Boana platanera, Escalona & Marca & Castellanos & Fouquet & Crawford & Rojas-Runjaic & Giaretta & Señaris & Castroviejo-Fisher, 2021

Escalona, Moisés, Marca, Enrique La, Castellanos, Michelle, Fouquet, Antoine, Crawford, Andrew J., Rojas-Runjaic, Fernando J. M., Giaretta, Ariovaldo A., Señaris, J. Celsa & Castroviejo-Fisher, Santiago, 2021, Integrative taxonomy reveals a new but common Neotropical treefrog, hidden under the name Boana xerophylla, Zootaxa 4981 (1), pp. 401-448 : 413-423

Boana platanera sp. nov. La Marca, Escalona, Castellanos, Rojas-Runjaic, Crawford, Señaris, Fouquet, Giaretta, and Castroviejo-Fisher

( Figs. 2 , 4–6 , 8J , 10 )

Suggested common name: Banana tree dwelling frog (rana platanera, in Spanish).

Boana xerophylla —populations north of the Orinoco River, Dubois (2017); Rojas-Runjaic & Infante-Rivero (2018); populations north of the Orinoco River, Barrio-Amorós et al. (2019); populations north of the Orinoco River, Escalona et al. (2019a); Cañizales (2020).

Hypsiboas xerophyllus —populations north of the Orinoco River, Orrico et al. (2017).

Hypsiboas crepitans —populations north of the Orinoco River, Faivovich et al. (2005); populations north of the Orinoco River, Köhler (2010); Lehtinen (2014); Guarnizo et al. (2015).

Hyla crepitans — Lutz (1927); Shreve (1947); Alemán (1953); Ginés (1959); Röhl (1959); Rivero (1961); Fouquette (1966); Solano de Chacín (1968); Kenny (1969); populations north of the Orinoco River, Duellman (1970, 1997, 2001); populations north of the Orinoco River, Cochran & Goin (1970); Donoso-Barros & León (1972); Yústiz (1976); Kluge (1979); La Marca (1992); Kime et al. (2000); Lynch & Vargas Ramírez (2000); Lynch & Suárez-Mayorga (2001); Bernal et al. (2004); Nava (2005); populations from Colombia and Panama, Vanzolini & Myers (2015).

Holotype. Adult male, ULABG 7758 (field number MES 018; Figs. 2 , 4 ), from 2 km of La Soledad , on the road from Barinitas to Santo Domingo, Barinas state, Venezuela (08º48’26’’ N, 70º30’46’’ W [datum: WGS 84]; 947 m asl; Fig. 7A ), collected by Enrique La Marca, Moisés Escalona, and Iván Mendoza, on June 13, 2012.

Paratypes. Three adult male specimens, ULABG 7759–7761 , topotypes, same data as holotype ( Fig. 5 ). Three adult male specimens, ULABG 7762–7764 , from La Soledad, Barinas state, Venezuela (08º48’35’’ N, 70º31’31’’ W [datum: WGS 84] 980 m asl), collected by Enrique La Marca, Moisés Escalona, and Iván Mendoza, on June 13, 2012 .

Referred specimens. Up to 497 specimens were used to assess the genetic and phenotypic variation, although they are not included in the type series. In addition, we analyzed the calls of eleven non-vouchered specimens. See Appendices 1–3 for details.

Generic placement. The new species is assigned to Boana , within the B. faber group (sensu Faivovich et al. 2005), based on the results of the phylogenetic analyses ( Fig. 1 ).

Definition. The new species is defined by the combination of the following characters: (1) moderately large size, snout-vent length (SVL) 54.33–74.74 mm in females (n = 76) and 46.57–63.34 mm in males (n = 195); (2) head narrower than body, flat in dorsal view; (3) snout truncated (slightly rounded in some individuals) in dorsal and lateral view; nostrils protuberant; (4) horizontal pupil; first half of palpebral membrane densely pigmented with black dots; (5) supratympanic fold extents posteriorly from posterior eye corner and slopes downward to a point above arm insertion; (6) distance between tympanum and eye equal to or shorter than half tympanum diameter; (7) forearm relatively more robust than arm; fingers moderately long and slender with large rounded discs; (8) adult males with prepollical spine, protruding through distal end in some cases; (9) hand-web formula I 0–0 II (1–1 +)– (1–1 +) III 1.5–(2 - –2 +) IV; (10) foot-web formula I 1.5–1 + II (2 - –2 +)–(2–3 -) III 3–(2 - –3 -) IV (3–3 -)–(3 - –3 +) V; (11) cloacal opening at thighs upper level, with supracloacal fold directed posteriorly; pericloacal region dark-brown with a white-colored crenulated supracloacal fold; (12) skin areolate on throat, chest, belly, and ventral surfaces of thighs; (13) dorsal skin smooth, slightly granulated on flanks; (14) poorly developed ulnar fold extending along finger IV external side; (15) tarsal fold poorly evident; (16) calcars absent; (17) adult males with vocal sac single, median, subgular, and moderately distensible ( Fig. 6H ); (18) dorsal coloration is variable at night, from almost uniform pale yellowish ocher, reddish brown, to conspicuously marbled with more or less well-defined dark-brown marks, and an irregular or broad X-shaped on the mid dorsum ( Fig. 6 ); (19) in preservative, palms and soles pale cream-colored, hand and foot webs cream with dark brown flecks, especially on foot.

Diagnosis. Character states of compared species are given in parentheses. Boana platanera is sister of B. xerophylla ( Fig. 1 ) and it can be diagnosed from the latter by the following combination of adult male characters: a pale orange-yellow or light brown coloration on dorsum (dark brown to green tones; Fig. 8I, J ); presence of black or dark brown speckles in the anterior half of the palpebral membrane (absent; Fig. 9 ); pericloacal region dark brown (cream); advertisement call with shorter, 28−71 ms, first note length (79–124 ms in call type I and 85−110 ms in call type II), composed of five notes (one note in call type I). Although not diagnostic, the following characters may help to differentiate both species: the new species has a statistically significant larger body (46.57–63.34 mm) than B. xerophylla (45.13–56.71 mm) ( Table 2 in Escalona et al. 2019a); in preservative, the gular region is cream with dark brown or black flecks near to the jaw border (cream and sometimes with few dark brown speckles on the border of the jaw; Fig. 9 ); advertisement call with statistically significant longer pulse interval (11−23 ms) than B. xerophylla (12–14 ms in call type I and 11−14 ms in call type II); statistically significant longer second phase length (137−312 ms) than B. xerophylla (73−150 ms in call type II).

Boana platanera is sympatric with B. pugnax in lowland western and central Venezuela, as well as some regions in Colombia and Panama. The new species is distinguished from B. pugnax by having a pale orange-yellow or light brown coloration on dorsum (dark-brown to dark-grey tones; Fig. 8G, J ); the anterior half of the palpebral membrane densely pigmented (poorly pigmented); dorsal surfaces of finger and toe discs usually cream, when pigmented, pale gray (dark); calcar absent (poorly developed); a complex advertisement call composed of five notes (simple call of one note). Although not diagnostic, B. platanera is distinguished from B. pugnax by having a statistically significant smaller body (males: B. platanera : 46.57–63.34 mm; B. pugnax : 55.85–74.80 mm [sample from western Venezuela]; Table 4 ); and a statistically significant longer advertisement call ( B. platanera : 200−451 ms; B. pugnax : 156–258 ms).

The new species differs from Boana crepitans by having a pale orange-yellow or light brown coloration on dorsum (gray-cream tones to dark brown; Fig. 8H, J ); advertisement call composed of five notes (two notes). Although not diagnostic, the new species has a statistically significant shorter ETD (eye to tympanum distance; B. platanera : 0.96–2.78 mm) than B. crepitans (1.99–3.29 mm) ( Tables 1–2 in Escalona et al. 2019a); statistically significant shorter advertisement call ( B. platanera : 200−451 ms; B. crepitans : 368−473 ms), with a statistically significant shorter second phase ( B. platanera : 137−312 ms; B. crepitans : 278−367 ms) ( Tables 2 , 3 in Escalona et al. 2019a).

Boana platanera differs from B. rosenbergi by having a smooth dorsum, thighs, and flanks (rough or with tubercles in at least one of these regions; Fig. 8F, J ); smaller body size (males: B. platanera : 46.57–63.34 mm; B. rosenbergi : 75.55–83.98 mm); presence of black dots on anterior half of the palpebral membrane (absent); ulnar fold smooth (slightly crenulated); and a complex advertisement call (simple in B. rosenbergi ) with shorter duration ( B. platanera : 200−451 ms; B. rosenbergi : 551–759 ms) ( Table 2 ).

The new species differs from Boana exastis by having a smooth dorsum, thighs, and flanks (rough or with tubercles in at least one of these regions; Fig. 8D, J ); smaller body size (males: B. platanera : 46.57–63.34 mm; B. exastis : 81.1–99.0 mm); presence of black dots on anterior half of the palpebral membrane (absent); poorly-defined dermal fringes on limbs (crenulated dermal fringes), less webbing in hands and feet (more extensive webbing in hands and feet), absence of cloacal plate (presence); absence of calcars (presence); complex advertisement call (simple in B. exastis ), with shorter duration ( B. platanera : 200−451 ms; B. exastis : 807–1229 ms) ( Table 2 ).

Boana platanera differs from B. lundii by having a smooth dorsum, thighs, and flanks (rough or with tubercles in at least one of these regions; Fig. 8E, J ); presence of black dots on anterior half of the palpebral membrane (absence); poorly-defined dermal fringes on limbs (well defined); less webbing in hands and feet (more webbing in hands and feet); absence of calcars (presence); complex advertisement call (simple), with shorter duration ( B. platanera : 200−451 ms; B. lundii : 834–1228 ms) ( Table 2 ).

The new species differs from Boana pardalis by having a smooth dorsum, thighs, and flanks (rough or with tubercles in at least one these regions; Fig. 8C, J ); presence of black dots on first half of the palpebral membrane (absence); poorly-defined dermal fringes on limbs (crenulated); less webbing in hands and feet (more webbing in hands and feet); absence of cloacal plate (presence); absence of calcars (presence); complex advertisement call (simple), with longer duration ( B. platanera : 200−451 ms; B. pardalis : 117–143 ms), composed of 4–6 notes (one) ( Table 2 ).

Boana platanera differs from B. faber by having a smaller body size (males: B. platanera : 46.57–63.34 mm; B. faber : 77.90–97.08 mm); presence of black dots on anterior half of the palpebral membrane (absence); ulnar fold present (absent); cream gular coloration with dark-brown flecks near to the jaw border (dark); complex advertisement call (simple), longer advertisement call ( B. platanera : 200−451 ms; B. faber : 75–121 ms), composed of 4–6 notes (one note), with higher fundamental frequency ( B. platanera : 542.7−1125.0 Hz; B. faber : 344.5–447.9 Hz) ( Table 2 ).

Boana platanera is readily distinguishable from B. albomarginata by having a smooth dorsal skin (slightly granular in B. albomarginata ), a pale orange-yellow coloration with dark-brown marks, irregular or broad X-shaped on the dorsum and dark-brown bars on flanks and thighs (green with some white spots on the dorsum; Fig. 8A, J ); supratympanic fold slopes downward to a point above arm insertion (supratympanic extends posteriorly until half of the laterodorsal region); absence of calcar (presence); complex advertisement call (simple), with longer duration ( B. platanera : 200−451 ms; B. albomarginata : 166–185 ms), with 4–6 notes (one note), and lower fundamental frequency ( B. platanera : 542.7−1125.0 Hz; B. albomarginata : 1136.9–1231.7 Hz) ( Table 2 ).Although not diagnostic, the new species is larger bodied (males: B. platanera : 51.97–59.08 mm; B. albomarginata : 44.68–53.6 mm).

Holotype description. Adult male in a good state of preservation ( Fig. 4 ), with a piece of tissue sampled from the right thigh. Morphometric measurements of the holotype are presented in Table 5 . Body slender; head wider than long (HW/HeL = 1.06); snout truncated in lateral and dorsal views; nostrils protuberant, oblique and directed dorsolaterally; internarial distance smaller (66 %) than eye-to-nostril distance; eye diameter, upper eyelid width and interorbital distance approximately equal to tympanum diameter; canthus rostralis rounded, almost indistinct; loreal region concave; lips not flared; eyes large (ED/HeL = 0.28), protruding; interorbital region and dorsal surface of snout slightly concave. Tympanum conspicuous, nearly circular; tympanic membrane differentiated; tympanic annulus well-defined, with upper border concealed by the tympanic membrane. Vocal sac single, subgular; vocal slits large, located at the posterolateral edge of the mouth floor. Tongue wider than long, cordiform, slightly notched behind; almost completely attached to mouth floor. Choanae elliptical, widely separate. Vomerine teeth massive; vomerine processes arched, slightly separate and between choanae.

Arms slender; forearms more robust than upper arms; an inconspicuous crenulated fringe along forearm ventroexternal surface and prolateral side finger IV (more conspicuous on right forearm). Hand with an elliptical and flat palmar tubercle and an inconspicuous thenar tubercle; prepollex well-developed, with a single curved spine ( Fig. 4 ); numerous small supernumerary tubercles scattered on the palm; relative fingers length: III>IV>II>I; subarticular tubercles single, rounded to conical and prominent; fingers discs large, expanded and nearly circular; disc on first finger slightly smaller than those of other fingers; third finger disc diameter about 50 % of eye diameter and approximately 60 % of tympanum diameter; hand-webbing formula: I 0–0 II 1 + –1 III 1.5–2 + IV.

Legs long and slender; thigh length slightly longer than shank length (FL/TL = 1.01); without calcars—the protrusion of the skin fold on the right heel, which resembles a calcar, is caused by the fixation position of the hind limbs, with the feet oriented posteriorly; tarsal fringe not evident; foot with an elliptical inner metatarsal tubercle; outer metatarsal tubercle not evident; subarticular tubercles single, rounded and prominent; supernumerary tubercles scarce, small, and flat; toes extensively webbed, with rounded and expanded disks; foot-web formula: I 1.5–1 + II 2 + –2 III 3–3 - IV 3–3 + V. Skin on dorsal surfaces smooth; throat, chest, belly and ventral surface of thighs areolate; supracloacal fold crenulated; pericloacal region tuberculate.

Color of euthanized holotype. The holotype was photographed two minutes after anesthesia and the following description was based in those photographs. Dorsum with a large dark brown irregular blotch over an olive background; flanks olive with vertical narrow brown bands; groin and postero-ventral part of flank orange; throat and chest dark gray; chin olive with dark brown markings; upper part of venter gray; middle and lower part of venter reddish orange, with gray tubercles; upper parts of extremities gray to olive, with brown bands or markings; foot web reddish; iris light greenish gray with fine dark venation.

Color of holotype in preservative. Dorsum pale brown with dark-brown irregular blotches without a defined pattern, darker on dorsum than on head ( Fig. 4 ); upper surface of arms pale brown, with one dark-brown incomplete band on forearm, wider on the external edge; dorsal surface of hands with one dark-brown incomplete band (more evident on the right hand); third and fourth fingers with the same coloration of arms; first and second fingers pale cream colored; dorsal surface of thighs pale brown with dark brown bars that form an irregular reticular pattern; dorsal surface of shanks, metatarsus and toes pale brown with dark brown blotches without a defined pattern; webbing on hands cream colored with dark brown flecks. Flanks cream with dark-brown vertical lines. Pericloacal region dark brown surrounded by white.

Ventral surfaces of arms, forearms and hand palms pale cream; pattern on dorsal surface of thigh partially extended to ventral surface; lower edge of thighs pale cream; shanks pale brown with dark brown blotches; metatarsus, feet and foot-web cream with numerous dark brown flecks; throat cream with dark brown flecks, mostly in borders of the mouth; chest and belly pale cream.

Variation in morphology. The variation of external measurements, as well as hand and foot webbing formula of some type specimens and from all the specimens examined is shown in Table 5 . There is sexual dimorphism in body size, with males generally smaller than females (males: 46.57–63.34 mm; females: 54.33–74.74 mm), and by the presence of a prominent prepollical spine in males ( Figs. 5 , 6 ).

Color variation in life. When active at night, adult specimens have a yellow, tan, to light brown coloration, with brown patterns (irregular in shape or with a broad X-shaped blotch near the head; some specimens have a brown vertebral line) on dorsum ( Figs. 6A , 7C ). Brown dots are spread on the flanks, above the upper lip, on the limbs, and on the webbings. Some specimens have a brown line delimiting the supratympanic fold. Some specimens have apparent scars on the dorsum. There is a brown blotch with a white line on the upper margin on heels and cloacal region. There are several vertical brown bars on flanks and transversal brown bars on the thighs. White coloration predominates along all the ventral surfaces, with strong orange or pinkish tones on the belly, arms, and thighs. In some specimens, the dorsal brown bars of the thighs are projected on the internal portion. The tarsus has two colorations delimited by the tarsal fold. The outer portion of the tarsus has a brown coloration with orange tones, while the internal portion has orange coloration. Some specimens have some brown spots around the mandible, on the upper portion of the thigh, and/or the webbing at the hands and foots. During the day, when normally are inactive, they have a pale cream coloration ( Fig. 6E ). The brown patterns and spots, when visible, are pale gray, and the bars on the flanks and thighs are dark brown almost black. Juvenile specimens have a pale green coloration with black diffuse dots on the dorsum ( Fig. 6F ). During the night, the iris around the pupil is metallic gray color and the outer area is pale emerald ( Fig. 6A, C, D ).

Color variation in preservative. Coloration pattern of paratopotypes in preservative is depicted in Fig. 5 . Specimens from localities other than the type locality show some color variation, as follows: dorsum occasionally may be uniformly pale-brown (e.g., ULABG 2950 , 2951 , 7755 , 7803 ) , sometimes bearing two scapular dark brown spots (e.g., ULABG 5338 ) , or few scattered small dots on sacral region (e.g., ULABG 5337 , 5338 ) or dorsum (e.g., ULABG 7745 ) . Some specimens tend to have an ashy background with irregular markings (e.g., ULABG 5060 , 6902 , 7753 ) and, very rarely, have a lichen aspect (e.g., ULABG 7756 , 7757 ) . More often, dorsum bears large blotches that may be irregular in shape (e.g., ULABG 1416 , 3725 ) , or sometimes with a broad X-shaped brown blotch (e.g., ULABG 2544 , 4749 , 4754 ) that could tend to be diffuse (e.g., ULAB 2949 ) . Specimen ULABG 7744 has a white supratympanic line from posterior border of eye to posterior part of tympanum, and a white line along border of upper lip. Specimen ULABG 1417 has a dark brown line on middle of dorsum, from almost tip-of-snout to sacral region. Forearms and wrists may have brown bands (e.g., ULABG 2544 ) . Shanks and thighs may have bands with ill-defined borders (e.g., ULABG 2544 ) .

Natural history. All specimens from the type series were collected after dusk (around 20:00 h) in a sewage system alongside a paved road, in steep terrain, 947 m asl. These were found active on the floor, many of them vocalizing. Surrounding vegetation was disturbed second-growth forest, with primary humid forest surrounding it ( Fig. 7B ). No other anuran species was heard or seen at this site.

The additional museum specimens reviewed came from the Tropical and Premontane altitudinal belts of the Holdridge’s system as applied to Venezuela ( Ewel et al. 1976), within the life zones of Tropical dry forest and Tropical humid forest, as well as Premontane dry forest and Premontane humid forest. The associated vegetation is usually that of open disturbed areas, with abundance of grasses and few trees. Field records of ELM for some of the referred specimens indicate that the species lives in places with annual mean temperatures higher than 17 ºC and mean annual precipitation roughly between 600 and 1,500 mm. Two precipitation regimes are associated with the distribution of this species. One is the bi-seasonal pluviometry regime, with a single wet season around the middle of the year (June and July). The other is the tetra-seasonal regime with two wet seasons separated by a short midyear period of less precipitation ( Monasterio & Reyes 1980).

Males vocalize on vegetation up to 5 m ( Fig. 6G ), but generally from the ground, on rocks, or partially submerged or floating in the water, although the latter behavior is usually observed in artificial and temporary pools ( Fig. 6H ). Field observations indicate that the amplexus in Boana platanera is axillary (sensu CarvajalCastro et al. 2020), with the toes projected above the female shoulders ( Fig. 10 ). Scars on the dorsum of several individuals suggest the existence of male to male combat in this species. This species can form choruses and males are separated from each other by at least one meter. In addition, this treefrog usually lays eggs in rafts in lentic aquatic systems. The following species were detected sharing the same breeding sites as B. platanera : B. lanciformis , B. pugnax , Dendropsophus microcephalus , Engystomops pustulosus , Leptodactylus fuscus , Pleurodema brachyops , Pseudis paradoxa , Rhinella horribilis , R. marina , Scarthyla vigilans , and Scinax rostratus .

Distribution. Boana platanera is widely distributed north of the Orinoco River in Venezuela (states of Anzoátegui, Apure, Aragua, Barinas, Carabobo, Cojedes, Distrito Capital, Lara, Mérida, Miranda, Portuguesa, Sucre, Táchira, Trujillo, Vargas, Yaracuy, and Zulia; Fig. 7A ; Appendix 3), although it is unclear if it is present in Nueva Esparta state and other islands near to mainland Venezuela. Based on our phylogenetic tree, it is also present in Panama, Colombia, and Trinidad and Tobago. Therefore, we assign all the populations of B. xerophylla s.l. north of the Orinoco River to B. platanera . The localities from Colombia and Panama published by Kluge (1979) and Lynch & Suárez-Mayorga (2001) are plotted in Fig. 7A .

Conservation status. Venezuelan populations of Boana platanera (as Hyla crepitans ) were referred as abundant by Donoso-Barros & León (1972). This abundance is indirectly evidenced by the hundreds of samples housed at Venezuelan museums. Two recent studies provided quantitative data about how its abundance varies geographically (referred as B. xerophylla ). Rojas-Runjaic & Infante-Rivero (2018) reported that B. platanera is rare between 1,500 and 2,000 m asl in the Sierra de Perijá. Cañizales (2020) reported this species as abundant in one locality (around 100 m asl) in western Venezuela. Our field observations agree with these observations, and point out that B. platanera is especially common and very abundant throughout the year in anthropic environments with aquatic habitats. Infection by the chytrid fungus Batrachochytrium dendrobatidis (Bd) was reported, with low prevalence (1/20), in samples from the Cordillera de Mérida (referred as Hypsiboas crepitans ) by Sánchez et al. (2008), absent in 16 tested individuals from Montalbán, Carabobo state, Venezuela (Señaris & Lampo, unpublished data), and present in 6 of 50 samples from across Colombia ( Flechas et al. 2017). Experiments by Márquez et al. (2010) found that B. platanera specimens (referred as Hypsiboas crepitans ) from the Cordillera de Mérida are susceptible to Bd infection, but they have the ability to overcome it within 38 days at a temperature (23 °C) purportedly ideal for Bd growth. Following the criteria of the IUCN Red List categories ( IUCN 2012), we propose that B. platanera should be listed as Least Concern (LC) based on its wide geographical and elevational distribution in a broad range of habitats, its tolerance for habitat modification, and presumed large and stable populations.

Etymology. The specific epithet platanera is an adjective used in apposition. It is the common name given in Colombia and Venezuela to this kind of relatively large treefrogs (i.e., Boana platanera and B. pugnax ). The epithet “platanera” refers to the plantain or banana plant (“ plátano ”; hence, “from the plantain”), where these frogs are commonly found in the daytime. This name is widely used since the species is abundant, easy to spot, present in anthropic environments, and widely distributed. Although B. platanera and B. pugnax have been misidentified in the past ( La Marca 1996; Escalona et al. 2017), our intention is to use the folk nomenclature as the species epithet so that it can be easily remembered by non-taxonomic experts.

Figures

FIGURE 1. Maximum likelihood phylogenetic tree of the Boana faber species group and outgroups inferred from concatenated DNA sequences of fragments of the mitochondrial genes 12S, 16S, and COI, totaling 2,820 aligned base pairs. The best models of molecular evolution were used per fragment and codon position. Numbers on internal branches are the percentage (≥ 50 %) of 1,000 nonparametric bootstrap pseudoreplicates. The updated taxonomy of B. xerophyla following the results of this work is on the right of the black bars.

FIGURE 2. Calls of Boana platanera sp. nov. (A, B, E, F, I and J) and B. xerophylla (C, D, G, H, K and L). (A and E) Oscillograms and (I) spectrogram of the advertisement call (call type I) of B. platanera. Illustrated calls from the holotype, ULABG 7758, temperature and humidity at time of recording unknown (recording “MES_SO1”). (B and F) Oscillograms and (J) spectrogram of the presumably territorial call (call type II) of B. platanera. Illustrated calls from a non-collected specimen from Caparú lagoon, Mérida state, Venezuela, temperature and humidity at time of recording unknown (recording “MES_CA6”). (C and G) Oscillograms and (K) spectrogram of the advertisement call (call type I) of B. xerophylla. (D and H) Oscillograms and (L) spectrogram of the presumably territorial call (call type II) of B. xerophylla. Both illustrated calls from specimen AAG- UFU 5550, from Rancho do Papito, road BR-432, on the road from Boa Vista to Cantá, Roraima state, Brazil, air and water temperature at time of recording 26 °C and 32 °C, respectively (recording “Hypsib_crepitansCantaRR2bAAG”).

FIGURE 4. Dorsal and ventral view of Boana platanera sp. nov. preserved holotype, ULABG 7758 (field number MES 018) from 2 km of La Soledad, on the road Barinitas to Santo Domingo, Barinas state, Venezuela. White scale bar represents 20 mm. Photos: Michelle Castellanos and Enrique La Marca.

FIGURE 5. Dorsal and ventral views of preserved paratypes, ULABG 7759–7764. Details in the text. Photos: Michelle Castellanos and Enrique La Marca.

FIGURE 6. Specimens of Boana platanera sp. nov. in life. Note the variation in coloration at night (A–D, G, H), the difference of the coloration during the day (E), and the coloration of a juvenile at night (F). All the specimens from Venezuela: (A, B) adult male not collected from Sabaneta sector, El Hatillo, Miranda state; (C) adult male from Río Cata, Aragua state; (D) adult male from Parque Minas de Aroa, Yaracuy state; (E) adult male from Ipika, Sierra de Perijá, Zulia state; (F) juvenile not collected, sex unknown, from Posada Ecológica Casa María, altos de Canoabo, Carabobo state; (G) adult male calling from Laguna de Caparú, Mérida state; (H) adult male calling from Posada Ecológica Casa María, altos de Canoabo, Carabobo state (not collected). The photos are not to the same scale. Photos: (A–C) Fernando J. M. Rojas-Runjaic; (D) Iván Mendoza; (E) Edwin Infante; (F, H) Celsa Señaris; (G) Moisés Escalona.

FIGURE 7. (A) Map of northern South America with confirmed localities of Boana platanera sp. nov. (circles and star), B. xerophylla (triangles), and B. sp. (squares). The type locality of B. platanera is indicated with a star; yellow symbols indicate localities of specimens with DNA sequence data analyzed in this study; white symbols indicate localities of specimens analyzed morphologically; gray symbols indicate localities of individuals according to the literature (Kluge 1979; Lynch & Suárez- Mayorga 2001). (B) Photograph illustrating the landscape around the type locality of B. platanera. (C) Paratype of B. platanera in life at night (ULABG 7763). Photos: (B) Jesús Molinari; (C) Enrique La Marca.

FIGURE 8. Species of the Boana faber species group in life. Note the difference in coloration, skin texture, supratympanic fold, calcar, tarsal fringe, and hand webbing. (A) Boana albomaginata; (B) B. faber; (C) B. pardalis; (D) B. exastis; (E) B. lundii; (F) B. rosenbergi; (G) B. pugnax; (H) B. crepitans; (I) B. xerophylla; and (J) B. platanera sp. nov. Photographs are not at the same scale. Photos: (A, B, J) Fernando J. M. Rojas-Runjaic; (C, E) Mario Sacramento; (D) Mauro Texeira Jr.; (F) Santiago Ron (Anfibios del Ecuador); (G) Iván Mendoza; (H) Omar Padilla; (I) Santiago Castroviejo-Fisher.

FIGURE 9. Lateral view of head of (A) Boana platanera sp. nov. (MHNLS 20970) and (B) B. xerophylla (MHNLS 15062). Ventral view of the anterior half of (C) B. platanera (MHNLS 5567) and (D) B. xerophylla (MHNLS 15062). The black arrows in the gular region indicate the difference in pigmentation between the two species. Photos are not at the same scale. Photos: Michelle Castellanos.

FIGURE 10. Axillary amplexus in (A) Boana platanera sp. nov. from Sabaneta sector, El Hatillo, state of Miranda, Venezuela, and (B) B. xerophylla from Reserva Biológica Ilha de Maracá, Roraima, Brazil. Photos: (A) Fernando J. M. Rojas-Runjaic and (B) Antoine Fouquet.