Dendropsophus ozzyi, Orrico, Victor G. D., Peloso, Pedro L. V., Sturaro, Marcelo J., Da Silva-Filho, Heriberto F., Neckel-Oliveira, Selvino, Gordo, Marcelo, Faivovich, Julián & Haddad, Célio F. B., 2014

Orrico, Victor G. D., Peloso, Pedro L. V., Sturaro, Marcelo J., Da Silva-Filho, Heriberto F., Neckel-Oliveira, Selvino, Gordo, Marcelo, Faivovich, Julián & Haddad, Célio F. B., 2014, A new “ Bat-Voiced ” species of Dendropsophus Fitzinger, 1843 (Anura, Hylidae) from the Amazon Basin, Brazil, Zootaxa 3881 (4), pp. 341-361 : 343-354

Dendropsophus ozzyi , new species

(Figs. 3–6)

Holotype: MPEG 27811, an adult male, collected on vegetation of pond within the forest, at Comunidade Bragança, Rio Paraconi, Floresta Nacional de Pau-Rosa ( FNPR), Maués, State of Amazonas, Brazil (03°56’50”S and 58°26’36”W, 45 meters above sea level [m a.s.l.]) on 25 February 2009, by M.J. Sturaro and P.L.V. Peloso (Figs 3–4).

Paratopotypes: MPEG 27809–27814, five adult males, collected on 20 February to 0 3 March 2009, other data same as holotype.

Paratypes: INPA-H 32742–32748, seven adult males on 23 June 2010, and 21 March 2011; CFBH 35760 and 35761, two adult males, on 23 June 2010, all collected by M. Gordo; MPEG 22400–22404, four adult males and one adult female, on 0 7 March 2008, collected by S. Neckel-Oliveira and H. F. Silva-Filho; all from Igarapé Prudente (02°27'13.03"S; 56°10'54.14"W), Juruti, State of Pará, Brazil. CFBH 35762–35763, two adults males, on 19 march 2011; CFBH 35764, an adult male, on 7 May 2013; MZUSP 154084–154088, five adult males, on 19 March 2011; MNRJ 86921–86925, five adult males, on 0 7 May 2013, all collected by M. Gordo; MPEG 22352–22355, three adult males and one adult female, on 0 8 March 2008, collected by S. Neckel-Oliveira and H. F. Silva-Filho; all from Igarapé Mutum (02°36'46.09"S; 56°11'38.53"W), Juruti, State of Pará, Brazil.

Generic placement: We assign the new species to the genus Dendropsophus on the basis of its overall morphological similarity with other species of the genus (Figs. 3–6). Overall morphology (particularly the combination of size, dorsum color, and flash colors) and advertisement calls of D. ozzyi are similar to those of species assigned to the D. microcephalus Group. Within this group, the new species is most similar to D. shiwiarum Ortega-Andrade and Ron 2013 because both species are unique in having pointed discs (Ortega- Andrade & Ron 2013). Dendropsophus shiwiarum is assigned to the D. microcephalus Group largely due to its resemblance to D. riveroi ( Ortega-Andrade & Ron 2013) ; the later was, however, originally assigned to the D. minimus Group and only recently shown to be part of the D. microcephalus Group ( Fouquet et al. 2011). For the above reasons, we tentatively assign D. ozzyi to the D. microcephalus Group.

We could not evaluate the putative morphological synapomorphies of the genus suggested by Faivovich et al. (2005) and we do not know the number of chromosomes for the species described here; a diploid number of 2N = 30 seems to be a synapomorphy for the genus (Suarez et al. 2013). We were also unable to check the proposed larval synapomorphies for the D. microcephalus Group ( Duellman & Trueb, 1983; Faivovich et al. 2005), as no tadpoles have been collected. Thus, we remark that a formal test of the relationships of D. ozzyi is still lacking.

Diagnosis. Dendropsophus ozzyi is diagnosable by the following combination of character states: (1) a medium sized member of the Dendropsophus microcephalus Group, (SVL 18.5–21.5 mm in adult males, N = 37); (2) head wider than body; (3) snout truncate in dorsal and lateral views; (4) nostrils protuberant; (5) large prominent eyes (EL/HW 0.3–0.5, N = 37); (6) nictitating membrane bearing brownish pigmentation on its border; (7) small tympanum (TD/ED 0.3–0.5, N = 37); (8) axillary membrane present; (9) glandular nuptial pads on finger II; (10) hands webbing formula II 2 - – 2- III 2 - – 2- IV 2 + – 2+ V; (11) feet webbing formula I 2 – 2 + II 1 + – 3- III 1 +– 2+ IV 2 + – 1+ V; (12) fingers and toes bearing pointed discs; (13) no inner tarsal fold; (14) calcar tubercle absent, heel with small, inconspicuous tubercles; (15) a dorsal sheath covering 1/3 of the cloacal opening; (16) in life, dorsal surfaces plain light brown with randomly distributed dark smudges; palmar, plantar and ventral surfaces of the axillary membrane, thigh, and shank vivid orange, belly creamy white with clear white spots and a rectangular bright white area, gular region orangish cream with brown flecks, vocal sac transparent; (17) advertisement call composed of a single, not pulsed, tonal, and highly pitched (dominant frequency around 9 kHz) note.

FIGURE 3. Dendropsophus ozzyi sp. nov. holotype (MPEG 27811). (A) Dorsal, and (B) lateral views of head; (C) palmar view of right hand, and (D) plantar view right foot. Scale bar = 5 mm.

Comparison with other species. Dendropsophus ozzyi differs from all other species of Dendropsophus , except D. shiwiarum —also in the D. microcephalus Group—by having pointed discs on fingers and toes. In the following paragraphs we first diagnose our new species from D. shiwiarum and then provided additional diagnostic characters that differentiate D. ozzyi from all other species of the genus arranged by species groups.

Dendropsophus ozzyi differs from D. shiwiarum by (character states of the later in parenthesis; all retrieved from Ortega-Andrade and Ron 2013) by the absence of dark brown interorbital, canthal, and post-orbital stripes extending to mid-flank (present); absence of suborbital bars or spots (present); shanks not presenting bars (present); gular region (and not the vocal sac) orangish cream with brown flecks (bright yellow, unflecked); by the absence of a distinct supratympanic fold (present). These species also differ by their color in life, while D. ozzyi presents vivid orange flash colors, D. shiwiarum presents “fleshy white” (sic) flash colors. “Fleshy” can be interpreted as a variety of colors, including orange. From pictures in Ortega-Andrade & Ron (2013) we understand that the color in D. shiwiarum is better described as pinkish-white.

Morphometrically, Dendropsophus shiwiarum and D. ozzyi are distinguishable by their eye-to-nostril distance (EN = 1.2–1.8 in D. shiwiarum and EN = 2.31–2.37 mm in D. ozzyi ). Advertisement calls of the two species differ by the longer duration of D. ozzyi (248 ms vs. 52.83 ± 31.87 ms of D. shiwiarum [original description does not present range]) and especially by the higher dominant frequency of D. ozzyi (9130.1–10136.7 Hz vs. 3983.6–5254.1 Hz of D. shiwiarum ).

From other species of the Dendropsophus microcephalus Group, D. ozzyi is distinguished from species of the D. rubicundulus Clade sensu Faivovich et al. (2005) (traits of the later in parentheses) by the stouter body (body slender), light brown dorsum in live and preserved specimens (dorsum dark green in life and violet in preserved specimens) with unaligned spots (stripes or aligned spots). From species of the D. decipiens Clade sensu Faivovich et al. (2005) by the absence of dorsal patterns (all species framed, except D. berthalutzae that presents an ‘><’ or an ‘X’) ( Pugliese et al. 2000).

The simple (monophasic) advertisement call also distinguishes D. ozzyi from most Dendropsophus species (see below). The high dominant frequency distinguishes D. ozzyi from all other congeners ( Table 2 ), except from D. minusculus (9020–9360 Hz see Duellman & Pyles 1983) although call duration of D. ozzyi is much longer [165–386 ms vs. 10–46 ms ( Duellman & Pyles 1983; Lescure & Marty 2000 respectively)].

Dendropsophus ozzyi is distinguished from species of the D. columbianus Group by the smaller SVL [combined SVL = 21.1 mm in D. columbianus , to 33.5 mm in D. bogerti (data of both species from Cochran & Goin 1970)], except that the maximum SVL of D. ozzyi is 0.5 mm longer than the minimum SVL of D. columbianus . Apart from that, members of the D. columbianus Group have flecked or marbled bellies, and dorsum with dark spots ( Duellman 1989; Rivera-Correa & Gutiérrez-Cárdenas 2012). Additionally, advertisement calls of species of the D. columbianus Group are diphasic (see Duellman & Trueb 1983) while D. ozzyi has a simple call.

Dendropsophus ozzyi is also smaller than species of the D. garagoensis Group [combined SVL ranges from 21.3 mm in D. virolinensis (see Kaplan & Ruiz-Carranza 1997) to 31.5 mm in D. praestans (see Duellman & Trueb 1983) specimens], except that the maximum SVL of D. ozzyi is 0.2 mm larger then the minimum SVL of D. virolinensis (21.3 mm). Species of the D. garagoensis Group also present different dorsal patterns, with paramedial white lines and dark blotches in the flanks (both traits absent in D. ozzyi ) ( Kaplan 1991; 1997; Kaplan & Ruiz- Carranza 1997). To date, advertisement calls of the D. garagoensis Group are unknown.

From species of the D. labialis , D. marmoratus and D. parviceps Groups , the new species differs by the homogeneous light brown dorsum whereas (green in life fading to gray in preserved specimens in the D. labialis Group and lichenous, both in life and in preserved specimens, in the D. marmoratus and D. parviceps Groups ). From the D. labialis and D. marmoratus Groups by the smaller SVL [combined SVL ranging from 27.5 mm in D. melanargyreus (VGDO unpub. data) to 61 mm in D. labialis , (from Guarnizo et al. 2012)]. Species of the D. marmoratus and D. parviceps Groups have black flash colors [species of the D. parviceps Group with or without bright yellow spots (see Duellman & Crump 1974)], and species of the D. labialis Group have blue colors (e.g., Guarnizo et al. 2012), while D. ozzyi has orange flash colors. Additionally, many species of the D. marmoratus and D. parviceps Groups have suborbital bars (e.g., Duellman & Crump 1974; Lutz & Bokermann 1963). These character states also distinguish D. ozzyi from D. yaracuyanus , a species without formal assignment to a species group ( Mijares-Urrutia & Rivero 2000) but quite similar to species of the D. parviceps Group. Additionally, calls of species of the D. parviceps Group are complex (e.g., Amezquita & Hodl 2004; Duellman & Crump 1974; Marquez et al. 1993) while D. ozzyi has a simple call. Calls of the D. labialis ( Guarnizo et al. 2012) and D. marmoratus Group ( Orrico et al. 2009) are longer and have much lower dominant frequencies.

Dendropsophus ozzyi is smaller than species of the D. leucophyllatus Group [combined SVL ranging from 23 mm in D. bifurcus (see Kaplan & Ruiz-Carranza 1997) to 40 mm in D. anceps specimens (see Duellman & Trueb 1983)]. Exceptions are D. rossalleni [19–22.3 mm ( De la Riva & Duellman 1997)] and smaller individuals of D. elegans [20 mm ( Lutz, 1973)] as D. ozzyi reaches a maximum SVL of 21.5 mm. Additionally, D. ozzyi does not have any interorbital bar or blotch of different color of the dorsum ground color, as observed in all species of the D. leucophyllatus Group. Furthermore, species of the D. leucophyllatus Group have pectoral glandular patches—a likely synapomorphy for the group (see comments in Rivera-Correa & Orrico 2013)—which are absent in D. ozzyi Additionally , species of the D. leucophyllatus Group present diphasic calls (e.g., Conte et al. 2010; Jungfer et al. 2010; Marquez et al. 1993) while D. ozzyi presents a monophasic call.

Dendropsophus ozzyi is distinct from species of the D. minutus Group by possessing a small cloacal sheath partially covering the cloaca (long cloacal sheath covering entirely the cloaca in D. minutus Group) and by the absence of white supracloacal and tarsal lines (present in species of the D. minutus Group). These traits also distinguish D. ozzyi from D. stingi and D. amicorum that present such structure and are quite similar to species of the D. minutus Group although not assigned to any species group in the original description (see Kaplan 1994; Mijares-Urrutia 1998). These same traits also distinguish D. ozzyi from D. aperomeus , a species of the D. minimus Group ( Duellman 1982).

The Dendropsophus minimus Group has been consistently recovered as paraphyletic but to date it is still recognized because D. minimus has never been included in a phylogenetic analysis ( Fouquet et al. 2011); therefore, the status of the group name remains unclear ( Frost 2014). We group the comparisons of D. ozzyi with the remaining species still associated with this possibly paraphyletic arrangement for convenience only, however, not formally recognizing it. Dendropsophus minimus presents a concealed tympanum (clearly visible in D. ozzyi ) a clear tarsal fold (absent in D. ozzyi ), a rostral white line (absent in D. ozzyi ) and a different dorsal pattern ( Ahl 1933). Dendropsophus miyatai presents a concealed tympanum (visible in D. ozzyi ), a bright red over bright yellow, dorsal pattern in life (light brown in D. ozzyi ), and absence of fimbriae on hind limbs (present in D. ozzyi ) ( Vigle & Goberdhan-Vigle 1990). Additionally, calls of the D. minutus Group are complex, diphasic or even triphasic (e.g., Cardoso & Haddad 1984; Duellman 1978; Heyer et al. 1990; Kohler & Lotters 2001b; Marquez et al. 1993; Morais et al. 2012) while D. ozzyi has a simple, monophasic, call.

The following comparisons pertain to species of the genus not assigned to a group ( Faivovich et al. 2005). Dendropsophus ozzyi differs from D. haraldschultzi by presenting a light brown smooth dorsum while D. haraldschultzi has a grayish cream dorsum with thin longitudinal stripes composed of small dots and with a series of glands, more densely distributed over the head ( Bokermann 1962). From D. battersbyi , D. ozzyi can be distinguished mainly by the smaller SVL—males of D. ozzyi range between 19.45–21.53 mm vs. 33 mm of the male holotype of D. battersbyi , and by different overall coloration; while D. battersbyi is gray and minutely dotted with reddish brown on all the upper and lower surfaces except the forelimbs and belly, D. ozzyi is light brown. Dendropsophus battersbyi presents a dark canthal and supratympanic streak, loreal region whitish with dark dots, limbs crossbarred and a pair of callous, white spots on the buttocks—features absent in D. ozzyi (see Rivero 1961). Dendropsophus tintinnabulum presents a concealed tympanum (visible in D. ozzyi ) and a white line dividing the dorsum from the flanks in preservative (VGDO observation on the syntypes). In life, D. tintinnabulum presents a grayish green dorsum and the venter is bluish green ( Melin 1941); D. ozzyi has a light brown dorsum and the venter is orange and white.

Description of holotype: Adult male, SVL 19.4 mm; head wider than long (HW/HL 1.18), widest below eyes; snout truncate in both dorsal and lateral views, eye-nostril distance longer than eye diameter ( END /ED 1.17); canthus rostralis indistinct, straight; loreal region slightly concave; lips thin; internarial area slightly depressed; nostrils protuberant, directed dorsolaterally; interorbital area flat; IOD/ED 1.48, IOD/HW 0.49; eyes large and protuberant (ED/HL 0.40, ED/HW 0.33); ELW 1.38 mm. Pupil horizontally elliptical; nictitating membrane transparent, its border slightly pigmented in the same pattern of the eyelid. Supratympanic fold barely visible, semi-circular in outline, less visible on the posterior portion; tympanum small (TD/HL 0.17), distinct, nearly round, separated from eye by a distance of 1.3 tympanum diameter; choanae small, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores very small, the right bearing two small, unaligned teeth while the left bears three; tongue cordiform, notched behind, posterior 1/3 not adhered to floor of mouth; vocal slits long, extending from midlateral base of tongue to almost the angle of jaws; vocal sac developed, single, and subgular, extending over the forearm (Figs. 3–4).

Arm slender, not hypertrophied; axillary membrane reaching half arm, delimited by the m. pectoralis portio axillaris (seen by transparency). Lateral border of arm with fimbria composed of small tubercles, not all clearly individualized. Fingers short, bearing discs; finger discs are slightly pointed, albeit disc of Finger II is slightly less, close to round; relative length of fingers II <III <V <IV; subarticular tubercles less wide than the digits, round on all fingers, bifid on penultimate articulation of Finger IV, most prominent on fingers II, III and IV; supernumerary tubercles present; inner metacarpal tubercle flat, elliptic; outer metacarpal tubercle bifid, nearly not visible; glandular nuptial pad covering the area corresponding to the prepollex in mesial view; webbing basal between fingers II and III; webbing formula II 2 -–2- III 2 -–2- IV 2 +–2+ V (Fig. 3C).

Hind limb long and slender (TL/SVL 0.53); no tarsal fold, but outer margins of tarsus and feet with fimbriae similar to the one found in arm; calcar tubercle absent, heel with small, inconspicuous tubercles; toes bearing pointed discs, smaller than those on fingers; feet slightly dehydrated, right one more than the left; relative lengths of toes I <II <III <V <IV; subarticular tubercles, round, bifid on penultimate articulation of Toe V, shallow; small supernumerary tubercles present on toes III and IV; inner metatarsal tubercle flat, elliptical; outer metatarsal tubercle indistinct, nearly not visible; webbing formula I 2–2 + II 1 +–3- III 1 +–2+ IV 2 +–1+ V. Toe IV bears a gland, visible at both dorsal and ventral views at the level of subarticular tubercles II and III; dorsally, a glandular patch is visible on Toe III ( Fig. 4 ).

Skin on dorsum, head, dorsal surfaces of forearms and thighs, flanks and groin smooth, finely shagreen on head, shank and tarsus; skin on belly and ventral surfaces of thighs granular. Cloacal opening directed posteriorly, above the thighs, covered by a small cloacal sheath dorsally; cloacal tubercles absent. Pectoral region, belly, ventral surfaces of forearm and thigh with many minute acini filled with dark yellow secretion. Pectoral acini not arranged as a glandular patch; loosely distributed over the area ( Fig. 4 )

Coloration in preservative: All dorsal surfaces yellowish brown, except thighs, fingers II, III, IV and toes I, II, III, IV that are cream. Dorsal ground color is darker at the head, fading posteriorly. Dorsum bears 13 brown smudges of irregular shape. Canthus rostralis, eyelid and the anterior third of the flanks have brown and white markings. Iris black with many minute silver spots of variable diameters. Ventral areas creamy white, except for the submandibular area that bears various brown flecks, palmar and plantar surfaces have a pattern resembling the dorsal pattern. The belly also presents a bright white rectangular area, roughly covering the area of the sternum. Dissections of paratype CFBH 35764 demonstrated that it is a layer of iridophores at the perimysium of the abdominal muscles.

Coloration in life (based on paratype MPEG 27810): Dorsal surfaces of head and body brown. Flanks, arms, hands (excluding fingers II to IV and finger disc of Finger V), tibia, and the external region of feet (including toes IV and V) light brown. Thighs, fingers II to IV and finger disc of Finger V, and internal region of feet (including toes I to III and toe discs of toes IV and V) vivid orange. Palmar, plantar and ventral surfaces of the axillary membrane, thigh and shank vivid orange. Ventrolateral areas of tarsus and feet dark brown. Belly creamy white with clear white spots and a rectangular bright white area (as described above). Gular region orangish cream with brown flecks. Iris copper with dark reticulations. Lateral areas of the head and flanks light brown, nearly white, composing a diffuse stripe starting at the loreal region and extending posteriorly to the groin through the trunk.

Variation. Measurements are shown in Table 1. External morphology of the type series is generally concordant with the holotype; including color ( Fig. 5–6 ). The number and conspicuousness of ventral acini varies from nearly none (MPEG 27809) to many (MPEG 27810, 27812 and MZUSP 154087 as also do the number of clear white flecks (while MPEG 27809 has none, MPEG 27810 has many) and the iridophore area of the abdominal perimysium. Dorsal coloration varies slightly, from lighter (MPEG 27814) to darker (MPEG 27809; CFBH 35764) shades of brown. The number, shape, and position of dorsal smudges also vary from absent (MPEG 27814; MZUSP 154086) to more then 20 (MPEG 27812). Left thigh muscles of MPEG 27809, MZUSP 154084–154088, and the left foot of MPEG 27810 were removed in the field as tissue samples for molecular analyses.

Advertisement call. Most of the recordings have more than one individual calling, and in many cases it is impossible to distinguish which one is emitting the note. Nonetheless, we have measured the note duration and spectral parameters for all the notes in the best recording (PLDR 035; N = 11 notes from up to three distinct calling males), and only the first three notes and the intervals between them in the second recording (PLDR 036; certainly emitted by the same individual). One of the individuals in recording PLDR 0 36 was collected (MPEG 27814, field number FPR 205) but the calls emitted by this male are unsuitable for bioacoustical analyses (saturated recording). Nonetheless, the call can be unambiguously identified as belonging to the same species as the ones for which analyses were performed.

Analyses of recordings reveal that the advertisement call of Dendropsophus ozzyi consists of an unusually high-pitched single note, emitted at relatively long intervals (Fig. 7). Amplitude modulation within the note shows higher energy at the beginning of the note, with energy decreasing abruptly and then continuing with small energy for the remaining of the note duration. Note duration is 165–386 ms (mean 248 ms; N = 14), interval between notes is 1965–5183 ms (N = 2), and the dominant frequency is 9130.1–10136.7 Hz (mean 9560.7 Hz; N = 14). Notes are not clearly multi-pulsed, except for a single note with four pulses.

Natural history. Dendropsophus ozzyi was found mainly on bushes close to water bodies, such as ponds and swamps, inside Terra Firme forest. The most commonly used type of vegetation were shrubs and small trees <5cm DBH (diameter at breast height), where individuals were observed on the leaves and branches in heights that varied from ground level (0 m) to 3.5 m, with prevalence between 1.0 and 1.5 m. No difference was observed in height of perch between sexes. Most individuals were observed 0.5 m away from water bodies. In addition, no individuals were observed farther than 5.0 m away from water bodies. Calling males were observed in the wettest months, from November to June. Time of observation varied overnight, from 18:00 to 23:00 and was more frequent Catalog Number Field Number Status SVL HL HW ED ELW END IOD TD THL TL FL MPEG 27809 FPR 105 Paratopotype 19.80 5.80 6.20 2.50 1.40 2.90 3.60 0.90 9.70 10.40 13.80 MPEG 27810 FPR 140 Paratopotype 20.00 6.50 7.20 2.60 1.40 2.50 3.60 1.40 10.80 11.10 14.10 MPEG 27811 FPR 141 Holotype 19.50 5.80 6.90 2.30 1.50 2.70 3.40 1.00 9.10 10.30 13.40 MPEG 27812 FPR 142 Paratopotype 18.50 5.70 6.50 2.30 1.50 2.90 3.60 1.10 9.60 10.10 13.30 MPEG 27813 FPR 143 Paratopotype 21.50 5.70 7.10 2.90 1.50 2.90 3.60 1.40 9.70 9.70 13.50 MPEG 27814 FPR 205 Paratopotype 19.80 6.10 6.80 2.30 1.50 2.30 3.30 0.80 10.20 9.80 13.50 INPA-H 32742 J123 Paratype 18.30 6.60 6.90 3.00 1.90 2.05 2.50 1.10 10.20 10.50 13.20 MZUSP 154088 J117 Paratype 20.10 7.40 8.00 2.90 1.90 2.00 2.60 1.00 10.20 10.80 13.00 MNRJ 86925 J264 Paratype 18.10 6.80 6.50 2.90 1.50 1.90 2.50 0.95 10.20 10.20 13.20 INPA-H 32744 J121 Paratype 19.00 6.60 7.00 2.80 2.10 2.10 2.50 0.75 10.00 10.40 13.40 CFBH 35764 J205 Paratype 19.40 7.10 7.30 2.90 1.70 1.90 2.50 0.85 10.25 10.50 13.50 CFBH 35762 J129 Paratype 18.60 7.20 7.30 2.85 2.40 2.00 2.10 0.90 10.40 10.90 13.90 MNRJ 86921 J206 Paratype 20.40 7.40 7.40 3.10 1.90 1.85 2.20 1.00 10.50 10.45 13.90 CFBH 35760 J125 Paratype 18.30 7.40 7.30 3.00 1.85 1.80 2.50 0.80 10.30 10.25 13.50 MNRJ 86923 J208 Paratype 20.50 7.10 7.30 3.00 1.90 1.90 2.60 1.00 10.85 10.90 14.10 MNRJ 86922 J207 Paratype 19.20 7.00 7.10 2.65 1.60 1.90 2.20 1.20 9.70 10.40 14.10 MZUSP 154087 J116 Paratype 20.65 7.10 7.60 3.00 2.30 1.85 2.80 1.05 11.00 10.80 14.15 CFBH 35761 J126 Paratype 20.90 7.40 7.20 2.90 2.30 1.90 2.55 0.95 11.70 11.50 13.70 INPA-H 32746 J18 Paratype 19.90 6.90 7.60 2.90 2.20 2.00 2.70 0.90 11.30 11.30 14.65 MNRJ 86924 J263 Paratype 19.50 7.50 7.10 3.00 1.60 1.90 2.30 1.05 10.00 10.45 13.80 MZUSP 154086 J115 Paratype 19.80 6.80 7.70 2.85 1.90 1.80 2.40 0.90 9.90 10.40 13.35 MZUSP 154085 J114 Paratype 19.40 7.10 7.40 2.70 1.70 2.10 2.20 0.95 10.65 10.60 12.70 INPA-H 32747 J19 Paratype 19.20 6.70 6.95 3.05 2.30 1.90 2.50 0.90 10.00 10.20 13.70 INPA-H 32748 J124 Paratype 18.00 6.45 6.85 2.65 1.80 1.75 2.30 0.90 10.15 10.00 12.90 INPA-H 32743 J20 Paratype 19.90 7.20 7.50 2.90 2.10 1.90 2.85 0.90 10.40 10.40 14.35 MZUSP 154084 J113 Paratype 19.30 7.00 7.25 2.90 1.85 2.05 2.60 0.95 10.85 11.05 13.00 INPA-H 32745 J122 Paratype 17.90 6.80 6.85 2.50 1.75 1.85 2.25 1.00 10.40 10.15 13.60 MPEG 22352 JUR246 Paratype 19.70 5.80 6.20 2.50 1.39 2.90 3.50 0.80 9.80 10.50 13.60 MPEG 22353 JUR247 Paratype 19.60 5.80 6.10 2.60 1.41 2.50 3.50 0.90 9.60 10.30 13.40 MPEG 22354 JUR248 Paratype 21.30 6.60 7.10 2.80 1.50 2.90 3.60 1.40 10.40 11.10 14.10 MPEG 22355 JUR249 Paratype 18.60 5.70 6.50 2.30 1.42 2.30 3.30 0.90 9.60 10.10 13.30 MPEG 22400 JUR221 Paratype 19.80 5.70 6.20 2.40 1.43 2.30 3.70 1.10 9.70 9.70 13.50 MPEG 22401 JUR222 Paratype 21.40 6.70 7.20 2.90 1.51 2.90 3.40 1.40 10.50 11.20 14.30 MPEG 22402 JUR223 Paratype 19.90 5.70 6.20 2.50 1.42 2.40 3.30 1.10 9.60 9.90 13.10 MPEG 22403 JUR224 Paratype 18.70 5.60 6.10 2.30 1.47 2.30 3.50 0.80 9.50 9.70 13.20 MPEG 22404 JUR225 Paratype 18.60 5.60 6.10 2.40 1.45 2.50 3.30 1.20 9.50 9.80 13.20 CFBH 35762 J130 Paratype 18.75 7.20 7.45 2.95 2.20 2.00 2.45 0.95 10.70 10.90 13.80

29

Species clade Calltype Minimum Maximum Average Reference

= Abrunhosa et al. (2001) report different values in their text and table. We reproduce the largest range reported.

**= Marquez (1993) reports dominant frequencies around 120 Hz, a curiously low value. Authors also report additional emphasized frequencies of higher values that are a more

robust test to our hypothesis that D. ozzyi sp. nov. has an unusually high dominant frequency. See Orrico et al. (2009) for a discussion on dominant frequency artifacts.

***= Bastos & Pombal (2003) do not report which frequency is dominant, just that there are emphasized harmonics around 3 and 8 kHz. Their figure 11 depicts a call with dominant

frequency around 7 kHz.

**** Lescure & Marty (2000) and Tárano (2010) report different values (~5 kHz and 3.8 kHz respectively).

between 19:00 and 21:00. Amplectant pairs and tadpoles were not observed. All collection sites are between the Tapajós and Madeira Rivers, and we expect more populations of D. ozzyi to be found within this area.

Etymology. The specific epithet is used as a noun in the genitive case and honors John Michael "Ozzy" Osbourne, a famous British rock singer, former vocalist of the rock band Black Sabbath, for its contribution to modern music and culture. Ozzy is commonly associated with bats because of the famous episode in which, on stage while he was performing a gig, he bit off a bat’s head thrown by a fan. He later stated that he thought it was a plastic toy.

When calling, this new Dendropsophus species can be vaguely associated with the high pitch sounds emitted by some bat species that are hearable to the human ear. When we heard this species in the field we immediately pictured a calling bat. Because of this “bat association” we take the opportunity to name this species after one of the biggest music legends of all time.

Figures

FIGURE 4. Dendropsophus ozzyi sp. nov. holotype (MPEG 27811). (A) Dorsal and (B) ventral views of the body. SVL = 19.6 mm. Arrowheads point to the glandular patch of Toe IV.

FIGURE 5. Dendropsophus ozzyi sp nov. Live male specimens: (A) MZUSP 154086, SVL = 19.8; (B) MZUSP 154086 (day coloration), SVL = 19.8; (C) MZUSP 154084, SVL = 19.3; (D) CFBH 35763, SVL = 18.8; (E) MNRJ 86923, SVL = 20.5; (F) MPEG 27810, SVL = 19.8 mm. (A) – (E) from Igarapé Mutum, Juruti, State of Pará, Brazil; (F) from Comunidade Bragança, Rio Paraconi, Floresta Nacional de Pau-Rosa (FNPR), Maués, State of Amazonas, Brazil (paratopotype).

FIGURE 6. Dorsal (a) and ventral (b) color patterns of 21 preserved paratypes of Dendropsophus ozzyi sp. nov. from Juruti, Pará, Brazil. Scale bar = 20.0 mm. (1) INPA-H 32746; (2) INPA-H 32747; (3) INPA-H 32743; (4) MZUSP 154084; (5) MZUSP 154085; (6) MZUSP 154086; (7) MZUSP 154087; (8) MZUSP 154088; (9) INPA-H 32744; (10) INPA-H 32745; (11) INPA-H 32742; (12) INPA-H 32748; (13) CFBH 35760; (14) MNRJ 86921; (15) MNRJ 86925; (16) CFBH 35763; (17) CFBH 35764; (18) CFBH 357602; (19) MNRJ 86922; (20) MNRJ 86923; (21) MNRJ 86924. For additional information see Table 1 and text.

Tables

TABLE 2. Advertisement call emission type and dominant frequency for species assigned to the Dendropsophus microcephalus Group. Dominant Frequency (kHz)

D. berthalutzae D. bipunctatus *	decipiens	monophasic 3.7 monophasic 4.8	4.7 5.6	3.8?	Forti et al. (2012); Moura et al. (2012) Abrunhosa et al. (2001)
D. branneri		monophasic 5.9	6.6	6.3	Nunes et al. (2007)
D. coffeus		diphasic 5.7	6	5.8	Köhler et al. (2005)
D. cruzi		monophasic 5.2	7	6.3	Bastos et al. (2003)
D. decipiens	decipiens	monophasic 4.7	5.2	5	Abrunhosa et al. (2001)