAVINg A POSTORBITAL INDENTATION (PRESENT IN OThER SPECIES). Hemiphractus panamensis MAY BE DISTINgUIShED FROM H. scutatus, AN AMAzONIAN SPECIES NOT OCCURRINg IN PANAMA, BY ITS SMALLER SIzE (MAXIMUM SVL IN MALES 52.5 VS. 57.4, FEMALES 64.7 VS. 80.5; DATA FOR H. scutatus FROM TRUEB 1974), AND BY hAVINg SMALL EXPANDED PADS ON ThE FINgERS AND TOES (ABSENT IN H. scutatus). Hemiphractus panamensis DIFFERS FROM H. elioti AND H. kaylockae BY hAVINg NEOPALATINE AND VOMERINE ODONTOIDS wIDELY SEPARATED (VS. IN CONTACT IN H. elioti AND NARROwLY SEPARATED IN H. kaylockae; FIg. 6). Hemiphractus panamensis APPEARS TO hAVE ONLY ABOUT TwO ODONTOIDS ON EACh VOMER, whEREAS H. elioti AND H. kaylockae BOTh hAVE ABOUT SEVEN (CONDITION UNkNOwN IN H. scutatus; FIg. 6). Hemiphractus panamensis MAY BE DISTINgUIShED FROM H. kaylockae BY hAVINg A STRAIghT LATERAL MARgIN OF ThE PARAOCCIPITAL hORNS (FIgS. 2, 9); SUPRAORBITAL RIDgES EVIDENT, wEAkLY DEVELOPED (ABSENT OR wEAkLY DEVELOPED IN H. elioti AND H. kaylockae; FIgS. 2, 9); CANThAL RIDgES EVIDENT, DISTINCT (ABSENT OR wEAkLY DEVELOPED IN H. elioti; FIgS. 2, 9). Hemiphractus panamensis hAS A VERY LARgE SUBTEMPORAL FENESTRA APPROXIMATELY 1.5 X SIzE OF ThE ORBIT IN FEMALES (MALES UNkNOwN), SUBTEMPORAL FENESTRA APPROXIMATELY ONE-ThIRD DIAMETER OF ThE ORBIT IN BOTh MALES AND FEMALES IN H. kaylockae, APPROXIMATELY ONE-hALF DIAMETER OF ORBIT IN FEMALES OF H. elioti (VERY SMALL IN MALES), APPROXIMATELY ONE-hALF DIAMETER OF ThE ORBIT IN FEMALES OF H. fasciatus (MALES UNkNOwN); LATERAL MARgINS OF ThE qUADRATOjUgAL EXTENDINg FAR BEYOND ThE PROFILE OF ThE PARAOCCIPITAL hORNS IN DORSAL VIEw (ABOUT TO LEVEL OF PARAOCCIPITAL hORNS, OR SLIghTLY BEYOND, H. elioti, H. fasciatus, AND H. kaylockae). Distribution and ecology. Hemiphractus panamensis IS kNOwN FROM SPECIMENS COLLECTED FROM CENTRAL PANAMA IN ThE PROVINCES OF COLON AND SAN BLAS wIThIN OR IN PROXIMITY TO whAT IS NOw ThE ChAgRES NATIONAL PARk (FIg. 5). SPECIMENS hAVE BEEN COLLECTED AT ~750 M ELEVATION IN LOwER MONTANE FORESTS wITh hIgh RAINFALL (MYERS 1966). TRUEB (1974) REPORTED A LARgE SPECIMEN (KU 93503) ThAT CONTAINED ������AN UNIDENTIFIED gASTROPOD, A LIzARD (Ptychoglossus), AND A DENDROBATID FROg (Colostethus) IN ThE STOMACh. MYERS (1966) REPORTED H. panamensis TO DISPLAY MOUTh gAPINg BEhAVIOR AND BITINg whEN ACCOSTED (FIg. 3B). AN INDIVIDUAL COLLECTED BY MYERS IN 1964 AT ALTOS DE PACORA (PANAMA PROVINCE) wAS ���ON A STICk LYINg ON BARE SOIL, IN A ROADSIDE DITCh AT NIghT. TwO LIVE SPECIMENS OBSERVED AT NIghT BY T. HERMAN AND COLLEAgUES (PERS. COMM.) AT CERRO BREwSTER (PANAM PROVINCE) wERE OBSERVED ON VEgETATION Remarks. WhILE OSTEOLOgICAL ChARACTERS OF ThE SkULL wOULD NOT SEEM TO BE USEFUL DIAgNOSTIC ChARACTERS IN ThE ABSENCE OF SkELETAL SPECIMENS OR RADIOgRAPhY, wE NOTE ThAT ThE DISTINCTLY ENLARgED SUBTEMPORAL FENESTRA OF H. panamensis CAN BE DETECTED BY MANUAL PALPATION AND IS ALSO VISIBLE EVEN IN PhOTOgRAPhS OF LIVINg SPECIMENS. DESPITE ThE RELATIVE PROXIMITY OF ThE RANgE OF ThIS SPECIES TO PANAMA CITY, AND INFRASTRUCTURAL DEVELOPMENTS SUCh AS ThE PANAMA CANAL AND VARIOUS SMIThSONIAN FIELD STATIONS, RELATIVELY FEw SPECIMENS OF ThIS SPECIES APPEAR TO EXIST IN MUSEUM COLLECTIONS. AS A RESULT, DATA REgARDINg VARIATION IN COLORATION AND MORPhOMETRY ARE UNAVAILABLE., Published as part of Hill, Robert L., Martin, Kathryn G., Stanley, Edward & Mendelson Iii, Joseph R., 2018, A taxonomic review of the genus Hemiphractus (Anura: Hemiphractidae) in Panama: Description of Two New Species, Resurrection of Hemiphractus panamensis (Stejneger, 1917), and Discussion of Hemiphractus fasciatus Peters, 1862, pp. 495-512 in Zootaxa 4429 (3) on pages 506-509, DOI: 10.11646/zootaxa.4429.3.3, http://zenodo.org/record/1285044, {"references":["Stejneger, L. (1917) A new species of horned tree-toad from Panama. Proceedings of the Biological Society of Washington, 301, 31 - 34.","Trueb, L. (1974) Systematic relationships of Neotropical horned frogs, genus Hemiphractus (Anura: Hylidae). Occasional Papers of the Museum of Natural History, University of Kansas, 29, 1 - 60. https: // doi. org / 10.5962 / bhl. part. 11750","Duellman, W. (2001) The Hylid Frogs of Middle America. Vol. 2. Society for the Study of Amphibians and Reptiles Contributions in Herpetology, Ithaca, New York, 1170 pp.","Crawford, A., Cruz, C., Griffith, E., Ross, H., Ibanez, R., Lips, K. & Crump, P. (2012) DNA barcoding applied to ex situ tropical amphibian conservation programme reveals cryptic diversity in captive populations. Molecular Ecology Resources, 13, 1005 - 1018.","Castroviejo-Fisher, S., Padial, J., de La Riva, I., Pombal Junior, J., da Silva, H., Rojas-Runjaic, F. J. M., Medina-Mendez, E. & Frost, D. (2015) Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development. Zootaxa, 4004 (1), 1 - 75. https: // doi. org / 10.11646 / zootaxa. 4004.1.1","Myers, C. (1966) The distribution and behavior of a tropical horned frog Cerathyla panamensis Stejneger. Herpetologica, 22, 68 - 71."]}

We reviewed the taxonomic status of populations of frogs in the genus Hemiphractus in Panama, which have all been referred to Hemiphractus fasciatus Peters, 1862 for over 40 years. Although relatively few specimens have been collected, mostly juveniles, it is clear that these frogs inhabit three separate upland regions of the country: The Cordillera de Talamanca in western Panama, the Chagres Highlands and Cordillera de San Blas in central Panama, and the Serranía de Pirre in the far eastern portion of the country. In accordance with previously published molecular data, we identified distinctive features of the skulls of frogs representing these three allopatric populations and herein revalidate H. panamensis (Stejneger, 1917), describe the new species Hemiphractus elioti sp. nov. from the Cordillera de Talamanca, and the new species Hemiphractus kaylockae sp. nov. from the Serranía de Pirre. We also propose that the taxon H. fasciatus is a South American species not occurring in Panama.

The amphibian fauna of the state of Rio de Janeiro, in southeatern Brazil, is characterized by high species richness and rates of endemism, and is still insufficiently known. A first list of amphibian species with occurrence in the state was published in 2004 and reported 166 taxa, but since then many new records, descriptions of new taxa, and revalidations and synonymizations of species have consistently improved the knowledge about the state’s amphibian biodiversity. Thus, a review and update of that list was deemed necessary. We herein present an updated and commented list of amphibian species occurring in the state of Rio de Janeiro based on a survey of the literature. We recorded the occurrence of a total of 201 species of amphibians (197 anurans and four caecilians) in Rio de Janeiro, with 54 of them (ca. 27%) considered to be endemic of the state. Our study presents an increase in species richness of 21% since the publication of the previous list, indicating a consistent advance in knowledge of the composition of the amphibian fauna in the state. In spite of its relatively small territorial extension (total area ca. 43,800 km²), the state of Rio de Janeiro contains nearly 20% of the amphibian species known to occur in Brazil and around 40% of those occurring in the Atlantic Forest biome. Thus, that state constitutes an important reservoir of amphibian biodiversity in the Atlantic Forest biome and in Brazil, as a whole.

Family Hemiphractidae Fritziana ulei was resurrected from synonymy with Fritziana fissilis by Folly et al. (2014)., Published as part of Dorigo, Thiago Arnt, Vrcibradic, Davor & Rocha, Carlos Frederico Duarte, 2018, The amphibians of the state of Rio de Janeiro, Brazil: an updated and commented list, pp. 1-11 in Pap��is Avulsos de Zoologia 58 on page 6, DOI: 10.11606/1807-0205/2018.58.05, http://zenodo.org/record/4614090