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Adenomera phonotriccus, n. sp. Figures 1‒3 REFERRED MATERIAL: Voucher specimens with DNA sequenced by Fouquet et al. (2014) and assigned to Adenomera sp. F (in part). SUGGESTED VERNACULAR NAME: Tody-tyrant-voiced nest-building frog. HOLOTYPE: MPEG 41155 (field number TRC 135), adult male, from the municipality of Palestina do Pará (5.70228° S, 48.24949° W; 166 m), on the western margin of the lower Araguaia River, state of Pará, northern Brazil, collected by P.L. V. Peloso, M.J. Sturaro, P. V. Cerqueira, G. Gonçalves, A.A. Giaretta, P. Marinho, and T. R. de Carvalho on January 9, 2018 (Zoobank LSID registration: urn:lsid:zoobank.org:act: 330AE4AB-1ABB-4B90-9245-1DBC2A1F661B). SOUND RECORDING: Adenomera _phonotriccusPalestinaPA2bAAGm661 MK 2. PARATOPOTYPES: MPEG 41156 and CFBH 43130‒43131, all adult males, collected (5.70300° S, 48.24833° W; 176 m) on January 9‒10, 2018, by the same collectors. ETYMOLOGY: The epithet phonotriccus is the combination of Greek phono - (from phoné, “sound, voice”) and triccus (a small bird whose species is not identifiable, though in modern times the name is applied to tyrant flycatchers; Jobling, 2010). This name is to be treated as a noun in apposition and is an allusion to the similarity between the vocalization of the new species and those of tody-tyrants. Tody-tyrants of the Neotropical genus Hemitriccus have peculiar vocalizations, which are reminiscent of the vocalization of Adenomera phonotriccus, especially the trilled song of H. cohnhafli (see Zimmer et al., 2013). Interestingly, several Hemitriccus species are morphologically very similar, and consequently best differentiated by their vocalizations, a pattern also observed in Adenomera. DIAGNOSIS: Adenomera phonotriccus is differentiated from its congeners by the following combination of characters: (1) small size (adult male SVL 19.8–21.6 mm; table 1); (2) robust body shape; (3) toe tips unexpanded to slightly swollen (character states B, C; Heyer, 1973); (4) distal antebrachial tubercle on underside of forearm; (5) throat and belly cream colored, mottled white and gray in some parts; (6) two color morphotypes (presence/absence of dorsolateral stripes); (7) advertisement call consisting of a single type of pulsed note, emitted regularly, not in calling bouts; (8) advertisement notes composed of complete pulses (pulses with periods of silence in between); (9) long-lasting call duration (213‒433 ms). COMPARISONS: Adenomera phonotriccus has adult males (SVL 19.8–21.6 mm; table 1) smaller than those of A. coca (23.6–25.6 mm; Angulo and Reichle, 2008), A. lutzi (25.7–33.5 mm; Kok et al., 2007), and A. simonstuarti (SVL 25.9–26.2 mm; Angulo and Icochea, 2010). Adenomera phonotriccus has a robust body shape (figs. 1–2), whereas A. diptyx, A. martinezi, and A. saci have a slender body (Carvalho and Giaretta, 2013a). Adenomera phonotriccus has toe tips unexpanded or slightly swollen (character states B, C), but not expanded into flattened discs (character state D) as in A. ajurauna, A. andreae, A. marmorata, A. lutzi, A. nana, and A. simonstuarti (Angulo et al., 2003; Kok et al., 2007; Kwet, 2007; Berneck et al., 2008; Angulo and Icochea, 2010; appendix 1). Adenomera phonotriccus is distinguished from congeners (except A. cotuba and A. lutzi; Kok et al., 2007; Carvalho and Giaretta, 2013b) by having antebrachial tubercles on underside of forearm (fig. 3), presumably absent in all remaining species. Adenomera phonotriccus differs from A. heyeri and A. lutzi by having ventral surfaces cream colored (mottled white and gray in some parts), yellow in the latter two species (Boistel et al., 2006; Kok et al., 2007). From A. engelsi, which does not have distinctive dorsolateral stripes and pigmentation on throat (Kwet et al., 2009), Adenomera phonotriccus differs by the occurrence of both color features (fig. 1). Adenomera cotuba does not have dorsolateral stripes (Carvalho and Giaretta, 2013b), whereas A. phonotriccus has two color morphotypes (presence/ absence of dorsolateral stripes). These species are better distinguished by their distinct vocalizations (see below; fig. 4). The advertisement call of Adenomera phonotriccu s (fig. 4; table 2) distinguishes it from all congeners by having pulsed notes composed of complete pulses (periods of silence in between), whereas the other species’ notes are formed by incomplete pulses, i.e., not separated one from the next by periods of silence (fig. 4). From the remaining species of Adenomera, the new spe- cies differs by having pulsed advertisement notes, whereas those species’ calls consist of nonpulsed notes (table 3). The advertisement call of A. phonotriccus is distinguished from that of A. cotuba by being composed of single notes emitted regularly (table 2), whereas A. cotuba’ s call is made up of many notes emitted as calling bouts, i.e., multinote advertisement call (Carvalho and Giaretta, 2013b: fig. 4A). In addition, to date, Adenomera phonotriccus has the longest note duration in the genus, whose range does not overlap with note-duration values for any congeners (table 3). DESCRIPTION OF HOLOTYPE: MPEG 41155 (figs. 1A, 2, 3A). Adult male. Body robust (figs. 1A, 2A). Snout subovoid in dorsal view (fig. 2A), acuminate in lateral view (fig. 2D). Nostrils closer to the snout tip than to the eyes; fleshy ridge on snout tip; canthus rostralis not marked; loreal region slightly concave; supratympanic fold from the posterior corner of the eye to the base of the arm; oval postcommissural gland; vocal sac subgular with a fold from jaw extending to forearm on each side, vocal slits present; vomerine teeth in two straight rows medial and posterior to choanae and oblique to sagittal plane. Tongue elongated, free behind. Relative finger lengths IV a Sinusoidal oscillation (Boistel et al., 2006) corresponds to incomplete pulses in Adenomera, based on (quasi-) periodic sinusoidal FMs (sometimes there is strong modulation unaccompanied by distinct pulses though). b Based on the call of Adenomera sp. 2 from Joinville. c See Materials and Methods for a detailed explanation. d Frequencies lower than 1 kH do not pertain to this species’ call (appendix 2). COLOR OF HOLOTYPE IN PRESERVATIVE (fig. 2): Snout tip with a faded white coloration (coincident with the fleshy ridge). Dorsum marble gray with a few darker specks and spots. Sacral stripe on the posterior third of body length, cream-colored. Black lumbar glands. Dorsal surface of limbs with dark brown stripes/blotches on a slightly lighter brown background. Dorsolateral, broad stripe from the posterior corner of the eye, passing over the flanks, predominantly cream-colored to a somewhat orange hue (faded coloration; orange in life). Upper and lower jaws covered with white-colored spots/blotches, whitish postcommissural gland, outlined at the bottom by a dark coloration. Tympanum reddish brown. Throat, belly, and ventral surface of limbs cream, with melanophores, throat with a black-dotted pattern, solid dark-colored laterally, coinciding with the expanded vocal sac. Posterior surface of thighs with a few dark brown spots on a light brown background, and paracloacal glands cream colored. COLOR OF HOLOTYPE IN LIFE (fig. 1A): Dorsum covered with black specks and spots irregularly distributed on medium brown background. Arms and legs reddish brown. Sacral and dorsolateral stripe, and postcommissural gland orange. Iris copper. Tympanum brown. Ventral surfaces cream colored and with mottling on throat and belly (laterally). Groin yellow. VARIATION IN THE TYPE SERIES: Despite minor variation in dorsal coloration and distribution of spots, specks, and warts on dorsal surfaces, characteristics of the three paratypes largely agree with the description of the holotype. Dorsolateral stripe is absent in two paratypes (MPEG 41156 and CFBH 43130). Shape and size of antebrachial tubercle varies among the type series; some of them are very discrete (fig. 2). ADVERTISEMENT CALL: The description is based on recordings of three males (appendix 3). Sample sizes for each acoustic trait and descriptive statistics, i.e., means and standard deviations, are provided in table 2. The call (fig. 4) consists of a single type of pulsed note, which is emitted at a rate of 26–35 calls per minute. Note duration varies from 213–433 msec. Notes have a smooth attack and decay, reaching the amplitude peak at 43%–85% of their duration. Pulse number varies from 14–26; pulse duration from 4–26 msec, emitted at a rate of 51–67 pulses per second; silence between pulses varies from nearly zero to 13 msec. The fundamental frequency of the note occupies a narrow bandwidth harmonic, with a peak frequency of 1857– 1997 Hz; the dominant frequency varies from 3639–4113 Hz and is contained within the second harmonic. Notes usually have a slight positive frequency modulation along their duration, but a few notes do not have any modulation, and a single note has negative modulation. Linear frequency modulation varies from -43–861 Hz. HABITAT AND NATURAL HISTORY: Adenomera phonotriccus is associated with forest habitats in Brazil’s eastern Amazonia in the state of Pará. The type locality is located on the west margin of the lower Araguaia River. This region is also occupied by savanna landscapes or ecotones between the Cerrado and Amazonian rainforest. The species was heard along an open area transect (by recent logging activity), especially at the forest edge, and inside the forest remnant. Males called while exposed or under leaf litter, and increased their calling activity during and shortly after rainfalls. The vocalization of Adenomera phonotriccus is very peculiar when heard in the field. Our first impression was that the calls might not be produced by a leptodactylid frog, or even an anuran species. Indeed, the vocalization is similar to that of some Neotropical suboscine birds (e.g., Rhynchocyclidae, Tyrannidae). There were two partially syntopic species of Adenomera also in calling activity at the study site: Adenomera sp. (A. heyeri clade) and A. aff. hylaedactyla. We heard the three species calling at the same period (late afternoon), even though A. phonotriccus and Adenomera sp. ceased most calling activity at dusk, whereas A. aff. hylaedactyla continued calling through the first hours of the night. The last species occupied open areas, e.g. pasture. In contrast, A. phonotriccus and Adenomera sp., are associated with forest habitats. DISTRIBUTION (fig. 5): Adenomera phonotriccus is known with certainty only from the type locality and Marabá. However, specimens from other regions in the state of Pará were also assigned to this lineage, referred as Adenomera sp. F, by Fouquet et al. (2014). Given the high levels of cryptic species diversity in Adenomera and complex genetic structure within the lineages identified by Fouquet et al. (2014), we conservatively restrict the species’ distribution to the type locality region until additional data, especially vocalizations, are obtained for the other populations assigned to Adenomera sp. F, a lineage widely distributed in central-east Pará, northern Brazil. CONSERVATION STATUS: Because the species is apparently restricted to forested habitats, or its immediate margins (forest edge), we infer that the current population trend for this species is “declining.” Major threats are deforestation (destruction of habitats) due to conversion of large areas where the species occur into pasture and agricultural land. The species appears to be locally abundant (although difficult to observe), which means that as long as its habitat is preserved the species should have its future secured. The known range of the species is hard to determine with current evidence. It may be much wider, if all the clade F specimens refer to A. phonotriccus, in which case the species would be found from the western margin of the Tocantins River to the western margins of the Tapajós River (Fouquet et al., 2014: fig. 2). Alternatively, it could have a much narrower distribution (southern Pará) if only specimens in subclade F3 correspond to A. phonotriccus —in the latter case, the species would be probably restricted to small forest fragments in one of the most highly threatened portions of the Amazonia (the so-called Arc of Deforestation). This is a typical example of the issues raised by Peloso (2010) and Angulo and Icochea (2010), where lack of taxonomic resolution hinders a proper assessment of threats to cryptic species. For now, we must unfortunately suggest this new species is yet another case of data-deficient taxa—a lot more studies and data are needed to define the conservation status of A. phonotriccus. REMARKS: Calls of the voucher PV 2597 (specimen no. DT 2123; recordings PLPDR 100; see appendix 3) were unambiguously assigned to A. phonotriccus (“F3” in fig. 6). This voucher was cited as collected in Araguatins (state of Tocantins), on the east margin of Araguaia River. However, this specimen was actually collected in Palestina do Pará, state of Pará—the animal was collected by P.L.V.P. in 2010 (coordinates given by Fouquet et al.’s appendix 1 are correct). The basecamp for this collection site was in Araguatins, and this may have been the source of confusion with the sample provenance (P.L.V.P., personal obs.). Another voucher specimen used in the phylogeny (PV 2412, specimen no. DT 2016), also assigned here to A. phonotriccus (“F3” in fig. 6) was collected in Marabá—this municipality is located in the state of Pará, not Amazonas (AM), as mentioned in appendix 1 of Fouquet et al. (2014). Of special interest is the acquisition of additional information for the subclade F1, which occurs in partial sympatry with A. phonotriccus (fig. 6). Meanwhile, we conservatively assign only subclade F3 (sensu Fouquet et al., 2014) to A. phonotriccus, and this species is an endemic of the central-east part of the state of Pará in association with the west margin of the lower Araguaia River (fig. 6). With respect to subclade F2, we suspect that it may not be conspecific with A. phonotriccus, given its distribution range associated with another region in the state of Pará (Xingu-Tapajós interfluve; fig. 6). Therefore, we maintain the original status of unconfirmed candidate species of both subclades F1 and F2 for the moment.

De Carvalho, Thiago R., Giaretta, Ariovaldo A., Angulo, Ariadne, Haddad, Célio F.B., Peloso, Pedro L.V. (2019): A new Amazonian species of Adenomera (Anura: Leptodactylidae) from the Brazilian state of Pará: a tody-tyrant voice in a frog. American Museum Novitates 2019 (3919): 1-24, DOI: https://doi.org/10.1206/3919.1

Governments have committed to conserving ?17% of terrestrial and ?10% of marine environments globally, especially “areas of particular importance for biodiversity” through “ecologically representative” Protected Area (PA) systems or other “area-based conservation measures”, while individual countries have committed to conserve 3–50% of their land area. We estimate that PAs currently cover 14.6% of terrestrial and 2.8% of marine extent, but 59–68% of ecoregions, 77–78% of important sites for biodiversity, and 57% of 25,380 species have inadequate coverage. The existing 19.7 million km2 terrestrial PA network needs only 3.3 million km2 to be added to achieve 17% terrestrial coverage. However, it would require nearly doubling to achieve, cost-efficiently, coverage targets for all countries, ecoregions, important sites, and species. Poorer countries have the largest relative shortfalls. Such extensive and rapid expansion of formal PAs is unlikely to be achievable. Greater focus is therefore needed on alternative approaches, including community- and privately managed sites and other effective area-based conservation measures.

LEPTODACTYLUS COCA SP. NOV. (FIGS 1, 2) Holotype: Adult male, NKA 3630, field number SR 99.134, collected near Carretera Antigua from Villa Tunari to Cochabamba (c. 800 m above sea level), just above the Village of El Palmar, Province of Chapare, Department of Cochabamba, Bolivia, on September 25, 1997 by S. Reichle and J. Aparicio. The holotype is also a call voucher for the species, recorded by S. Reichle. Paratypes: Two additional specimens were collected at the type locality: a gravid female (CBF 4298), and a male, which is also a call voucher (CBF 2619), recorded by S. Reichle and collected by S. Reichle and J. Aparicio on September 25, 1997. DIAGNOSIS Leptodactylus coca differs in its advertisement call from all other known members of the L. marmoratus group for which acoustic data are available. Leptodactylus araucaria (Kwet & Angulo, 2002) and L. nanus can be distinguished from L. coca by their small size [maximum SVL in L. araucaria males, 18.8 mm (Kwet & Angulo, 2002); maximum SVL in L. nanus males, 19.4 mm (Kwet, 2007)], whereas Leptodactylus lutzi (Heyer, 1975) differs from the new species by its large size [SVL in males, 25.7���33.5 mm (Kok et al., 2007)], the presence of a dark triangular seat patch, and distinct spotting on the posterior face of the thigh (Heyer, 1975). Individuals of Leptodactylus bokermanni Heyer, 1973 have not been found to possess a light mid-dorsal stripe extending from above the vent to the middle of the body, whereas this is a possible pattern for individuals of L. coca. The new species differs from Leptodactylus martinezi Bokermann, 1956 in lacking four longitudinal series of dark glands on the dorsum (Bokermann, 1956). L. andreae, L. diptyx, L. hylaedactylus, Leptodactylus marmoratus Steindachner, 1867, and Leptodactylus thomei Almeida & Angulo, 2006 all have distinct, different advertisement calls to those of L. coca. There is a recently described species of Leptodactylus of the marmoratus group from French Guiana, Leptodactylus heyeri (Boistel, de Massary & Angulo, 2006), which has a similar call to that of L. coca. However, even with similar calls, there are differences in acoustic patterns. For example, even if the call rate is often influenced by the motivation of callers, the call of L. heyeri is issued at a much lower call rate than that of L. coca, and the call of L. heyeri does not show evidence of pulse structure, whereas it is present in L. coca. The species also differ morphologically: L. heyeri has a relatively broader head in proportion to the body; the dorsal coloration is very distinct, with stripes and bands, and well-defined black lumbar glands; and males of this species have a yellow throat and belly. DESCRIPTION OF HOLOTYPE Body small, robust, with relatively short limbs. Dorsal outline of snout rounded (Fig. 1), in profile snout nearly acuminate; head wider than long; nostrils positioned dorsolaterally, closer to tip of snout than to corner of eye; internarial distance about a quarter of head width. Tympanum distinct, slightly more than half the diameter of eye; tympanic membrane translucid, possible to observe tympanic canal; supratympanic fold weakly developed, extending from back of eye to arm insertion, dark, contouring outline above fold extending from back to eye to about half or two thirds of the way down to the arm; oval creamcoloured gland below angle of jaw, and supratympanic fold present and distinct; canthus rostralis indistinct. Vocal sac single and internal, paired elongate vocal slits present. Vomerine teeth posterior to choanae in transverse series parallel to choanae, separated from each other by approximately the length of one vomerine tooth row. Arms short and robust; fingers slender, finger lengths III > I = II > IV; finger tips rounded, without fringes or expansions; palms of hand with two large ovoid-shaped cream-coloured metacarpal tubercles; smaller inner metacarpal tubercle a little less than half the size of larger metacarpal tubercle; fingers with conspicuous, distinct cream-coloured rounded subarticular tubercles, absence of nuptial asperities. Hindlimbs robust, shank slightly longer than thigh; toe lengths IV > III > V > II > I; toe tips very slightly flattened or not, with very minor expansions (character state B of Heyer, 1973); toes without fringes. Metatarsal tubercles conspicuous, distinct; inner tarsal fold weakly developed, lined with scant, small white tubercles; tarsus also with small white tubercles, although not profuse; sole of foot with very sparse, scant white tubercles. Texture on dorsal surface mostly smooth, becoming tuberculate around the sacral area; thighs mostly smooth, shanks with very small, mostly scant, tubercles. Dorsolaterally, a dark longitudinal glandular fold starting just behind shoulder and running discontinuously posterior to sacral region; beneath it a second cream coloured glandular lateral fold extending along the same length as the darker fold. Surface of venter smooth. * Holotype. COLOUR IN PRESERVATIVE Preserved in 70% alcohol, the dorsum is a light brown colour, with the discontinuous upper dorsolateral glandular lines a darker shade of brown. There is a dark marking in the shape of an inverted U on the dorsum between the shoulders. Tips of glands on lower back dark brown; dorsal surface of limbs, a uniform light brown, except for the white-tipped tubercles and a dark bar on left foot. Tip of snout to back of head at about tympanum level a darker shade of brown, with a slight milky/foggy aspect to it. Venter white, immaculate; skin on edges of lower lip sparsely dotted with melanophores. COLOUR IN LIFE The coloration in life does not differ much from the colour described in preservative, just more intense. MEASUREMENTS OF HOLOTYPE (IN MM) SVL, 24.1; HL, 8.0; HW, 8.7; ED, 2.6; TYD, 1.4; END, 1.6; IOD, 2.4; IND, 2; FAL, 4.3; HDL, 5; THL, 9.7; SL, 10.3; TSL, 6.5; FL, 11.5. MORPHOLOGICAL VARIATION Female slightly larger than males (Table 1). The dorsal pattern of the male specimen CBF 2619 is similar to that of the holotype, except that it lacks the fogginess observed in the snout and head of the holotype; it does not have the inverted U-shaped dark marking between the shoulders; the two dark brown dorsolateral discontinuous glandular lines are ventrally flanked by broad cream lateral stripes that run parallel to them. In addition, it appears to have an inverted triangular-shaped interorbital spot, which, although fainter on the left side of the specimen, is dark brown from its base between the eyes to the back of the head, with a lighter spot within the triangle close to the right eye. Furthermore, specimen CBF 2619 has two symmetrical inguinal dark blotches on either side of the lower dorsum, and another two symmetrical dark blotches lower down, flanking the base of the hairline cream stripe that extends from the base of the vent to the lower back, and very sparse dark spots on the upper dorsum. The supratympanic fold is more pronounced from just behind the eye to behind the tympanum; it is weaker from the back of tympanum to the arm. An accompanying dark brown bar follows the outline of the supratympanic fold only to the back of the tympanum. In contrast with the holotype, this specimen shows distinguishable irregular crossbars on the hindlimbs. The toe tips are expanded (character states C-D or D sensu Heyer, 1973) and flattened. * Holotype. Female specimen CBF 4298 is morphologically the most different of the three specimens collected. It has a cream-coloured patch on the dorsum between shoulders; an inverted triangular-shaped interorbital spot that is dark brown from its base between the eyes to the level of the posterior edge of the tympanum, with the exception of a couple of lighter-coloured irregular spots within the triangle that take the appearance of eyes on a mask; it then takes a lighter shade from the apex of this spot posteriorly until it reaches the anterior edge of the shoulder-level cream patch. The ���mask��� is flanked by two lighter coloured trapezoidalshaped spots, one on either side of the head, behind the eyes. The supratympanic fold is well developed from the back of the eye to the arm, with a dark brown bar following the contour of the fold nearly to the arm. There is a cream-coloured mid-dorsal hairline stripe running from above the vent to the creamcoloured shoulder patch, becoming progressively thinner anteriorly, and two lateral thin creamcoloured stripes run from the back of the tympanum to the groin, one on either side of the body. Two symmetrical inguinal dark blotches are located on either side of the lower dorsum. The remainder of the dorsum is marked with an irregular pattern of darker spots against a lighter brown background, which roughly resembles broken discontinuous lines running along the anterior���posterior axis. Irregular crossbars are present on both arms and hindlimbs. The toe tips are slightly expanded (character state C sensu Heyer, 1973) and mildly flattened or not flattened. In contrast to the holotype, both paratypes have profuse white-tipped tubercles on the soles of the feet. All specimens have rounded snouts in dorsal outline, although in profile the snout of the female appears to be more rounded than that of the males; all specimens have white-tipped tubercles on the dorsal surfaces of shanks and thighs (although these are more profuse on the shanks than they are on the thighs), and they all have immaculate creamish-coloured venters. ADVERTISEMENT CALL Acoustic parameter measurements for the two recorded voucher males are listed in Table 2. Figure 3 depicts both temporal and spectral patterns of the call. The advertisement call of L. coca has been previously described in Angulo (2004) as Adenomera ���Chapare���; it is here directly associated to the species name L. coca. The description is based on the combined calls from both vouchers, which were recorded at 21.1 ��C and 20.8 ��C, respectively. The advertisement call of the new species can be heard as a sound file on track 6, CD 2 of M��rquez et al. (2002). In comparison with other species of the L. marmoratus group (see Kwet & Angulo, 2002; Angulo et al., 2003; and references within; Kok et al., 2007; and this study, for acoustic parameters for other species) the advertisement call of L. coca is intermediate in duration, ranging from 110 to 145 ms, emitted at an intermediate to low call rate (66 and 84 calls/min). Call rise time is highly variable (11.6���96 ms), a fact reflected by associated standard deviations for both call vouchers. The signal undergoes quite strong amplitude modulation for most of its duration, before going into its final decay phase. This amplitude modulation creates some pulse structure, but because these pulses tend to meld together, it is difficult to specify an amplitude pattern. In one of the vouchers these modulations vary between 10 and 15 per call, emitted at an average rate of 120 pulses/s. The fundamental frequency ranges between 1695 and 1914 Hz, and a harmonically related second frequency, ranging between 3449 and 3748 Hz, is the main carrier. Up to five or six frequency bands can be detected at any time (this is also a function of recording conditions); nevertheless, given the fact that the energy peaks of upper harmonics are considerably lower than that of the main carrier, these frequency peaks are not depicted in the spectrum image of Figure 3, as they fall below the cut-off point of -30 dB. There is either no or some frequency modulation of the main carrier (0���775 Hz). NATURAL HISTORY The new species was found on open ground and at the base of dense, grassy vegetation in montane rainforests at about 800 m above sea level in the cocagrowing region of Chapare, Bolivia. Voucher males were recorded in the evening after a heavy downpour during the day; individuals vocalized from dense clusters of grass as well as from open areas. Many individuals were heard calling, and some of them were very close (within 5 m) to each other. No water bodies were found close by, and we suspect that the species could display direct development (e.g. as has been reported for L. marmoratus; Lutz, 1931, 1947). ETYMOLOGY The new species is named after the coca plant, Erythroxylon coca, given its occurrence near coca plantations of the Chapare region, in Bolivia. CONSERVATION STATUS The three type specimens are the only known representatives of the new species. They were all collected on the same day, after heavy rain. Given the available information, we suggest that this species be considered as Data Deficient following the IUCN���s Red List categories (IUCN, 2001)., Published as part of Angulo, Ariadne & Reichle, Steffen, 2008, Acoustic signals, species diagnosis, and species concepts: the case of a new cryptic species of Leptodactylus (Amphibia, Anura, Leptodactylidae) from the Chapare region, Bolivia, pp. 59-77 in Zoological Journal of the Linnean Society 152 (3) on pages 62-67, DOI: 10.1111/j.1096-3642.2008.00447.x, http://zenodo.org/record/4687663, {"references":["Kwet A, Angulo A. 2002. A new species of Adenomera (Anura, Leptodactylidae) from the Araucaria forest of Rio Grande do Sul (Brazil), with comments on the systematic status of southern populations of the genus. Alytes 20: 28 - 43.","Kwet A. 2007. Bioacoustic variation in the genus Adenomera in southern Brazil, with revalidation of Leptodactylus Nanus Muller, 1922 (Anura: Leptodactylidae). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe Supplement 83: 56 - 68. doi: 10.1002 / mmnz. 200600027","Heyer WR. 1975. Adenomera lutzi (Amphibia: Leptodactylidae), a new species of frog from Guyana. Proceedings of the Biological Society of Washington 88: 315 - 318.","Kok PJR, Kokubum MNC, MacCulloch RD, Lathrop A. 2007. Morphological variation in Leptodactylus lutzi (Anura, Leptodactylidae) with description of its advertisement call and notes on its courtship behavior. Phyllomedusa 6: 45 - 60.","Heyer WR. 1973. Systematics of the marmoratus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Contributions in Science, Natural History Museum, Los Angeles County 251: 1 - 50.","Bokermann WCA. 1956. Sobre una nueva especie de Leptodactylus del Brasil (Amphibia). Neotropica 2: 37 - 40.","Almeida AP, Angulo A. 2006. A new species of Leptodactylus (Anura: Leptodactylidae) from the state of Espirito Santo, Brazil, with remarks on the systematics of associated populations. Zootaxa 1334: 1 - 25.","Boistel R, Massary J-C (de), Angulo A. 2006. Description of a new species of the genus Adenomera (Amphibia, Anura, Leptodactylidae) from French Guiana. Acta Herpetologica 1: 1 - 14.","Angulo A. 2004. The evolution of the acoustic communication system in members of the genus Adenomera (Anura: Leptodactylidae): a comparative approach. Unpublished PhD Thesis, University of Toronto.","Marquez R, De la Riva I, Bosch J, Matheu E. 2002. Sounds of frogs and toads of Bolivia. Barcelona: Alosa, sonidos de la naturaleza.","Lutz A. 1931. Sur la biologie des batraciens du Bresil. Comptes Rendus de la Societe de Biologie, Paris 109: 755 - 756.","Lutz B. 1947. Trends towards non-aquatic and direct development in frogs. Copeia 1947: 242 - 252.","IUCN. 2001. IUCN red list categories and criteria, Version 3.1. IUCN Species Survival Commission. Gland: IUCN."]}

Leptodactylus thomei sp. nov. (Figures 1���3) Holotype. MBML 2521, adult male, collected at Povoa����o, state of Esp��rito Santo, Brazil, on 28 August 2002, by Antonio de Padua Almeida, and recorded by Ariadne Angulo. Paratypes. Twenty-seven specimens collected at the type locality: MBML 2515 ���2517, 25 August 2002; MBML 2518 ���2520, 26 August 2002; MBML 2522 ���2526, 27 August 2002; MBML 2527, 28 August, 2002, collected by Antonio de Padua Almeida and some recorded by Ariadne Angulo; MBML 2302 ���03, 19 September 2001; MBML 2306, 21 September 2000; MBML 2307, MBML 2311, MBML 2314, 0 2 May 2001; MBML 2309 ��� 10 and MBML 2312 ���13, 18 September 2001, collected by Antonio de Padua Almeida and Rogerio Penha da Silva; MBML 2297, MBML 2300, 0 1 May 2002; MBML 2304 ���05, 0 7 June 2001; MBML 2308, 16 September 2001, collected by Antonio de Padua Almeida. Diagnosis. The advertisement call of Leptodactylus thomei differs from the advertisement calls of all other described species of the L. marmoratus group with known vocalizations. Leptodactylus thomei further differs from Leptodactylus andreae M��ller, 1923 by the lack of flattened toe discs and the presence of a mask-like pattern formed by an inverted triangle found in the interorbital region. The new species differs from L. araucaria (Kwet & Angulo, 2002) in: (1) size (maximum SVL of L. thomei males 23.2 mm and females 23.1, in L. araucaria males 18.8 mm and females 19.9 mm), (2) the presence of visible white tubercles on tibia (absent in L. araucaria), and (3) lack of dorsal longitudinal dark marks and light dorsolateral and mid-dorsal stripes in L. thomei (present in most L. araucaria). Leptodactylus thomei differs from L. bokermanni Heyer, 1973 by its marginally smaller size and by having a mask-like pattern embedded in the tan triangle patch of the interorbital region (but see discussion). It can also be distinguished from Leptodactylus diptyx Boettger, 1885 by the advertisement call and egg clutch size [58 unpigmented eggs in L. diptyx (De la Riva 1995), 29-33 cream-coloured eggs in L. thomei (Almeida & Angulo 2002 and this study)]. It can be distinguished from Leptodactylus heyeri (Boistel, Massary & Angulo, 2006) by: (1) advertisement call, (2) the lower surface of the foot with several small and distinct tubercles, (3) males lacking a yellow throat, and (4) a distinct dorsal colouration pattern. Leptodactylus thomei can be distinguished from Leptodactylus hylaedactylus (Cope, 1868) by its aquatic reproductive mode (Almeida & Angulo 2002). The new species differs from Leptodactylus lutzi (Heyer, 1975) given that L. lutzi is a larger species (males 30 mm and females up to 34 mm) and has a characteristic spotting on the posterior face of the thigh (Heyer 1975). It can be distinguished from Leptodactylus marmoratus (Steindachner, 1867) by (1) not possessing flattened toe discs, (2) possessing a mask-like patch in the interorbital region, and (3) having an aquatic reproductive mode. Leptodactylus thomei lacks the four longitudinal series of black dorsal spots that characterize L. martinezi Bokermann, 1956. Description of holotype. Body small, robust, with short limbs (see Table 1). Outline of snout subelliptical to subovoid, laterally the snout is acuminate, head as long as broad (Fig. 1). Nostrils anterior, dorsolateral, equally distant from snout and eye. Tympanum distinct, tympanum diameter almost half of eye diameter. Supratympanic fold poorly developed, but crevice extending from back of tympanum to arm insertion. Supratympanic melanophore markings dark and continuous from back of eye to back of tympanum, sparser and discontinuous from back of tympanum to arm insertion. Canthus rostralis indistinct, loreal region slightly concave. A rounded, cream-coloured oval gland is present at the angle of jaw. Vocal sac single and internal, a pair of vocal slits present; tongue oval to elongate. Short series of vomerine teeth posterior to choanae. Arms short, robust. Finger lengths III>I>II>IV; finger tips rounded, not expanded; fingers without webbing or fringes (Fig. 2 a). Two large metacarpal tubercles; size of outer, circular, metacarpal tubercle about three times that of inner, ovoid, metacarpal tubercle; prominent, rounded subarticular tubercles on fingers, very pronounced on the thumb. Nuptial asperities absent. Hindlegs short, thigh and shank of similar length. Toe lengths IV>III>V>II>I; toe tips rounded, neither expanded nor flattened; toes without webbing or fringes (Fig. 2 b). Two distinct, light, metatarsal tubercles; inner metatarsal tubercle ovoid, about twice as large as outer, rounded metatarsal tubercle. Sole of foot and lower surface of tarsus with several small, but relatively distinct tubercles. Upper shank surface smooth, with perceptible but sparse white tubercles. Dorsal texture slightly rugose, with more conspicuous tubercles placed in two longitudinal, discontinous, and glandular lateral folds extending from tympanum to inguinal region. A pair of cream oval glands present on either side of the anus. Ventral surface smooth. Colour in life. The dorsum is greyish-brown, with two dark dorsolateral melanophore lines running discontinuously from the back of the eye to thigh insertion, with a greater discontinuous patch in the shoulder area. Medially, the shoulder area also has a chevron shaped mark. There are two light and irregular blotches within the dark triangular patch of the interorbital region, giving the patch the appearance of a mask. This patch extends from the interorbital region to the back of head, roughly terminating at the tympanum level. Posteriorly and dorsally, there is a tan blotch on either side of the insertion of the legs. There are faded to bright orange patches on the heels, flanks, and dorsal scapular area. Venter is immaculate white. Iris is reticulated and copper-like in colour. Colour in preservative. Dorsum greyish-brown, with dark melanophore marks of dorsal pattern as previously described. Venter white, immaculate; throat white, with a slight shadow following the contour of lower lip. Measurements of holotype (in mm). SVL 21.2, HL 5.9, HW 5.9, ED 2.4, TYD 1.1, END 1.52, IOD 1.9, IND 2.3, FAL 1.4, HDL 3.6, TL 8.6, SL 8.1, TSL 3.9, FL 8.2. Morphological variation. Two irregular lighter-coloured blotches within the dark triangular patch of the interorbital region, which give the patch the appearance of a mask, are commonly found in members of the new species (93 % of examined specimens, n = 28, in other specimens the interorbital triangular patch is not continuous, as the inner blotches can be joined to the lighter dorsal pattern outside the triangle). Most specimens examined Males (n = 20) Females (n = 4) commonly have a light mid-dorsal hairline stripe whose extension can vary from above the vent to the sacral region (79 % of examined specimens, n = 28, absent in the other 21 %); white or cream oval glands on either side of the anus (60 % of examined specimens, n = 28, not visible in the other specimens), toe tips either slender, pointed, or with very slight expansions (stages 1 A and B of Heyer 1973), without discs. Larval morphology. The tadpole of Leptodactylus thomei is illustrated in Figure 4. Body piriform in dorsal view, almost twice as long as it is wide; snout narrowly rounded in profile; nostril marginally nearer to tip of snout than to eye; distance between nares slightly less than interorbital width; nares somewhat triangular, dorsolateral and directed anterolaterally; eyes dorsal; opening of the lateroventral sinistral spiracle directed posterolaterally, with short spiracular tube, positioned around halfway between tip of snout and body-tail juncture. Vent tube short and median; maximum body height slightly greater than maximum tail height; dorsal fin origin on the body-tail juncture; dorsal and ventral fins slightly arched; tail musculature reaching the tail tip; myotomes distinct over the last two thirds of tail; tail tip rounded to pointed. Mouth anteroventral; oral disc Range Mean S Total length (TL) 11.3���17.9 16.1 1.45 Body length (BL) 4.3���6.4 5.7 0.48 Tail length (TAL) 7.0��� 11.8 10.3 1.06 Maximum tail height (MTH) 2.0��� 3.4 2.8 0.32 Tail muscle height (TMH) 0.8���1.8 1.4 0.21 Tail muscle width (TMW) 0.7���1.4 1.2 0.19 Body height (BH) 2.5���3.8 3.1 0.29 Body width (BW) 3.0��� 4.4 3.9 0.33 Internarial distance (IND) 0.5 ���1.0 0.8 0.12 Interorbital distance (IOD) 0.8���1.1 0.9 0.09 Snout-nostril distance (SND) 0.3���0.6 0.4 0.08 Snout-eye distance (SED) 0.8���1.2 1.1 0.11 Eye-nostril distance (END) 0.3���0.6 0.5 0.08 Oral disc width (OD) 1.1���1.8 1.6 0.16 Eye diameter (ED)* 0.52���0. 62 0.58 0.06 ED/BL* 0.09���0.12 0.1 0.01 OD/BL 0.22���0.34 0.28 0.02 surrounded by a single row of rounded marginal papillae with a broad rostral gap; submarginal papillae absent; tooth row formula 2 (2)/ 3; rows of similar length; beak cornified, serrated; lateral line system not visible. Measurements are presented in Table 2; an illustration of the oral disc can be seen in Figure 5. Larvae colour in life. Dorsum tan to dark-brown, with scattered golden speckles; spiracle transparent; ventral wall transparent; internal organs visible; the posterior half with golden spots which present an iridescent aspect under stereomicroscopic examination; tail musculature mottled, with the fins slightly mottled; conspicuous mottling present in upper fin. Dorsal aspect of hind limbs with dark-brown bands alternated with unpigmented bands; this pattern is also found in the toe rudiments. Advertisement call. This call was first described and figured in Angulo (2004) under Adenomera ���Linhares���. One of the recordings used in Angulo���s (2004) analysis was replaced with another one in this study, given that the removed recording could potentially constitute an intermediate signal or a signal with a different function. The holotype MBML 2521 was recorded at 01:08 hrs on August 27, 2002, at an air temperature of 17.7 ��C and 92 % relative humidity. A total of six calls were recorded for this male and were used to describe the acoustic parameters measured herein (see Table 3, Figure 6). Call length average is 189.5 ms (range 165-210 ms), emitted at a low call rate (18 calls/min), with a mean call rise time of 71.73 ms, but ranging from 53-90 ms. The call is distinctly pulsed, audible to the human ear with a deep and regular amplitude modulation, consisting of 17���21 pulses/call, an average pulse rate of 111 pulses/s, average pulse duration of 9 ms, and average pulse rise time of 6 ms. Pulses rise gradually in increasing order of amplitude over the first third of the call, are roughly sustained over the middle third, and then smoothly decrease in amplitude, tapering off to background noise levels over the final third of the signal. The waveform representation of the call looks, overall, symmetrical. The dominant frequency oscillates around 5223 Hz and is approximately twice the fundamental frequency (average of 2505 Hz), comprising a harmonically-related formant. The maximum number of harmonically-related frequencies detected for this call is six or seven. The call has an average upward change in dominant frequency of 1881 Hz, but ranges from 1809���1981 Hz. Advertisement call variation. Table 3 shows measurements of acoustic parameters of calls for five vouchered males, including the holotype, and three additional unvouchered recordings (but see Table 3 for unavailable voucher 1). Temperatures are also provided for each set of recordings. Overall, the call of L. thomei is of intermediate to long duration (call length range 120���210 ms), when compared to advertisement calls of other Leptodactylus of the L. marmoratus group; it is emitted at low call rates (10���24 calls/min), albeit widely variable, very likely influenced by the motivation of callers and interactions with neighbouring males. All advertisement calls examined exhibit a pulsed structure, although it is not always possible to distinguish individual pulses from the waveform. Calls usually sound pulsed due to the very deep and regular amplitude modulation, although the pressure level of this modulation does not go down to zero in examined calls. If pulses can be discretely observed, the number varies from 10���21 per call, and ranges *One of these callers is in fact vouchered (MBML 2520); however, since both were recorded simultaneously, and since it is difficult to determine which of the calls was issued by the voucher, we have not assigned calls to this voucher. from 97���154 pulses/s. The amplitude patterns of calls vary according to the individual caller. The dominant frequency ranges from 4566 to 5562 Hz, and it corresponds to the next harmonic following the fundamental frequency (range 2153���2811 Hz). Between three to possibly seven harmonics can be detected for this call. The call shows upward change in the main carrier which varies from mild to very strong modulation (range 603 to 2153 Hz). This change in dominant frequency is variable both within and among individuals. Agonistic calls. When offered playbacks of their own call type or when neighbouring males call in close proximity, often males will change their call type. The acoustic signals produced under these circumstances differ from the advertisement call and do not resemble any other known advertisement call. There appear to be at least two types of vocalizations in situations of conflict for this species. One is a short ���chuckle��� (depicted in Figure 7), comparable to aggressive signals in other species of the L. marmoratus group (Angulo 2004), while the other is a signal which seems to be intermediate between the ���chuckle��� and advertisement call. The ���intermediate��� signal is often times heard as a single call, whereas ���chuckles��� have been heard as quick, consecutive calls or issued in groups of three. Agonistic signals were recorded for three males on 28 August 2002 at 20.4 �� C (n = 1 call), 19.9 ��C (n = 7 calls) and 21.5 ��C (n = 7 calls). Of these recordings, one set corresponds to what we believe may be ���intermediate��� calls (n = 7 recorded at 21.5 ��C) and the other two to ���chuckles���. Angulo (2004) provided descriptions for both these calls types, which are summarised herein. The ���intermediate��� call length varies from 28���127 ms with a mean duration of 101 ms. The call rate is ca 53 calls/minute, call rise time varies from 3 to 66 ms. There is pulse structure, with about 3���13 pulses per call. The fundamental frequency varies between 2173 and 2253 Hz, and dominant frequencies can range between 4526 and 4705 Hz; the dominant frequencies are the second harmonicallyrelated bands of the signal. Change in dominant frequency varies from moderate to intense (range 948���1723 Hz), and the number of harmonic bands detected is about 3. ���Chuckles��� (see Figure 7), on the other hand, are shorter (average 64 ms, range 50���91 ms), and with a much higher call rate, ranging from 205���227 calls/min if they were to be issued continuously. The number of pulses per call decreases to 3���6, and pulses seem to change in amplitude pattern as compared with the advertisement call. With two exceptions, where the main carrier seems to become the third harmonically-related frequency (6041 and 6300 Hz), the dominant frequency is the second harmonically-related band (range 3848���4885 Hz), and a fundamental frequency that ranges between 1874���2412 Hz. ���Chuckles��� contain up to five or six emphasized harmonic bands, with the first three harmonically-related frequencies with the greatest energy content. ���Chuckles��� are noisier across the frequency spectrum (see Figure 7 d). This acoustic signal also shows an upward change in dominant frequency towards the end of the call. Natural history. Individuals of the new species are found inhabiting the extensive leaf litter carpet which covers cocoa plantations within alluvial forests, although they display a patchy distribution during the reproductive season, with males aggregating into vocalizing groups. Reproductive activity starts around mid-August and continues until March, which coincides with the rainy season in this area. Calling activity can usually start around midafternoon or dusk, sometimes as early as midday during rainy or cloudy days, and continue well into the night. It increases dramatically at the beginning of and after showers. The species is fairly abundant during the rainy season. No other Leptodactylus of the L. marmoratus group is known to occur in this habitat. Reproduction. Reproduction of this species involves egg deposition in foam nests in earthen chambers and a subsequent aquatic phase necessary for the tadpoles to complete development (Almeida & Angulo 2002). During field work, we collected a gravid female (MBML 2522), which upon capture expelled an egg. She was kept in a plastic bag overnight. On the following day, the female was sitting on about 0.5 cm of a layer of foam, where all remaining eggs (n = 32) had been released. Eggs were cream-coloured, with an average diameter of 2.3 mm (range 1.1���2.85 mm, SD= 0.36). Etymology. The new species is named after Jo��o Carlos Alciati Thom��, in recognition for his conservation efforts on the Doce River Coastal Plain, northern Esp��rito Santo, which encompasses the type locality of the new species. Known geographic distribution. The new species has been collected at one locality in the state of Esp��rito Santo (Povoa����o, municipality of Linhares) and it probably also occurs in Governador Lindemberg, Esp��rito Santo (based on heard calls and one collected specimen from this site���MBML 5736) and in southern Bahia (based on examination of museum specimens���see Appendix). Although uncommonly encountered in the dry season, in the rainy season this species is frequently heard and observed., Published as part of Almeida, Antonio De Padua & Angulo, Ariadne, 2006, A new species of Leptodactylus (Anura: Leptodactylidae) from the state of Esp��rito Santo, Brazil, with remarks on the systematics of associated populations, pp. 1-25 in Zootaxa 1334 on pages 4-17, DOI: 10.5281/zenodo.174274, {"references":["Muller, L. (1923) Neue oder seltene reptilien und batrachier der Zoologischen Sammlung des bayerischen Staates. Zoologischer Anzeiger, 57, 39 - 54.","Kwet, A. & Angulo, A. (2002) A new species of Adenomera (Anura, Leptodactylidae) from the Araucaria forest of Rio Grande do Sul (Brazil), with comments on the systematic status of southern populations of the genus. Alytes, 20 (1 - 2), 28 - 43.","Heyer, W. R. (1973) Systematics of the marmoratus group of the frog genus Leptodactylus (Amphibia, Leptodactylidae). Contributions in Science, Natural History Museum, Los Angeles County, 251, 1 - 50.","Boettger, O. (1885) Liste von reptilien und batrachiern aus Paraguay. Zeitschrift fur Naturwissenschaft, 58, 1 - 36.","Marquez, R., De la Riva, I. & Bosch, J. (1995) Advertisement calls of Bolivian Leptodactylidae (Amphibia, Anura). Journal of Zoology, London, 237, 313 - 336.","Almeida, A. P. & Angulo, A. (2002) Adenomera aff. marmorata (NCN). Reproduction. Herpetological Review, 33 (3), 197 - 198.","Boistel, R., Massary, J. - C. (de) & Angulo, A. (2006) Description of a new species of the genus Adenomera (Amphibia, Anura, Leptodactylidae) from French Guiana. Acta Herpetologica, 1, 1 - 14.","Cope, E. D. (1868) An examination of the reptilian and Batrachia obtained by the Orton Expedition to Ecuador and upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences, Philadelphia, 1868, 96 - 140.","Heyer, W. R. (1975) Adenomera lutzi (Amphibia: Leptodactylidae), a new species of frog from Guyana. Proceedings of the Biological Society of Washington, 88 (28), 315 - 318.","Steindachner, F. (1867) Reise der Osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859. Amphibien. Kaiserlich-Konigliche Staatsdruckerei. Wien, 70 pp.","Bokermann, W. C. A. (1956) Sobre una nueva especie de Leptodactylus del Brasil (Amphibia). Neotropica, 2 (8), 37 - 40.","Angulo, A. (2004) The evolution of the acoustic communication system in members of the genus Adenomera (Anura: Leptodactylidae): A comparative approach. Doctoral thesis, University of Toronto, Toronto, 232 pp."]}

We describe a new species of the genus Adenomera from French Guiana. Collecting conditions, details about the localities and microhabitats are given. The advertisement call was recorded and is herein analyzed and described. Morphological and bioacoustic comparisons are drawn with other species of the genus. The new species is readily distinguished from other taxa by its distinctive coloration pattern, the occurrence of dorsolateral ridges and inguinal glands, and by the longer duration of the notes of its advertisement call. The taxonomy of Adenomera is evaluated with regard to current available knowledge., Acta Herpetologica, Vol 1 No 1 (2006)