Systematics of the Dendropsophus leucophyllatus species group (Anura, Hylidae) from the Chocó region of Ecuador, with description of a new species Author Aguirre, P. Doménica https://orcid.org/0009-0006-9146-9374 Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17 - 01 - 2184, Quito, Ecuador & Department of Biological Science, Florida State University, Tallahassee, FL 32306, USA Author Apunte, Katherine 0000-0003-4300-5111 Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17 - 01 - 2184, Quito, Ecuador Author Ron, Santiago R. 0000-0001-6300-9350 Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17 - 01 - 2184, Quito, Ecuador text Evolutionary Systematics 2025 2025-02-21 9 1 7 31 journal article 10.3897/evolsyst.9.135431 044BF460-92DB-4715-A639-DF2CF34AA07C Dendropsophus cannatellai sp. nov. Dendropsophus ebraccatus : Ortega-Andrade et al. 2010 . Dendropsophus ebraccatus : Ron et al. 2011 . Dendropsophus ebraccatus : Simpson et al. 2013 . Dendropsophus ebraccatus : Caminer et al. 2017 . Dendropsophus ebraccatus (in part): Pirani et al. 2020 . Dendropsophus ebraccatus (in part): Orrico et al. 2021 . Dendropsophus ebraccatus : Pintanel et al. 2022 . Dendropsophus ebraccatus (in part): Touchon et al. 2024 . Holotype . • QCAZ 40847 (field no. SC-PUCE 19751), adult male from Ecuador , Esmeraldas Province , Cantón San Lorenzo , Playón San Francisco ( 1.0873 ° N , 78.6905 ° W ), 64 m , collected by Luis A. Coloma , Santiago R. Ron , and Mark Weiner on 06 August 2007 . Paratypes (130) : All specimens were collected in Ecuador . Pichincha Province : • QCAZ 3768 –3774, 4840–4843 Cantón Pedro Vicente Maldonado , 1 km E from Maldonado ( 0.0832 ° N , 79.0394 ° W ), 745 m , collected by Stephan Lötters and Luis A. Coloma on 13 March, 1993 ; • QCAZ 9457 , 9489, Cantón Pedro Vicente Maldonado , Río Pitzará ( 0.1192 ° N , 79.2344 ° W ), 280 m , collected by Giovanna Rodríguez on 01 January, 1996 and 01 August, 1995 ; • QCAZ 10434 , 27207–27209 Cantón Puerto Quito , Aldea Salamandra ( 0.1192 ° N , 79.2344 ° W ), 149 m , collected by David Cannatella on 08 April, 2004 ; • QCAZ 27623 –27624 Cantón Pedro Vicente Maldonado , 1 km E from Maldonado ( 0.0829 ° N , 79.0383 ° W ), 670 m , collected by John J. Wiens on 30 March, 1990 ; • QCAZ 78254 –78259, 78309, 78576 Cantón Pedro Vicente Maldonado , Río Silanche ( 0.1278 ° N , 79.1414 ° W ), 369 m , collected by Gabriela B. Bittencourt on 19 January, 2023 ; • QCAZ 78809 –78814 Cantón Pedro Vicente Maldonado , Río Silanche ( 0.1278 ° N , 79.1414 ° W ), 369 m , collected by Santiago R. Ron on 17 July, 2023 . Esmeraldas Province : Santo Domingo de Onzole ( 0.8003 ° N , 79.0709 ° W ), 110 m , collected by Néstor Acosta on 18 January, 1997 ; • QCAZ 10704 , 22893, Cantón Pedro Vicente Maldonado ( 0.0878 ° N , 79.0491 ° W ), 670 m , collected by Luis Coloma on 13 March, 1993 and Sebastián Valdiviezo on 31 March, 1990 . Esmeraldas Province : • QCAZ 15519 , 23156, 23160, 23371, Cantón San Lorenzo , 3 km west of Durango ( 1.0643 ° N , 78.6443 ° W ), 158 m , collected by Ítalo Tapia on 15 June, 2001 and David Cannatella on 03 March, 2003 ; • QCAZ 23162 Cantón San Lorenzo , Durango ( 1.0127 ° N , 78.8954 ° W ), 14 m , collected by David Cannatella on 04 March, 2003 ; • QCAZ 26098 , 26100 Cantón San Lorenzo , Durango – San Lorenzo ( 1.0793 ° N , 78.6695 ° W ), 74 m , collected by Néstor Acosta on 08 June, 2003 ; • QCAZ 29489 –29511, 30329–30330, 55529–55530 Cantón San Lorenzo , Durango ( 1.0427 ° N , 78.6244 ° W ), 85 m , collected by Ítalo Tapia on 02 February, 2005 , Elicio Tapia on 19 December, 2004 and Justin Touchon on 20 April, 2013 ; • QCAZ 29799 –29809, 30255 Cantón San Lorenzo , Durango – San Francisco ( 1.0872 ° N , 78.6905 ° W ), collected by Ítalo Tapia on 17 September, 2004 ; • QCAZ 32136 San Francisco , 1.3 km east ( 1.0888 ° N , 78.69563 ° W ), 85 m , collected by Diego Almeida on 26 May, 2006 ; • QCAZ 35709 –35710 La Mayronga ( 1.05 ° N , 79.27 ° W ), 150 m , collected on 27 November, 1998 ; • QCAZ 37741 , 37845 Cantón San Lorenzo , 21 km of Durango ( 1.0248 ° N , 78.6175 ° W ), 284 m , collected by Ítalo Tapia on 03 January, 2008 ; • QCAZ 37749 Cantón San Lorenzo , 21 km of Durango ( 1.0470 ° N , 78.6182 ° W ), 104 m , collected by Ítalo Tapia on 15 December, 2007 ; • QCAZ 40276 , 40279 Cantón San Lorenzo , 2 km east of Durango ( 1.0247 ° N , 78.6174 ° W ), 85 m , collected by Ítalo Tapia on 13 March, 2009 ; • QCAZ 51854 , 51863, 51877 Cantón San Lorenzo , Durango ( 1.0379 ° N , 78.6222 ° W ), 283 m , collected by Andrés Merino on 10 September, 2011 ; • QCAZ 55531 –540, 55545–55550, 55555–55558 Cantón San Lorenzo , 9 km west of Durango ( 1.2083 ° N , 78.7413 ° W ), 91 m , collected by Justin Touchon on 26 April, 2013 ; • QCAZ 55541 –55544, 55551–55554 Cantón San Lorenzo , 7 km west of Durango ( 1.2044 ° N , 78.8447 ° W ), 139 m , collected by Justin Touchon on 22 April, 2013 ; • QCAZ 66629 –66635 Cantón San Lorenzo , Durango ( 1.0874 ° N , 78.6221 ° W ), 265 m , collected by Diego Almeida on 10 February, 2017 . Common names. English: Cannatella’s treefrog. Spanish: Ranita de Cannatella. Definition (Figs 8 , 9 ). In this section, we describe coloration in life. We assign the new species to the genus Dendropsophus based on the phylogeny (Fig. 1 ). The new species is characterized by: (1) mean SVL 23.4 mm in males (range 21–26; n = 115); 30.7 mm in females (range 28–33; n = 14); (2) axillary membrane reaching arm halfway to elbow; (3) reduced webbing on fingers; (4) webbing on feet; (5) palmar tubercle single; (6) pectoral glands present; (7) dorsal background coloration brown with white to bright yellow dorsolateral bands, extending to the eyes and top of head; white or bright yellow well defined sacral mark, fused with dorsolateral bands in some individuals; (8) absence of a light labial stripe in contact with the orbit; (9) dorsal surfaces of the limbs brown, with white to bright yellow ovoid marks, with the forearm typically bearing one to two marks, and the shank one to three; (10) webbing and ventral surfaces vary from orange to yellow; (11) iris dull bronze to coppery bronze, and (12) advertisement call pulsed with a mean dominant frequency = 3274.7 Hz (range 3169.0–3498.7) and mean pulse rate of the primary note = 95 pulses / s (range 87–100). Figure 8. Life coloration of Dendropsophus cannatellai sp. nov. First and third rows, dorsolateral views; second and fourth, ventral views. From left to right, first and second rows: QCAZ 51877 (SVL = 22.26 mm, male); QCAZ 78809 (SVL = 33.61 mm, female); QCAZ 55558 (SVL = 21.96 mm, male); third and fourth rows: QCAZ 40847 (holotype; SVL = 22.32 mm, male); QCAZ 78812 (SVL = 25.86 mm, male); QCAZ 78813 (SVL = 24.94 mm, male). See type series for locality data. All specimens are shown at the same scale. Figure 9. Adult preserved specimens of Dendropsophus cannatellai sp. nov. First and third rows, dorsolateral views; second and fourth, ventral views. From left to right, first and second rows: QCAZ 9457 (SVL = 28.65 mm, female); QCAZ 23162 (22.83 mm, male); QCAZ 26098 (30.73 mm, female); third and fourth rows: QCAZ 35710 (24.74 mm, male); QCAZ 78254 (24.71 mm, male); QCAZ 78257 (31.94 mm, female). See type-series for locality data. All specimens are shown at the same scale. Diagnosis. In this section, coloration refers to live adult individuals, unless otherwise noted. Adult individuals of Dendropsophus cannatellai sp. nov. can be distinguished from other species of the genus Dendropsophus that do not belong to the D. leucophyllatus group by the presence of two pectoral glands. Within the D. leucophyllatus group, we focus our comparisons with species from Central America and the Chocó region , also the most closely related. Other species of the group have a cis-Andean distribution and, therefore, cannot be confused with the new species. Nevertheless, we include diagnosis for species from the western Amazon basin, which are geographically closest to the new species range. Figure 10. Tadpole ( QCAZ 78576) of Dendropsophus cannatellai sp. nov , stage 25 ( Gosner 1960 ). Dorsal, ventral, and lateral views. Body length = 8.14 mm; tail length = 16.24 mm; total length = 24.38 mm. Dendropsophus ebraccatus differs from the new species (characters of the later in parenthesis) in having a light labial stripe reaching the orbit (area below the eye has dark background color, sometimes with one or two small light spots). In addition, D. ebraccatus either lacks dorsolateral yellow bands or, if present, they are wide and sinuous (if present, yellow dorsolateral bands straight to moderately curved; variation in coloration of D. ebraccatus from Duellman 1970 and Ohmer et al. 2009 ). Dendropsophus cannatellai sp. nov. clearly differs from its sister species, D. gryllatus , in dorsal coloration and by being larger (although SVL ranges overlap: male average SVL = 21.95, range 18.71–24.69 mm vs. SVL = 23.43, range 21.05–26.58 mm in D. cannatellai sp. nov. ) The dorsal pattern of D. gryllatus consists of two longitudinal dark brown lines against a pale-yellow background (Fig. 14 ; brown background with clear yellow dorsolateral bands in the new species). The new species also differs from D. gryllatus by having bright orange flash coloration (pale yellow in D. gryllatus ). Dendropsophus gryllatus exhibits a sacral spot that extends forward, making it more elongated and continuous with other dorsal markings (the sacral spot of D. ebraccatus is distinct, circular or slightly oval, and symmetrically positioned near the lower back, with well-defined edges that contrast against its darker dorsal pattern; sacral spot is absent in some individuals). The Amazonian species D. sarayacuensis and D. bifurcus also has a brown background color with yellow bands and marks. However, D. sarayacuensis differs in that the yellow areas in the shanks have irregular borders (borders are more linear in D. cannatellai sp. nov. ) Dendropsophus bifurcus differs by having less extensive yellow bands and marks, especially in the shanks where, in most individuals, they are absent, except for the heel (extensive yellow marks along the shank in D. cannatellai ). Dendropsophus reticulatus differs by having a uniform clear dorsal color pattern, sometimes with one or several dark brown round marks ( Caminer et al. 2017 ) (clear dorsal pattern restricted to head, dorsolateral bands, and rump in D. cannatellai ). Dendropsophus triangulum and D. arndti differ by being larger, SVL> 28.0 mm in males and> 33.0 mm in females ( Caminer et al. 2017 ) ( D. cannatellai males SVL range 21.05–26.58 mm , females = 28.25–31.94 mm ). Dendropsophus cannatellai sp. nov. can be distinguished from Dendropsophus carnifex ( D. columbianus group), a species from the Andean Chocó , by having a short axillary membrane that reaches halfway to the elbow (axillary membrane absent in D. carnifex ). Description of holotype (Figs 12 , 13 ). Adult male, SVL 22.32 mm , foot length 3.04 mm , head length 7.42 mm , head width 7.58 mm , eye diameter 3.34 mm , tympanum diameter 1.25 mm , tibia length 11.8 mm , femur length 10.88 mm , arm length 4.55 mm , eye-nostril distance 1.66 mm . Body as wide as head, head wider than long; snout short, truncated in dorsal view and profile; distance from nostril to eye shorter than eye diameter; canthus rostralis barely distinct, rounded; loreal region slightly concave; internarial area subtly depressed; nostrils slightly protruding, directed posterolaterally; interorbital region flat; eyes large, prominent; eye diameter 2.7 times tympanic annulus diameter; tympanum hidden beneath skin; tympanic membrane absent, tympanic annulus visible under skin, oval, longer dorsoventrally and concealed dorsally by supratympanic fold, separated from eye by about 57 % of its diameter; faint supratympanic fold, extending posteriorly from rear corner of eye to anterior edge of arm insertion. Arm sturdy and short, axillary membrane extends halfway to elbow; relative length of fingers I <II <IV <III; fingers with large rounded discs, that of Finger III approximately the same diameter as tympanum; subarticular tubercles prominent, round, subconical, single; distal tubercle on Finger IV bifid; supernumerary tubercles present; palmar tubercle indistinct; prepollex sharp-pointed (based on x-ray image); nuptial excrescences absent; hand webbing formula I basal II 1 ½ — 2 ½ III 2 ⅔ — 2 IV. Hindlimbs large; toes with discs slightly wider than long, smaller than those on fingers; relative toe length I <II <V <III <IV; outer metatarsal tubercle indistinct, small, round; inner metatarsal tubercle large, elongated, elliptical; subarticular tubercles single, round, flat; supernumerary tubercles limited to soles, small; foot webbing formula I 1 + — 2 II 1 + — 2 – III 1 + — 2 IV 2 + — 1 + V (Fig. 11 ). Two pectoral glands bellow the insertion of the arms, separated from each other by about half their individual width. Skin on dorsum, head, flanks, and dorsal surfaces of limbs smooth. Ventral skin coarsely granular, except for pectoral glands, which are finely granular. Ventral surfaces of body and thighs granular. Longitudinal wrinkles of vocal sac from throat to front of the insertion of the arms. Skin on arms and shanks smooth. Cloacal opening directed posteriorly at upper level of the thighs, covered by a long, broad cloacal sheath. Tongue broadly heart-shaped, free laterally and posteriorly, widely attached to mouth floor. Vomerine teeth between choanae, in two slightly angled rows, not contacting each other, smaller than ovoid choanae. In life dorsal background coloration is brown with yellow dorsolateral bands, extending to the eyes and top of head (Fig. 8 ); bright yellow well defined sacral mark; absence of a light labial stripe and large suborbital mark; the dorsal surfaces of the thighs is light brown; the forearm and shank have two yellow transversal marks each; coloration of webbing and ventral surfaces yellow; iris coppery bronze. Color of holotype in preservative shown in Fig. 12 . Figure 11. Metamorph, juvenile and subadults of Dendropsophus cannatellai sp. nov. Dorsolateral views of individuals raised in captivity in Balsa de los Sapos. From left to right, first row: QCAZA l 78576 (total length = 23.27 mm); QCAZA n 79747 (SVL = 11.57 mm); second row: (18.46 mm); (19.59 mm); third row: (19.38 mm); (17.89 mm). All subadults approximately 1 year old since hatching. Figure 12. Dorsal, ventral and lateral views of the holotype of Dendropsophus cannatellai sp. nov. ( QCAZ 40847, SVL 22.32 mm). Figure 13. Ventral views of the holotype’s hand (left) and foot (right) of Dendropsophus cannatellai sp. nov. ( QCAZ 40847). Variation. Sexual dimorphism is observed in the size of adult individuals, where males are smaller (SVL males = 21.05–26.58 mm ; SD = 1.235. SVL females = 28.25–31.94 mm ; SD = 2.078). Color variation in life and preservative is shown in Figs 8 , 9 . In some individuals (e. g., QCAZ 55555) the dorsolateral bands are extensive and fused with the sacral mark. There is variation in the extent of membrane between the digits of the forelimbs, particularly between fingers II, III, and IV, where adult females have more extensive membrane. In dorsal view, snout shape varies from truncate to slightly rounded. In preserved male specimens, the vocal sac is evident as wrinkles in the gular region. However, in some individuals, these wrinkles are less noticeable. Distribution, natural history, and conservation status. Dendropsophus cannatellai sp. nov. occurs in Western Foothill Forest and the Chocó Tropical Rainforest in NW Ecuador and SW Colombia . Although the occurrence in Colombia needs to be corroborated, we found the following evidence of its presence: (1) localities in Ecuador near the border with Colombia , and (2) records on iNaturalist (observation numbers 147931034 in Tumaco, Nariño , and 36196673 in San Andrés de Tumaco), morphologically indistinguishable from the new species. Figure 14. Life coloration of Dendropsophus gryllatus . First and third rows, dorsolateral views; second and fourth, ventral views. From left to right, first and second rows: QCAZ 75534 (SVL = 20.80 mm); QCAZ 75535 (SVL = 21.19 mm); QCAZ 75536 (SVL = 22.24 mm); third and fourth rows: QCAZ 75538 (SVL = 18.71 mm); QCAZ 75539 (SVL = 21.89 mm); QCAZ 75540 (SVL = 20.26 mm); all adult males. See Suppl. material 1 , table S 1 for locality data. All specimens are shown at the same scale. Within its distribution range, climatic conditions are generally warm and humid, with an annual average temperature ranging from 22.5 to 25.4 ° C and an average annual precipitation with a range from 1025 to 3348 mm . The physiognomy of these ecosystems consists of big trees that can reach almost 30 m tall, and a great abundance of palms. The Western Foothill Forest has an elevation range from 300 to 1300 m , with a dominance of plants from the families Mimosaceae , Fabaceae , and Burseraceae with a high percentage of endemism. The presence of epiphytes is common. The Chocó Tropical Rainforest has an elevation from 0 to 300 m , with an abundance of ferns and a high diversity of trees ( Ron et al. 2024 ). Figure 15. Adult preserved specimens of Dendropsophus gryllatus showing variation in dorsal and ventral coloration. From left to right, first and second rows: QCAZ 75532 (22.54 mm); QCAZ 75533 (21.96 mm); QCAZ 75535 (21.19 mm); third and fourth rows: QCAZ 75536 (22.24 mm); QCAZ 75537 (22.85 mm); QCAZ 75539 (21.89 mm); all adult males. See Suppl. material 1 , table S 1 for locality data. All specimens are shown at the same scale. The new species is found in artificial open areas, near ponds and puddles where herbaceous vegetation is abundant. It is less frequent in secondary forest. Males call from leaves or grasses less than 2 m above water, between the 18: 00 and 0: 00 hours. A couple collected in Río Silanche, laid a clutch in captivity with 190 eggs . Experiments in captivity report average clutch sizes of 84 eggs under shaded conditions and 98 under unshaded conditions ( Touchon et al. 2024 ). In the same experiments, most females laid terrestrial eggs, but 30 % of females laid a few eggs in water, under unshaded conditions. Figure 16. Distribution of the species Dendropsophus cannatellai sp. nov. and D. gryllatus . Localities are based of specimens deposited at Pontificia Universidad Católica del Ecuador, QCAZ Museum, Duellman (1974) , and Michelsohn et al. (2022) . Shapes with white outlines represent type localities. All known localities for both species are shown. The new species is assigned to the Red List category Near Threatened ( NTU ). Its extent of occurrence ( EOO ) has < 20000 km 2 (minimum convex polygon of the known localities of D. cannatellai sp. nov. ~ 6134 km 2 ). However, populations are predominantly located on disturbed and deforested areas, near roads. The species appears to thrive in human modified habitats including pastures for cattle raising and ponds in artificial open areas with abundant herbaceous vegetation. Etymology. The specific name cannatellai is a noun in the genitive case and is a patronym for David C. Cannatella, who collected type specimens of the new species. Moreover, during his career he has contributed extensively to the study of systematics and evolution of neotropical amphibians. He has been integral part of scientific societies, exemplified by his presidencies of the American Society of Ichthyologists and Herpetologists (2002–2003) and the Society of Systematic Biologists (2004–2005). Currently he is Dean of the Department of Integrative Biology at the University of Texas at Austin, USA .