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As more land is altered by human activity and more species become at risk of extinction, it is essential that we understand the requirements for conserving threatened species across human-modified landscapes. Owing to their rarity and often sparse distributions, threatened species can be difficult to study and efficient methods to sample them across wide temporal and spatial scales have been lacking. Passive acoustic monitoring (PAM) is increasingly recognized as an efficient method for collecting data on vocal species; however, the development of automated species detectors required to analyse large amounts of acoustic data is not keeping pace. Here, we collected 35 805 h of acoustic data across 341 sites in a region over 1000 km2 to show that PAM, together with a newly developed automated detector, is able to successfully detect the endangered Geoffroy's spider monkey (Ateles geoffroyi), allowing us to show that the Geoffroy's spider monkey was absent below a threshold of 80% forest cover and within 1 km of primary paved roads and occurred equally in old growth and secondary forests. We discuss how this methodology circumvents many of the existing issues in traditional sampling methods and can be highly successful in the study of vocally rare or threatened species. Our results provide tools and knowledge for setting targets and developing conservation strategies for the protection of the Geoffroy's spider monkey.

The Amazon is one of the most diverse biomes around the globe, currently threatened by economic and industrial development and climate change. Large mammals are keystone species, playing an important role in ecosystem structure and function as ecological engineers, while being highly susceptible to deforestation, habitat degradation, and human exploitation. Using a unifying hierarchical Bayesian spatial approach, we examine the site-use patterns of four large Amazonian Forest mammals and their relationships to anthropogenic factors at a biome-wide scale. Our results showed that species’ patterns of site use are correlated with human induced habitat changes, and that this correlation is species-specific. The white-lipped peccary shows highest site-use estimates within strict protected areas, affected by proximity to urban areas and benefiting from indigenous territories, the tapir responding slightly to proximity to burned forested areas, while the giant armadillo and the jaguar were primarily affected by vegetation cover loss; disturbances related to the colonization of the Amazon. Our findings contribute to the understanding of how human-induced environmental changes influence the site-use patterns of these four large mammals, and inform future conservation and land use planning. Transboundary conservation efforts, empowering and integrating native (indigenous and non-indigenous) communities in land governance schemes, involving the private sector and securing the commitment of developed countries are important paths for the protection and sustainability of the globally-crucial Amazon rainforest.

Supplemental Material for Indigenous Lands are Better for Amphibian Biodiversity Conservation Than Immigrant-Managed Agricultural Lands: A Case Study From Manu Biosphere Reserve, Peru by Shirley J. Serrano-Rojas, Andrew Whitworth, Julio A. Paredes-Garcia, Ruthmery Pillco-Huarcaya, Lawrence Whittaker, Karl H. Huaypar-Loayza, and Ross MacLeod in Tropical Conservation Science

Supplemental material, sj-pdf-1-trc-10.1177_19400829211028118 for Unexpected Diversity in Regenerating Sites Stresses the Importance of Baselines: A Case Study With Bats (Order Chiroptera) on the Osa Peninsula, Costa Rica by Elène Haave-Audet, Doris Audet, Michelle Monge-Velazquez, Eleanor Flatt and Andrew Whitworth in Tropical Conservation Science

Riparian zones are one of the most productive ecosystems in the world, but are at risk due to agricultural expansion and climate change. To maximize return on conservation investment in mixed-use landscapes, it is important to identify the minimum intact riparian forest buffer sizes to conserve riparian ecosystem services. The minimum riparian forest buffer width necessary to maintain tropical river water quality remains unclear, and there is little analysis of effective riparian buffer lengths. Also, in studies on the effect of land use on river water quality globally, there is little standardization in the area where land use is analyzed. Here, these challenges were addressed in the Osa Peninsula in southwestern Costa Rica. Water quality parameters and social variables were sampled at 194 locations across the region. For each sample, land use was calculated in nine different riparian buffer sizes and at the sampling location. Riparian forest cover had a positive effect on water quality parameters, while agricultural cover had a negative effect. The longer the length of the buffer considered, the greater the relative support for influencing water quality (1000 m > 500 m > 100 m). All buffer widths yielded similar support within each length class. These results indicate that length of riparian forest buffers, not width, drives their ability to conserve water quality. While wide and long riparian forests are ideal to maximize the protection of river water quality and other ecosystem services, in landscapes where that is impractical, the 15-m-wide riparian forest buffers that are supported by Costa Rican legislation could improve water quality, providing that they are at least 500 m long. The results also indicate the importance of methodological standardization in studies that monitor land use effects on water quality. The authors propose that studies in similar regions analyze land use in riparian zones 15-m-wide by 1000 m upstream. Conserving and restoring narrow, long riparian forest buffers could provide a rapid, economical management approach to balance agricultural production and water quality protection.

Euspondylus excelsum was described from 8 specimens from the Region Huánuco in central Peru. We obtained an adult female from the Region Junín and a subadult male (photo vouchers only) from the Region Madre de Dios. Our new records of E. excelsum extend the range of this species by 592 km to the south and increase the previously known elevational range from 1023–1159 m a.s.l. to 550–1550 m a.s.l. The adult female is the largest known specimen extending the maximum SVL to 90.1 mm. Our findings indicate that E. excelsum is relatively widely distributed in the montane forests and foothills of the eastern Andes. Its arboreal mode of life is probably one reason for its rarity and limited records during traditional field surveys.

Ameerega shihuemoy sp. nov. (Figures 1–3, 6) Cryptophyllobates sp: Chaparro & Ochoa 2005 p.7 (MHNC 4779 collected on 0 7 December 2004 by J. C. Chaparro & J. A. Ochoa at Erika Lodge, Departamento Madre de Dios). Ameerega gr. pictus: Chaparro et al. 2016 p. 2 (from Amarakaeri Communal Reserve, Departamento Madre de Dios). Ameerega sp1: Whitworth & Villacampa 2014 p. 3 (from Manu Learning Centre, Departamento Madre de Dios). Holotype. MHNC 15488 (Fig. 1), an adult female, from near Cupudnoe river, 440 m elevation, Peru (Coordinates: 12°47'26.70" S, 70°58'21.30" W), Distrito Madre de Dios, Provincia Manu, Departamento Madre de Dios, collected on 25 April 2010 by R. C. Gutiérrez. Paratopotypes. Seven specimens: Five adult females (MUSA 3178, MUSA 3180–3182, MHNC 12988), and two immature females (MUSA 3177, MUSA 3 179), collected with the holotype by R. C. Gutiérrez. Paratypes. Fifteen specimens, all from Provincia Manu, Departamento Madre de Dios, Peru. Two adult females (MHNC 10406–10407), and one adult male (MHNC 10408), from the Reserva Comunal Amarakaeri, 32 km east of Huepetuhe, near the headwaters of Colorado river, 846 m elevation, (Coordinates: 12°59'59.25'' S, 70°50'31.32'' W), collected on 0 7 November 2010 by J. Delgado and J. G. Estrada. Two adult females (MHNC 10525–10526), and two adult males (MHNC 10536, MHNC 10542), from the Reserva Comunal Amarakaeri, 21.5 km southwest of Shintuya, near Azul river, 480 m elevation, (Coordinates: 12°48'50.8'' S, 71°06'07.8'' W), collected on 23 May 2011 by J. Delgado. An adult male (MHNC 15863), from the Reserva Comunal Amarakaeri, Pad A, Lote 76, near Dahuene river, 530 m elevation, (Coordinates: 12°58'38.47'' S, 71°01'30.73'' W), collected on 20 July 2016 by R. Coronel and G. Valencia. Two immature specimens (MHNC 10806, MHNC 10811), from 3.5 km south of Shintuya, near Serjali river, 450 m elevation, (Coordinates: 12°42'49.02'' S, 71°15'17.51'' W), collected on 20 January 2011 by J. G. Estrada. Two immature specimens (MHNC 10816, MHNC 10826), from 10 km southwest of Itahuania, near Shilive river, 450 m elevation, (Coordinates: 12°43'02.93'' S, 71°09'07.56'' W), collected on 26 January 2011 by J. G. Estrada. An immature female (MHNC 14561; Figs. 2A, B), from the Reserva Comunal Amarakaeri, Pad A, Lote 76, near Colorado river, 825 m elevation, (Coordinates: 12°59'17.97'' S, 71°0'52.67'' W), collected on 0 4 February 2015 by T. Gregory. An adult specimen (MHNC 5012), from Aguas Calientes, 2.75 km east (downstream) from Shintuya, (Coordinates: 12°40'7.27" S, 71°16'12.20'' W), collected on 0 9 July 2014 by S. J. Serrano. An immature specimen (MHNC 4779; Figs. 2 C, D), from Erika Lodge, 500 m elevation, (Coordinates: 12°45'23.53" S, 71°22'48.68" W), collected on 0 7 December 2004 by J. C. Chaparro and J. A. Ochoa. Referred specimens. Five specimens: MUSM 31611, MUSM 31664, MUSM 31691, MUSM 31692 and MUSM 31730, from Manu Learning Centre Research Station, 460 m elevation, Peru, (Coordinates: 12°47'21.849" S, 71°23'28.06" W), Distrito Manu, Provincia Manu, Departamento Madre de Dios, Peru, collected on 12 February, 0 5 July, 21July, and 25 July 2012, and on 0 3 February 2013, respectively, by A. Whitworth. Etymology. The specific name shihuemoy (English pronunciation: shee-way-moy) corresponds to the Harakmbut word for "poison dart frog". The Amarakaeri are aboriginals from Amazonian Peru and their language belongs to the Harakmbut linguistic group. They coexist with the new species. Diagnosis and comparisons with other species. Following Grant et al. (2006), the new species is assigned to the genus Ameerega on the basis of the following characters: dorsal skin finely granular; Finger I> Finger II when adpressed; toe and finger webbing absent; presence of bright flash marks and absence of ventrolateral line. This is a medium species of Ameerega with an adult SVL of 23.3 ± 2.7 (19.3–30.4 mm, N = 25). Vocal slits present; snout subacuminated, short and protruding. The head and dorsal surfaces are blackish with bronze-brown shades. The flanks are black with metallic bluish-green blotches, with a distinct bright orange or coral stripe extending from groin to above the eye and forward along the canthus rostralis to join around the snout. A flash spot is observed on the hip and upper surface of thighs, and a pale orange spot is located on the anterodorsal base of the thighs. It lacks concealed axillary and calf spots. The ventral surfaces have black reticulations with blue. The maxillary arch has teeth and the dentigerous process of vomers is lacking. The choanae are partly concealed by the palatal shelf of maxilla. The finger discs are weakly to moderately expanded, and when adpressed the first finger is slightly longer than second, and the fourth is shorter than first and second fingers, while the third is longer than all others. The finger discs are weakly to moderately expanded. The finger disc of the third finger of adults is about 1.2 times wider than the immediately adjacent part of the finger. The third finger shows no sexual dimorphism. Basal webbing is present on toes II–IV. Morphologically, divergence of A. shihuemoy from other species is supported by a unique combination of characters: the lack of the conspicuous axillary, thigh and calf flash marks, characteristics of many other species of Ameerega; black granular dorsum with cream to light orange dorsolateral lines, metallic blue venter and underside of extremities with black reticulations and spots. Within Ameerega, it shows the general appearance of A. altamazonica Twomey & Brown 2008, A. boliviana (Boulenger 1902), A. hahneli (Boulenger 1883), A. ignipedis Brown & Twomey 2009, A. petersi (Silverstone 1976), A. picta (Bibron 1838), A. pongoensis (Schulte 1999), A. pulchripecta (Silverstone 1976), A. simulans (Myers, Rodriguez & Icochea 1998), A. smaragdina (Silverstone 1976), and A. yungicola (Lötters, Schmitz & Reichle 2005), with which it shares a granular black to brown dorsum, a light labial bar, a conspicuous dorsolateral line running from the snout to the groin, and a metallic blue belly and underside of arms and hind limbs. From all these species it can be distinguished by lacking flash marks on the axillae, thighs, and calves (absent only in A. boliviana, A. simulans, A. smaragdina, most A. petersi, and some A. pongoensis), by having bright cream to orange dorsolateral stripes (white, pale yellowish-green, intense yellow, or green in all other species but A. picta), and by its blue belly reticulated with black (bluish white and black in A. boliviana, green and blue with black marbling in A. petersi, and green and blue lacking black marbling in A. smaragdina). Measurements (in mm) of the holotype. The female holotype (Fig. 1) has SVL, 25.7; TL, 12.3; GBW, 8.2; HW, 7.9; HL, 8.0; ED, 5.2; IOD, 3.0; TSCN, 2.0; NED, 2.6; IND, 2.9; EL, 3.3; HDT, 2.0; MTD, 0.8; HaL, 7.0; WTFD, 0.7; WTF, 0.5; WTTD, 0.8; WTT, 0.4; WFTD, 0.9; WFT, 0.4. Description of the type series. External morphology: A small to medium-size Ameerega, with adult males attaining approximately 19.2–21.8 mm SVL and adult females approximately 21.5–25.7 mm SVL (measurements and proportions summarized in Table 1). Dorsal skin of head, body, shank, thigh, and hind limbs coarsely and conspicuously granular; skin smooth or nearly smooth on forelimbs, and smooth on sides of head, body and on ventral surfaces. Head slightly wider between jaw articulations than between outer edges of upper eyelids; head usually narrower than body or about as wide. Head width between jaws 29–33% of SVL. Snout sloping, bluntly pointed or rounded in profile, truncate to rounded (usually) or bluntly pointed in dorsal and ventral view. Nares situated near tip of snout, directed slightly posterolaterally; nares visible from front and from below but not from above. Canthus rostralis short, protruding; loreal region nearly vertical, slightly concave to flat. Interorbital distance wider than upper eyelid. Eye shorter than snout length; distance from center of naris to eye 60–87% of eye length. Tympanum 35–65% of eye size. Hand (Fig. 1 F) moderately large, its length 23–28% of SVL and 74–90% of head width between angles of jaws. Relative lengths of adpressed fingers IV> I> II> III; fingers I, II, and IV approximately similar in size when adpressed, with overlapping discs; adpressed first finger varies from slightly shorter to slightly longer than second. Finger discs weakly to moderately expanded; third finger disc 1.2–1.8 times wider than distal end of adjacent phalanx. A large outer metacarpal (= palmar) tubercle on the base of the palm and a smaller inner metacarpal tubercle on the base of first finger, these being relatively small, with rounded surfaces. One subarticular tubercle on fingers I & II, and two subarticular tubercles on fingers III & IV; subarticular tubercles well developed and prominently raised, although distal one on finger III and both in IV sometimes weaker; supernumerary tubercles, finger keels, and outer metacarpal fold absent; tibia length 47–54% of SVL; relative lengths of adpressed toes, IV> III> V> II> I; first toe usually reaching base of subarticular tubercle of second toe; basal webbing on toes II–IV; toe fringes absent; outer metatarsal fold absent; a large inner metatarsal fold located on the distal half of tarsus and extended near the inner metatarsal tubercle (Fig. 1 G); ventrolateral side of tarsus relatively smooth, not especially rugose or tubercular; toes with moderately expanded discs, wider than finger discs (Fig. 1 G); one to three moderately raised subarticular tubercles (one each on toes I & II, two each on III & V, and three on IV); two large metatarsal tubercles with low, rounded surfaces; inner metatarsal tubercle slightly larger than outer metatarsal tubercle; supernumerary tubercles absent. Color pattern in life. (Based on living specimens and photographs; Figs. 2, 3). The noticeably granular body is blackish with bronze shades middorsally, turning black and blue dorsolaterally. The flanks are mostly black with metallic bluish green, with a distinct bright orange, coral, or cream stripe extending obliquely from groin to above the eye, and forward along the canthus rostralis to join around the snout. A pale whitish-bronze labial stripe commences between naris and eye, and extends posteriorly under the eye and tympanum to the base of the upper arm. A bright orange to coral spot is present on the hip and upper thigh; there is no calf spot and no pale markings in axilla or groin. The limbs are bronze-brown with shades of green and black. The ventral surfaces are overall blue with variable black reticulation, while in some individuals the ventral surfaces are overall black with blue reticulation. Color in preservative (Fig. 1). The bright orange and coral colors fade to pale grey; the blue ventral surfaces fade to grey in a dark reticulum of variable distinctiveness. Vocalization. Recordings of three A. shihuemoy males were taken at the Manu Learning Centre reserve. The advertisement call of this new species can be characterized as a series of chirp-like, pulsed notes. Notes are repeated at a rate of 0.8–1.0 notes per second (0.9 ± 0.1), duration of individual notes range from 84–109 ms (98.4 ± 6.8 ms), spaced 969–1196 ms apart (1042.5 ± 186.6 ms), with eight pulses (8.2 ± 0.8) per note. Dominant frequency ranges from 4478.9–4909.6 Hz (4672.72 ± 251.0 Hz) and is not frequency-modulated. Calling activity happens most frequently in the early mornings between 05:00 to 09:00 and late afternoon between 16:00 to 18:00. We also recorded a second call in A. shihuemoy consisting of three notes in quick succession (within 174.9 ± 16.6 ms of each other), repeated once every four to five seconds. The three notes in this call have different duration; the first note (87.2 ± 2.5 ms) typically has a longer duration than the second (69.4 ± 2.6 ms) and the third note (72.8 ± 2.8 ms). This three-note call seems to function as an aggressive or territorial call. We compared the call of the following species with Ameerega shihuemoy: two recordings of A. simulans (74 advertisement calls) by J. M. Padial from Marcapata, Quispicanchis Province, Department Cusco, Peru, were obtained from Fonoteca Zoologica (Museo Nacional de Ciencias Naturales 2016); two recordings of A. boliviana (45 advertisement calls) by J. Bosch & I. De la Riva from Correo-Apolo, La Paz, Bolivia; one recording of A. yungicola (30 advertisement calls) by M. Pacheco-Suarez from Caranavi, Yungas, Bolivia obtained from “Guía fotográfica de los anfibios de la region de las yungas Bolivia ” (Pacheco-Suarez 2015); two recordings of A. hahneli (46 advertisement calls) by J. Serrano-Rojas from Shintuya, Madre de Dios, Peru; two recordings of A. picta (63 advertisement calls) by J. M. Padial from Madidi National Park, Northern Bolivia and from Paractito, Cochabamba Department, Central West Bolivia deposited at Fonoteca Zoologica (Museo Nacional de Ciencias Naturales 2016); two recordings of A. macero (55 advertisement calls) by J. Serrano-Rojas from Manu Learning Centre, Madre de Dios, Peru. The advertisement call of A. shihuemoy is easily distinguished (Fig. 4) from the morphologically similar Ameerega species of the Ameerega picta group (for measurable call parameters and intra- and interspecific variation see Table 2). Parameters A. shihuemoy A. simulans A. picta A. hahneli Mean SD Mean SD Mean SD Mean SD Calls/N° of individuals 80/3 - 74/2 - 63/2 - 46/2 - Notes/min 52.8 5.4 79.2 3.6 130.2 6.0 511.8 4.2 Notes/s 0.9 0.1 1.3 0.1 2.2 0.1 8.5 0.1 Duration of the note (ms) 98.4 6.8 104.9 10.2 45.8 2.4 13.4 2.8 Break between calls (ms) 1042.5 186.6 691.4 162.7 429.9 44.6 107.4 12.6 Fundamental frequency (Hz) 4237.0 281.9 4060.9 74.6 3770.7 76.7 2516.8 83.7 Dominant frequency (Hz) 4672.7 251.0 4460.3 157.7 4044.2 94.7 4550.0 49.1 continued. Parameters A. boliviana A. yungicola A. macero Mean SD Mean SD Mean SD Calls/N° of individuals 45/ 2 - 30 - 55 /2 - Notes/min 72.0 4.2 313.2 0.0 519.4 2.2 Notes/s 1.2 0.1 5.2 0.0 8.7 0.0 Duration of the note (ms) 80.9 8.6 47.8 4.7 37.6 1.3 Break between calls (ms) 783.2 88.6 148.1 7.3 76.1 2.7 Fundamental frequency (Hz) 3416.1 68.2 3475.7 43.5 3353.7 38.1 Dominant frequency (Hz) 3846.0 46.3 3703.7 0.0 3617.6 0.0 The note duration of the new species (84–109 ms) is longer than A. hahneli (11–18 ms), A. macero (36–40 ms), A. picta (41–52 ms) and A. yungicola (40–57 ms), but similar to A. boliviana (75–94 ms) and A. simulans (91–118 ms). The space between calls in A. shihuemoy is longer than in the other six species. The number of notes per second in the advertisement call of A. shihuemoy is lower than the other six species. The call of the new species has a higher fundamental frequency than all other species but similar to A. simulans, and a higher dominant frequency than all other species but similar to A. hahneli and A. simulans, except the latter species has pronounced upward frequency-modulated notes by about 950 Hz (Fig. 5). It is important to take into consideration that this comparison was performed on recordings from three individuals of A. shihuemoy from a single location. Tadpoles. Ontogenetic variation of 11 characteristics measured for 16 tadpoles in stages 24–46 (Gosner 1960) are summarized in Table 3. The tadpole of A. shihuemoy sp. nov. belongs to the exotrophic ecomorphological guild, benthic type as defined by Altig & Johnson (1989). This description is based on a tadpole (MHNC lot 12987) at developmental Stage 25 (Gosner 1960; Figs. 3 A, C) and measuring a total length 17.2 mm; body length 6.2 mm; maximum width 4.1 mm; depth 3.0 mm; internarial distance 0.6 mm; eye to nares distance 0.5 mm; eye diameter 0.6 mm; interorbital distance 1.0 mm, oral disc width 1.2 mm; tail length 11.0 mm. The body is globular, compressed in lateral view, and ovoid in dorsal view. The snout is rounded in dorsal view (Figs. 6 A and C). The mouth is located anteroventrally and is surrounded by a small oral disc (Figs. 6 B and D). Papillae are laterally emarginated, simple and conical. Marginal papillae absent on anterior labium, present in one complete row on posterior labium. Anterior jaw sheath has a medial indentation with reduced serration, posterior jaw sheath Vshaped and has serration throughout. Lateral processes long, extending well past lower jaw. Labial Tooth Row Formula (LTRF) is 2(2)/3(1). A-1 complete, A-2 with medial gap, same width as A-1. P-1, P-2, and P-3 complete; P-1 and P-2 equal width, P-3 shorter. Nares are oval, small, without projections and inflexions, and are located dorsolaterally. Eyes are small and oriented dorsolaterally. The spiracle is single, sinistral, and located just before mid-body. Fins are concave and the posterior end is rounded, reaching their maximum height at the last third of the tail; the dorsal fin does not extend onto the body. The maximum tail height is reached at about the end length and is as high as body height. In preservative, the body is dark grey, the belly is translucent with intestines slightly visible; caudal musculature creamy white with small, irregular grey flecks and slightly translucent fins. In life, the body is dark brown with black spots, the belly is transparent but slightly pigmented posteriorly, intestines well visible through skin, caudal musculature poorly pigmented and transparent tail fin with melanophores in small and irregular clusters along the tail. Gosner stages In addition, we described observations in the field following Gosner (1960). We found two clutches of eggs in small rocky cavities; they were monitored every two days in order to take notes about the development of eggs. The first set was found on 25 June 2013, with 22 eggs at stage 13; a stage in which the neural plate develops on the dorsal surface, and eggs were covered by a transparent orange mucilage. After nine days, eggs become tadpoles, where they reach stage 17 (tail bud). The same embryos were abandoned by the father after two days on 0 5 July 2013. The embryos were getting dirty and dry, and inside we could observe the larvae of an insect, which could likely prey upon the embryos. The second set was found on 10 July 2013, containing 25 eggs in stage 13 covered by a transparent orange mucilage. After ten days, eggs were observed at stage 19 (heart beat), which is represented mainly by the development of the gills and tail. Seven days later tadpoles were at stage 21 (cornea transparent), in which the tail becomes increasingly transparent. Finally, three days later (30 July 2013) only five tadpoles were found at stage 22 (tail fin circulation); we assumed that the father was carrying the rest of tadpoles. One year

Ameerega shihuemoy sp. nov. (Figures 1���3, 6) Cryptophyllobates sp: Chaparro & Ochoa 2005 p.7 (MHNC 4779 collected on 0 7 December 2004 by J. C. Chaparro & J. A. Ochoa at Erika Lodge, Departamento Madre de Dios). Ameerega gr. pictus: Chaparro et al. 2016 p. 2 (from Amarakaeri Communal Reserve, Departamento Madre de Dios). Ameerega sp1: Whitworth & Villacampa 2014 p. 3 (from Manu Learning Centre, Departamento Madre de Dios). Holotype. MHNC 15488 (Fig. 1), an adult female, from near Cupudnoe river, 440 m elevation, Peru (Coordinates: 12��47'26.70" S, 70��58'21.30" W), Distrito Madre de Dios, Provincia Manu, Departamento Madre de Dios, collected on 25 April 2010 by R. C. Guti��rrez. Paratopotypes. Seven specimens: Five adult females (MUSA 3178, MUSA 3180���3182, MHNC 12988), and two immature females (MUSA 3177, MUSA 3 179), collected with the holotype by R. C. Guti��rrez. Paratypes. Fifteen specimens, all from Provincia Manu, Departamento Madre de Dios, Peru. Two adult females (MHNC 10406���10407), and one adult male (MHNC 10408), from the Reserva Comunal Amarakaeri, 32 km east of Huepetuhe, near the headwaters of Colorado river, 846 m elevation, (Coordinates: 12��59'59.25'' S, 70��50'31.32'' W), collected on 0 7 November 2010 by J. Delgado and J. G. Estrada. Two adult females (MHNC 10525���10526), and two adult males (MHNC 10536, MHNC 10542), from the Reserva Comunal Amarakaeri, 21.5 km southwest of Shintuya, near Azul river, 480 m elevation, (Coordinates: 12��48'50.8'' S, 71��06'07.8'' W), collected on 23 May 2011 by J. Delgado. An adult male (MHNC 15863), from the Reserva Comunal Amarakaeri, Pad A, Lote 76, near Dahuene river, 530 m elevation, (Coordinates: 12��58'38.47'' S, 71��01'30.73'' W), collected on 20 July 2016 by R. Coronel and G. Valencia. Two immature specimens (MHNC 10806, MHNC 10811), from 3.5 km south of Shintuya, near Serjali river, 450 m elevation, (Coordinates: 12��42'49.02'' S, 71��15'17.51'' W), collected on 20 January 2011 by J. G. Estrada. Two immature specimens (MHNC 10816, MHNC 10826), from 10 km southwest of Itahuania, near Shilive river, 450 m elevation, (Coordinates: 12��43'02.93'' S, 71��09'07.56'' W), collected on 26 January 2011 by J. G. Estrada. An immature female (MHNC 14561; Figs. 2A, B), from the Reserva Comunal Amarakaeri, Pad A, Lote 76, near Colorado river, 825 m elevation, (Coordinates: 12��59'17.97'' S, 71��0'52.67'' W), collected on 0 4 February 2015 by T. Gregory. An adult specimen (MHNC 5012), from Aguas Calientes, 2.75 km east (downstream) from Shintuya, (Coordinates: 12��40'7.27" S, 71��16'12.20'' W), collected on 0 9 July 2014 by S. J. Serrano. An immature specimen (MHNC 4779; Figs. 2 C, D), from Erika Lodge, 500 m elevation, (Coordinates: 12��45'23.53" S, 71��22'48.68" W), collected on 0 7 December 2004 by J. C. Chaparro and J. A. Ochoa. Referred specimens. Five specimens: MUSM 31611, MUSM 31664, MUSM 31691, MUSM 31692 and MUSM 31730, from Manu Learning Centre Research Station, 460 m elevation, Peru, (Coordinates: 12��47'21.849" S, 71��23'28.06" W), Distrito Manu, Provincia Manu, Departamento Madre de Dios, Peru, collected on 12 February, 0 5 July, 21July, and 25 July 2012, and on 0 3 February 2013, respectively, by A. Whitworth. Etymology. The specific name shihuemoy (English pronunciation: shee-way-moy) corresponds to the Harakmbut word for "poison dart frog". The Amarakaeri are aboriginals from Amazonian Peru and their language belongs to the Harakmbut linguistic group. They coexist with the new species. Diagnosis and comparisons with other species. Following Grant et al. (2006), the new species is assigned to the genus Ameerega on the basis of the following characters: dorsal skin finely granular; Finger I> Finger II when adpressed; toe and finger webbing absent; presence of bright flash marks and absence of ventrolateral line. This is a medium species of Ameerega with an adult SVL of 23.3 �� 2.7 (19.3���30.4 mm, N = 25). Vocal slits present; snout subacuminated, short and protruding. The head and dorsal surfaces are blackish with bronze-brown shades. The flanks are black with metallic bluish-green blotches, with a distinct bright orange or coral stripe extending from groin to above the eye and forward along the canthus rostralis to join around the snout. A flash spot is observed on the hip and upper surface of thighs, and a pale orange spot is located on the anterodorsal base of the thighs. It lacks concealed axillary and calf spots. The ventral surfaces have black reticulations with blue. The maxillary arch has teeth and the dentigerous process of vomers is lacking. The choanae are partly concealed by the palatal shelf of maxilla. The finger discs are weakly to moderately expanded, and when adpressed the first finger is slightly longer than second, and the fourth is shorter than first and second fingers, while the third is longer than all others. The finger discs are weakly to moderately expanded. The finger disc of the third finger of adults is about 1.2 times wider than the immediately adjacent part of the finger. The third finger shows no sexual dimorphism. Basal webbing is present on toes II���IV. Morphologically, divergence of A. shihuemoy from other species is supported by a unique combination of characters: the lack of the conspicuous axillary, thigh and calf flash marks, characteristics of many other species of Ameerega; black granular dorsum with cream to light orange dorsolateral lines, metallic blue venter and underside of extremities with black reticulations and spots. Within Ameerega, it shows the general appearance of A. altamazonica Twomey & Brown 2008, A. boliviana (Boulenger 1902), A. hahneli (Boulenger 1883), A. ignipedis Brown & Twomey 2009, A. petersi (Silverstone 1976), A. picta (Bibron 1838), A. pongoensis (Schulte 1999), A. pulchripecta (Silverstone 1976), A. simulans (Myers, Rodriguez & Icochea 1998), A. smaragdina (Silverstone 1976), and A. yungicola (L��tters, Schmitz & Reichle 2005), with which it shares a granular black to brown dorsum, a light labial bar, a conspicuous dorsolateral line running from the snout to the groin, and a metallic blue belly and underside of arms and hind limbs. From all these species it can be distinguished by lacking flash marks on the axillae, thighs, and calves (absent only in A. boliviana, A. simulans, A. smaragdina, most A. petersi, and some A. pongoensis), by having bright cream to orange dorsolateral stripes (white, pale yellowish-green, intense yellow, or green in all other species but A. picta), and by its blue belly reticulated with black (bluish white and black in A. boliviana, green and blue with black marbling in A. petersi, and green and blue lacking black marbling in A. smaragdina). Measurements (in mm) of the holotype. The female holotype (Fig. 1) has SVL, 25.7; TL, 12.3; GBW, 8.2; HW, 7.9; HL, 8.0; ED, 5.2; IOD, 3.0; TSCN, 2.0; NED, 2.6; IND, 2.9; EL, 3.3; HDT, 2.0; MTD, 0.8; HaL, 7.0; WTFD, 0.7; WTF, 0.5; WTTD, 0.8; WTT, 0.4; WFTD, 0.9; WFT, 0.4. Description of the type series. External morphology: A small to medium-size Ameerega, with adult males attaining approximately 19.2���21.8 mm SVL and adult females approximately 21.5���25.7 mm SVL (measurements and proportions summarized in Table 1). Dorsal skin of head, body, shank, thigh, and hind limbs coarsely and conspicuously granular; skin smooth or nearly smooth on forelimbs, and smooth on sides of head, body and on ventral surfaces. Head slightly wider between jaw articulations than between outer edges of upper eyelids; head usually narrower than body or about as wide. Head width between jaws 29���33% of SVL. Snout sloping, bluntly pointed or rounded in profile, truncate to rounded (usually) or bluntly pointed in dorsal and ventral view. Nares situated near tip of snout, directed slightly posterolaterally; nares visible from front and from below but not from above. Canthus rostralis short, protruding; loreal region nearly vertical, slightly concave to flat. Interorbital distance wider than upper eyelid. Eye shorter than snout length; distance from center of naris to eye 60���87% of eye length. Tympanum 35���65% of eye size. Hand (Fig. 1 F) moderately large, its length 23���28% of SVL and 74���90% of head width between angles of jaws. Relative lengths of adpressed fingers IV> I> II> III; fingers I, II, and IV approximately similar in size when adpressed, with overlapping discs; adpressed first finger varies from slightly shorter to slightly longer than second. Finger discs weakly to moderately expanded; third finger disc 1.2���1.8 times wider than distal end of adjacent phalanx. A large outer metacarpal (= palmar) tubercle on the base of the palm and a smaller inner metacarpal tubercle on the base of first finger, these being relatively small, with rounded surfaces. One subarticular tubercle on fingers I & II, and two subarticular tubercles on fingers III & IV; subarticular tubercles well developed and prominently raised, although distal one on finger III and both in IV sometimes weaker; supernumerary tubercles, finger keels, and outer metacarpal fold absent; tibia length 47���54% of SVL; relative lengths of adpressed toes, IV> III> V> II> I; first toe usually reaching base of subarticular tubercle of second toe; basal webbing on toes II���IV; toe fringes absent; outer metatarsal fold absent; a large inner metatarsal fold located on the distal half of tarsus and extended near the inner metatarsal tubercle (Fig. 1 G); ventrolateral side of tarsus relatively smooth, not especially rugose or tubercular; toes with moderately expanded discs, wider than finger discs (Fig. 1 G); one to three moderately raised subarticular tubercles (one each on toes I & II, two each on III & V, and three on IV); two large metatarsal tubercles with low, rounded surfaces; inner metatarsal tubercle slightly larger than outer metatarsal tubercle; supernumerary tubercles absent. Color pattern in life. (Based on living specimens and photographs; Figs. 2, 3). The noticeably granular body is blackish with bronze shades middorsally, turning black and blue dorsolaterally. The flanks are mostly black with metallic bluish green, with a distinct bright orange, coral, or cream stripe extending obliquely from groin to above the eye, and forward along the canthus rostralis to join around the snout. A pale whitish-bronze labial stripe commences between naris and eye, and extends posteriorly under the eye and tympanum to the base of the upper arm. A bright orange to coral spot is present on the hip and upper thigh; there is no calf spot and no pale markings in axilla or groin. The limbs are bronze-brown with shades of green and black. The ventral surfaces are overall blue with variable black reticulation, while in some individuals the ventral surfaces are overall black with blue reticulation. Color in preservative (Fig. 1). The bright orange and coral colors fade to pale grey; the blue ventral surfaces fade to grey in a dark reticulum of variable distinctiveness. Vocalization. Recordings of three A. shihuemoy males were taken at the Manu Learning Centre reserve. The advertisement call of this new species can be characterized as a series of chirp-like, pulsed notes. Notes are repeated at a rate of 0.8���1.0 notes per second (0.9 �� 0.1), duration of individual notes range from 84���109 ms (98.4 �� 6.8 ms), spaced 969���1196 ms apart (1042.5 �� 186.6 ms), with eight pulses (8.2 �� 0.8) per note. Dominant frequency ranges from 4478.9���4909.6 Hz (4672.72 �� 251.0 Hz) and is not frequency-modulated. Calling activity happens most frequently in the early mornings between 05:00 to 09:00 and late afternoon between 16:00 to 18:00. We also recorded a second call in A. shihuemoy consisting of three notes in quick succession (within 174.9 �� 16.6 ms of each other), repeated once every four to five seconds. The three notes in this call have different duration; the first note (87.2 �� 2.5 ms) typically has a longer duration than the second (69.4 �� 2.6 ms) and the third note (72.8 �� 2.8 ms). This three-note call seems to function as an aggressive or territorial call. We compared the call of the following species with Ameerega shihuemoy: two recordings of A. simulans (74 advertisement calls) by J. M. Padial from Marcapata, Quispicanchis Province, Department Cusco, Peru, were obtained from Fonoteca Zoologica (Museo Nacional de Ciencias Naturales 2016); two recordings of A. boliviana (45 advertisement calls) by J. Bosch & I. De la Riva from Correo-Apolo, La Paz, Bolivia; one recording of A. yungicola (30 advertisement calls) by M. Pacheco-Suarez from Caranavi, Yungas, Bolivia obtained from ���Gu��a fotogr��fica de los anfibios de la region de las yungas Bolivia ��� (Pacheco-Suarez 2015); two recordings of A. hahneli (46 advertisement calls) by J. Serrano-Rojas from Shintuya, Madre de Dios, Peru; two recordings of A. picta (63 advertisement calls) by J. M. Padial from Madidi National Park, Northern Bolivia and from Paractito, Cochabamba Department, Central West Bolivia deposited at Fonoteca Zoologica (Museo Nacional de Ciencias Naturales 2016); two recordings of A. macero (55 advertisement calls) by J. Serrano-Rojas from Manu Learning Centre, Madre de Dios, Peru. The advertisement call of A. shihuemoy is easily distinguished (Fig. 4) from the morphologically similar Ameerega species of the Ameerega picta group (for measurable call parameters and intra- and interspecific variation see Table 2). Parameters A. shihuemoy A. simulans A. picta A. hahneli Mean SD Mean SD Mean SD Mean SD Calls/N�� of individuals 80/3 - 74/2 - 63/2 - 46/2 - Notes/min 52.8 5.4 79.2 3.6 130.2 6.0 511.8 4.2 Notes/s 0.9 0.1 1.3 0.1 2.2 0.1 8.5 0.1 Duration of the note (ms) 98.4 6.8 104.9 10.2 45.8 2.4 13.4 2.8 Break between calls (ms) 1042.5 186.6 691.4 162.7 429.9 44.6 107.4 12.6 Fundamental frequency (Hz) 4237.0 281.9 4060.9 74.6 3770.7 76.7 2516.8 83.7 Dominant frequency (Hz) 4672.7 251.0 4460.3 157.7 4044.2 94.7 4550.0 49.1 continued. Parameters A. boliviana A. yungicola A. macero Mean SD Mean SD Mean SD Calls/N�� of individuals 45/ 2 - 30 - 55 /2 - Notes/min 72.0 4.2 313.2 0.0 519.4 2.2 Notes/s 1.2 0.1 5.2 0.0 8.7 0.0 Duration of the note (ms) 80.9 8.6 47.8 4.7 37.6 1.3 Break between calls (ms) 783.2 88.6 148.1 7.3 76.1 2.7 Fundamental frequency (Hz) 3416.1 68.2 3475.7 43.5 3353.7 38.1 Dominant frequency (Hz) 3846.0 46.3 3703.7 0.0 3617.6 0.0 The note duration of the new species (84���109 ms) is longer than A. hahneli (11���18 ms), A. macero (36���40 ms), A. picta (41���52 ms) and A. yungicola (40���57 ms), but similar to A. boliviana (75���94 ms) and A. simulans (91���118 ms). The space between calls in A. shihuemoy is longer than in the other six species. The number of notes per second in the advertisement call of A. shihuemoy is lower than the other six species. The call of the new species has a higher fundamental frequency than all other species but similar to A. simulans, and a higher dominant frequency than all other species but similar to A. hahneli and A. simulans, except the latter species has pronounced upward frequency-modulated notes by about 950 Hz (Fig. 5). It is important to take into consideration that this comparison was performed on recordings from three individuals of A. shihuemoy from a single location. Tadpoles. Ontogenetic variation of 11 characteristics measured for 16 tadpoles in stages 24���46 (Gosner 1960) are summarized in Table 3. The tadpole of A. shihuemoy sp. nov. belongs to the exotrophic ecomorphological guild, benthic type as defined by Altig & Johnson (1989). This description is based on a tadpole (MHNC lot 12987) at developmental Stage 25 (Gosner 1960; Figs. 3 A, C) and measuring a total length 17.2 mm; body length 6.2 mm; maximum width 4.1 mm; depth 3.0 mm; internarial distance 0.6 mm; eye to nares distance 0.5 mm; eye diameter 0.6 mm; interorbital distance 1.0 mm, oral disc width 1.2 mm; tail length 11.0 mm. The body is globular, compressed in lateral view, and ovoid in dorsal view. The snout is rounded in dorsal view (Figs. 6 A and C). The mouth is located anteroventrally and is surrounded by a small oral disc (Figs. 6 B and D). Papillae are laterally emarginated, simple and conical. Marginal papillae absent on anterior labium, present in one complete row on posterior labium. Anterior jaw sheath has a medial indentation with reduced serration, posterior jaw sheath Vshaped and has serration throughout. Lateral processes long, extending well past lower jaw. Labial Tooth Row Formula (LTRF) is 2(2)/3(1). A-1 complete, A-2 with medial gap, same width as A-1. P-1, P-2, and P-3 complete; P-1 and P-2 equal width, P-3 shorter. Nares are oval, small, without projections and inflexions, and are located dorsolaterally. Eyes are small and oriented dorsolaterally. The spiracle is single, sinistral, and located just before mid-body. Fins are concave and the posterior end is rounded, reaching their maximum height at the last third of the tail; the dorsal fin does not extend onto the body. The maximum tail height is reached at about the end length and is as high as body height. In preservative, the body is dark grey, the belly is translucent with intestines slightly visible; caudal musculature creamy white with small, irregular grey flecks and slightly translucent fins. In life, the body is dark brown with black spots, the belly is transparent but slightly pigmented posteriorly, intestines well visible through skin, caudal musculature poorly pigmented and transparent tail fin with melanophores in small and irregular clusters along the tail. Gosner stages In addition, we described observations in the field following Gosner (1960). We found two clutches of eggs in small rocky cavities; they were monitored every two days in order to take notes about the development of eggs. The first set was found on 25 June 2013, with 22 eggs at stage 13; a stage in which the neural plate develops on the dorsal surface, and eggs were covered by a transparent orange mucilage. After nine days, eggs become tadpoles, where they reach stage 17 (tail bud). The same embryos were abandoned by the father after two days on 0 5 July 2013. The embryos were getting dirty and dry, and inside we could observe the larvae of an insect, which could likely prey upon the embryos. The second set was found on 10 July 2013, containing 25 eggs in stage 13 covered by a transparent orange mucilage. After ten days, eggs were observed at stage 19 (heart beat), which is represented mainly by the development of the gills and tail. Seven days later tadpoles were at stage 21 (cornea transparent), in which the tail becomes increasingly transparent. Finally, three days later (30 July 2013) only five tadpoles were found at stage 22 (tail fin circulation); we assumed that the father was carrying the rest of tadpoles. 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Elachistocleis muiraquitan was recently described from fifteen specimens found at two sites in Acre state, northwestern Brazil. Prior to the description of E. muiraquitan, individuals fitting the description of this species found in southeastern Peru and northwestern Bolivia were identified as Elachistocleis bicolor, a species associated with markedly different habitat and environmental conditions. Here, we re-identified these specimens and also propose the first map of E. muiraquitan's potential distribution, based on known localities along with climatic and environmental parameters.