Dendropsophus shiwiarum Ortega-Andrade & Ron, 2013, sp. nov.
Creators: name: Ortega-Andrade, H. Mauricio
name: Ron, Santiago R.
Publisher: name: Zenodo
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Hylidae
Dendropsophus
Dendropsophus shiwiarum

description: Dendropsophus shiwiarum sp. nov.
(Figs. 1–4, Table 1)
“ Hyla riveroi ”.—Duellman (1978); Almendáriz (1989); Duellman & Mendelson (1995). “ Dendropsophus sp.”.—Ortega-Andrade (2010).
Holotype: DHMECN 4393, an adult female collected at Kurintza, 2.060° S, 76.780° W, 354–m elevation, Pastaza province, Republic of Ecuador, between 26–28 April 2007 by H. Mauricio Ortega-Andrade, Miguel Cachay, Jorge Santi and Tseremp Timias.
Paratopotypes: Two adult males (DHMECN 4394, 4399), with same collection data as holotype.
Paratypes: ECUADOR: Nineteen adult males (DHMECN 4430–32, QCAZ 8250, 8252, 8253, 8308, 21936– 937, 22885, 27045–046, 28086), two juvenile females (QCAZ 11544, 27791) and one adult female (QCAZ 11542). Napo province: QCAZ 27045, 27046 were collected at Ahuano, Runa Huasi, 1.060° S, 77.550° W, 394 m a.s.l, on 25 May 2005 by E. Carrillo and M. Read. Orellana province: QCAZ 8250, 8252, 8253, 8308 were collected at 42 km carretera Pompeya–Iro, Parque Nacional Yasuní, 0.650° S, 76.450° W, 236 m a.s.l; QCAZ 11542, 27791, collected at Apaika, Parque Nacional Yasuní, 0.870° S, 75.860° W, 209 m a.s.l.; QCAZ 22752 collected at Estación Científica Yasuni de la Pontificia Universidad Católica del Ecuador, Parque Nacional Yasuní, 0.680° S, 76.410° W, 241–m a.s.l. Pastaza province: DHMECN 4430–32, were collected at Juyuintza, 2.110° S, 76.190° W, 200 m a.s.l, between 26–28 April 2007 by H. M. Ortega-Andrade, M. Cachay, J. Santi and T. Timias. Sucumbíos province: QCAZ 21936, 21937, 22885 were collected at 2.5 km N de Lago Agrio, 0.120° N, 76.880° W, 302 m a.s.l, on 17 March 2003 by S. Valdiviezo, D. Tirira, J. Wiens and L. Coloma; QCAZ 11544, collected at Hostería La Selva, 0.450° S, 76.280° W, 232 m a.s.l; QCAZ 28086, collected at Reserva de Producción Faunística Cuyabeno, 0.320° S, 75.770° W, 223 m a.s.l; QCAZ 52969, 52970 collected at Limoncocha, 0.40688° S, 76.62063° S, 256 m a.s.l., on 13 March 2012 by H. Mauricio Ortega-Andrade.
Diagnosis. A small species of Dendropsophus distinguished by the following combination of traits: (1) females similar in size (up to 18.8 mm SVL, N = 2) to males (up to 18.5 mm, N = 19); (2) snout short, truncate in both dorsal and lateral profiles; (3) canthus rostralis rounded in cross-section, loreal region slightly concave; (4) tympanic membrane non-differentiated and annulus evident only ventrally, supratympanic fold weakly developed; (5) vomerine odontophores small, oblique, separated medially between choanae, bearing 1–5 teeth; (6) skin on dorsum, head and dorsal surfaces of limbs finely shagreen; skin on belly and proximal surfaces of thighs coarsely granular, skin on other surfaces smooth; (7) ulnar fold and tubercles absent, tarsal fold and tubercles on outer edge of tarsus absent, inner metatarsal tubercle distinct and oval, outer metatarsal tubercle absent; (8) axillary membrane developed, reaching a half or three-fourths of arm length; (9) fingers about onethird webbed, webbing formulae I2 ½–2 II2 -– 3III 2–2+IV, toes about four-fifths webbed, webbing formulae I1 ½– 2 II1 +–2 III1 +–2- IV2 +–1+V; (10) disc of Finger III and Toe IV with pointed tip, conical tubercle on dorsal surface of each disc which represents the projection of distal phalanx, distal subarticular tubercle of fourth finger bifid in all specimens; (11) in life, dorsum coppery tan, with or without dark reddish brown spots or an inverted triangular blotch on scapular area; (12) belly immaculate white, ventral and hidden surfaces of thighs translucent fleshy white, vocal sac bright yellow; (14) iris coloration varies from silvery copper to reddish pale yellow with a copper ring around pupil; (15) bones white.
Comparisons with other species. Among other small species of Dendropsophus in the Amazon Basin, the new species is most similar to Dendropsophus riveroi. Both species have similar body size, but the latter have rounded discs on tips of Finger III and Toe IV. The type material of Dendropsophus riveroi and specimens from the type locality (e.g., ICN 50611, 50613, 50615, 50617) can be distinguished from D. shiwiarum (whose traits are given in parenthesis, Figs. 2, 4) by having: (1) smooth skin on the dorsum (finely shagreen); (2) tympanic membrane and tympanic annulus prominent (tympanic membrane non-differentiated and annulus evident only ventrally); (3) snout rounded in both dorsal and lateral views and in profile (both truncated); and (4) dorsal surfaces of digital discs lacking a conical tubercle as result of projection of distal phalanx (tubercle present).
Amazonian species of the Dendropsophus microcephalus group (Duellman & Fouquette 1968) [D. coffeus (Köhler, Jungfer & Reichle, 2005), D. joannae (Köhler & Lötters, 2001a, 2001b), D. leali (Bokermann, 1964), D. walfordi (Bokermann, 1962)] are distinguished by their larger body size, iris and dorsal coloration and rounded discs. The Cerrado dweller D. cruzi (Pombal & Bastos, 1998) differs in having elliptic discs on digits. The holotype and allotype of Dendropsophus leali (MZUSP 74210, 74211) differ from the new species by its larger size (males up to 22 mm in D. leali; 18.85 mm in D. shiwiarum), prominent tympanic membrane and annulus, rounded tips of digits, and commonly with “X” or “)(“ –shaped dark marks in the scapular region (holotype), or uniform reddish tan dorsum with a narrow, barely visible, dark brown interorbital mark (allotype). Dendropsophus coffeus, an inhabitant of southwest Amazonian ecoregion, Departamento La Paz, Bolivia, is similar to D. shiwiarum in snout shape, dorsal color pattern and the presence of a dark brown lateral stripe, but is distinguished by its larger size (SVL up to 21.2 mm in males, 26.0 mm in females), prominent tympanic membrane and annulus, smooth dorsal skin with minute scattered tubercles. Dendropsophus joannae has protuberant eyes, tuberculate dorsal skin, a red inner iris in life, whereas D. juliani Moravec, Aparicio & Köhler (2006) differs by exhibiting a greenish ventral surfaces and vocal sac in life; as well both species present differences in advertisement call (Köhler & Lötters 2001a; Moravec et al. 2006).
Dendropsophus aperomeus (Duellman 1982), is an inhabitant of Amazonian slopes of Andes in northern and central Peru, distinguished from D. shiwiarum by its larger size (males up to 21.3 mm of SVL, females up to 25.0 mm), having a single distal subarticular tubercle on the fourth finger, a white supra-anal stripe, and rounded tips of digits. Dendropsophus minimus (Ahl, 1933) is only known from the type locality at Rio Amazonas region in central Brazil, and is distinguished from the new species by having a concealed tympanum, rounded tips of digits and pale dorsolateral stripes which extend to the groin (dorsolateral stripes in D. shiwiarum only extend to the mid-flank). Dendropsophus miyatai (Vigle & Goderdhan-Vigle 1990) is an inhabitant of upper Amazon Basin of Ecuador and northeastern Peru, distinguished by having rounded tip of digits, concealed tympanum, and dorsal color pattern with bright red and yellow blotches.
Members of the Dendropsophus minutus group [Dendropsophus delarivai (Köhler & Lötters 2001b), D. minutus (Peters, 1872), D. xapuriensis (Martins & Cardoso, 1987)] differ from D. shiwiarum in having a white supra-anal stripe and a longer cloacal sheaths.
Description of holotype. Head about as wide as body, wide as long, widest below eyes; snout short (30% of HL), truncate in both dorsal and lateral profiles (Fig. 2); distance from nostril to corner of eye shorter than diameter of eye (80% of ED); canthus rostralis rounded and distinct; loreal region slightly concave; lips not flared; internarial area not depressed, nostrils barely protuberant, directed anterolaterally, at about three-quarters the distance from eyes to tip of snout; interorbital area flat, in about 36% of head width; eye large, protuberant, its diameter about two times the distance from upper lip to orbit and about 30% of head length. Tympanic membrane is concealed under skin; thin supratympanic fold extends laterally from posterior corner of eye to point above arm insertion; fold hides upper edge of tympanic annulus. Ventral border of the tympanic annulus barely evident, round, located behind mid-orbit, its diameter about two-thirds of eye length, separated from eye by a distance of about half its diameter. Arm not hypertrophied; dorsal surfaces of arms shagreen; axillary membrane extends to mid-length of upper arm; ulnar folds and tubercles absent; fingers short and broad, relative length of fingers I <II <IV <III; large discs, diameter of disc on third finger about as wide of tympanic membrane; disc pointed at tip on Finger III, and nearly round on other fingers (Figs. 3 – 4); conical tubercle on dorsal surface of each disc; subarticular tubercles small, round; distal tubercle of fourth finger prominent, bifid; supernumerary tubercles small, round, and flat; barely evident supernumerary tubercles form a row along the proximal one third of each finger; palmar tubercle small, ovoid; prepolical tubercle large, flat, ovoid; webbing of hand I2 ½–2 II2 -– 3III 2–2+IV. Legs slender and long (FL+TL+FoL= 1.7 times the SVL); tarsal fold and tarsal tubercles absent. Toes moderately large; discs about the same size than those of fingers; disc pointed on Toe IV, nearly round in other toes (Figs. 3 – 4); conical tubercle present in dorsal surface of each disc; relative length of toes I <II <III = V <IV; outer metatarsal tubercle absent; inner metatarsal tubercle moderately small, elliptical; subarticular tubercles small, round; supernumerary tubercles absent; webbing on foot I1 ½–2 II1 +–2 III1 +–2- IV2 +–1+V. Skin of dorsum, head and dorsal surfaces of limbs finely shagreen to tuberculate; skin on flanks shagreen; skin on venter areolate; skin on chest and throat smooth. Cloacal opening directed posteriorly at upper level of thighs; cloacal sheath very short, covering 1/6 upper edge of cloacal opening; cloacal folds and tubercles absent. Tongue cordiform, posterior fourth not attached to mouth floor, vomerine odontophores small, but prominent, about half the diameter of choanae, each bearing five vomerine teeth; choanae moderately sized, ovoid, medially separated, not covered by palatal shelf.
Color of holotype in life. Dorsal surfaces pinkish tan, paler toward posterior third of body; blurred brown transversal stripe on sacrum; dark brown interorbital, canthal, and post-orbital stripes extending to mid-flank; white bar bordered with brown below each eye. Three pale brown narrow stripes on dorsal surfaces of shanks; dorsal surfaces of arms uniform tan. Outer edge of venter, chest, throat, webbing, hidden surfaces, ventral surfaces of forearms and limbs flesh-colored and unpigmented; belly immaculate, whitish-cream. Iris silvery copper with a dark red ring around pupil.
Measurements of holotype (in mm). Specimen DHMECN 4393 is a gravid female. SVL = 18.8; HW = 6.4; HL = 6.8; ED = 2.1; EN = 1.7; TD = 1.4; FL = 9.5; TL = 9.7; FoL = 13.5. Proportions: HL/SVL = 0.36; HW/HL = 0.94; FL/SVL = 0.51; ED/HL = 0.31; EN/HL = 0.25; EN/ED = 0.81; TL/SVL = 0.52; FoL/SVL = 0.72; TD/ED = 0.67.
Variation. Dendropsophus shiwiarum exhibits considerable variation in coloration and pattern, even among individuals of the same population (Fig. 5). In preservative, the dorsum of most specimens is reddish with irregular small marks (e.g. DHMECN 4394, 4430) or pinkish tan with blotches or large marks (DHMECN 4393); some specimens also have a reddish inverted triangular mark on the scapular region (QCAZ 21936, DHMECN 4432) or a wide interorbital bar and an inverted triangular reddish mark on the sacral region (QCAZ 11542). Most specimens also have a narrow dark brown interorbital bar, which can be complete (e.g. DHMECN 4394), or barely visible (DHMECN 5020), wider than the horizontal canthal bar (QCAZ 11542), or incomplete (DHMECN 4393). At night, males are pale reddish brown with a dark brown dorsolateral stripe and a yellow vocal sac (Fig. 5 E–F); by day, the dorsum is creamy tan with or without reddish brown markings. Iris coloration varies from silvery copper (e.g. DHMECN 4393) to reddish pale yellow (e.g. DHMECN 4432) with a copper ring around pupil. Stripes, unpigmented spots or bars below the eye are variable in number, even between sides in the same individual: 3–4 in about 30%, 2–3 in about 6%, 1–2 in about 7%, 1 in about 30%, lacking in about 7%. Males have long vocal slits, extending from the first quarter of the mandible to the angles of the jaws; vocal sac large, single, median, subgular and greatly distensible. The condition of pointed discs in D. shiwiarum is variable, from prominent as occur in most specimens (~90%, e.g. QCAZ 48105) to less evident in other specimens (e.g. QCAZ 51162). Anyway, the evidence suggests that it is not an artifact of preservation given live individuals present such pointed discs (see living specimens of D. shiwiarum in http://zoologia.puce.edu.ec/gallery/main.php?g2_itemId=3646&g2_page=1). Variation in measurements and proportions of specimens from the type series is summarized in Table 1.
Color in life. Background dorsal coloration varies from bright yellow, to yellowish tan, pinkish tan or coppery red (Figs. 1, 5). Irregular triangular marks or blotches or dark brown to pale red flecks are always present. Vocal sac in males and tips of digits are bright yellow.
Color in preservative. In alcohol, all bright yellow dorsal surfaces turn into cream; coppery red and pinkish tan areas become paler than in life; dark flecks in reddish brown individuals (e.g. DHMECN 4430) become barely visible. Ventral surfaces of forearms and limbs, outer edge of venter, chest, throat and webbing are transparent; belly immaculate cream.
Advertisement call (Fig. 6). A total of 35 calls were analyzed from two males recorded from Ecuador. Specimen DHMECN 4431 was recorded at 21:06 h on 17 May 2007 (air temperature ca. 22 °C) at Juyuintza, Pastaza province. Specimen QCAZ 48101 was recorded at 21:08 h on 3 April 2010 (air temperature ca. 24.2 °C), at km 93 on Pompeya – Iro road, Yasuni National Park, Orellana province. Both males were part of a small chorus (approximately six to eight calling males, not collected), calling from shrubs, 0.5 – 1.5 m above the water, on a swamp dominated by the palm Mauritia flexuosa. The call consists of one high pitched pulsed note, with length 52.83 ± 31.87 ms, dominant frequency range 3983.6– 5254.1 Hz (4714.5± 353.1 Hz), call rise time 23.1±7.8 ms, interval between notes 1946.15 ± 587.24 ms (range 814.0 – 2966.9) and a call rate ~32.9 calls/minute.
Natural history and distribution. Dendropsophus shiwiarum is known from 22 localities in eastern Ecuador, provinces of Napo, Orellana, Pastaza and Sucumbíos, up to 550 m a.s.l (Fig. 7). This species occurs in about 34,861 km 2 along the evergreen lowland rainforest, but it is expected to be more widely distributed along the upper Amazon basin. Duellman & Mendelson (1995) describe nine specimens of “ D. riveroi ” from the Amazon Basin of northern Loreto, Peru (ca. 108 km E from the type locality of D. shiwiarum and ca. 50 km E from Juyuintza) that, based on their morphological similarity, may correspond to D. shiwiarum.
Dendropsophus shiwiarum is an opportunistic breeder at temporary ponds and swamps on secondary forest, flooded and open areas, being very rare through dry seasons. Males were calling from leaves of bushes, trees and Heliconia at secondary forest, 0.05–1.5 m above the water, in the border of a temporary swamp at Kurintza on 26 and 28 April 2007, and from leaves of bushes and low vegetation at flooded areas dominated by the palm Mauritia flexuosa (vegetation type locally known as moretal) in Juyuintza, on 17 May 2007. The holotype is a gravid female which, after capture, deposited a clutch of 121 unpigmented eggs in a single layer of clear gelatin inside a plastic bag; it was captured in amplexus with DHMECN 4394 (16.8 mm in SVL) perched on a Heliconia leaf at 18:33 h at Kurintza. Two days after the collection of the holotype, the rain stopped, the swamp dried-out and D. shiwiarum was not found anymore.
Other hylid frogs found at these swamps were Dendropsophus brevifrons (Duellman & Crump 1974), D. parviceps (Boulenger 1882), D. rhodopeplus (Günther 1858), D. sarayacuensis (Shreve 1935), Hypsiboas calcaratus (Troschel 1848), H. cinerascens (Spix 1824), H. geographicus (Spix 1824), H. lanciformis (Cope 1869), Osteocephalus deridens (Jungfer, Ron, Seipp & Almendariz 2000), O. fuscifacies (Jungfer, Ron, Seipp & Almendariz 2000), O. taurinus (Steindachner 1862) and Scinax funereus (Cope 1874). The aquatic snake Helicops angulatus (Linnaeus 1758) was also recorded at one of the swamps. The new species is sympatric with Dendropsophus bifurcus (Andersson 1945), D. bokermanni (Goin 1959), D. aff. leali, D. marmoratus (Laurenti 1768) D. miyatai, D. parviceps, D. rhodopeplus, D. sarayacuensis, and D. triangulum (Günther 1868) along its distributional range in eastern Ecuador.
Etymology. The new species is named for the Shiwiar indigenous nation of the Pastaza Trench in the upper Amazon Basin of Ecuador, as recognition for sharing their invaluable knowledge about biodiversity and rich culture, maintaining through the generations a close relationship with nature. Moreover, they generously granted permission to HMO?A to sample frogs from their ancestral territories. The epithet is a genitive noun in apposition.
description: Published as part of Ortega-Andrade, H. Mauricio & Ron, Santiago R., 2013, A new species of small tree frog, genus Dendropsophus (Anura: Hylidae) from the eastern Amazon lowlands of Ecuador, pp. 163-178 in Zootaxa 3652 (1) on pages 164-173, DOI: 10.11646/zootaxa.3652.1.6, http://zenodo.org/record/222516