Addenda and errata to “Description of a new species of the genus Adenomera (Amphibia, Anura, Leptodactylidae) from French Guiana"

In the first issue of Acta Herpetologica, Boistel et al. (2006) described a new species of Adenomera (Amphibia, Anura, Leptodactylidae). Because of an unfortunate set of events, there are several aspects of this paper that have to be corrected, or elaborated on. We address these issues herein

The Impact of Conservation on the Status of the World\u27s Vertebrates

Using data for 25,780 species categorized on the International Union for Conservation of Nature Red List, we present an assessment of the status of the world\u27s vertebrates. One-fifth of species are classified as Threatened, and we show that this figure is increasing: On average, 52 species of mammals, birds, and amphibians move one category closer to extinction each year. However, this overall pattern conceals the impact of conservation successes, and we show that the rate of deterioration would have been at least one-fifth again as much in the absence of these. Nonetheless, current conservation efforts remain insufficient to offset the main drivers of biodiversity loss in these groups: agricultural expansion, logging, overexploitation, and invasive alien species

Assessing the Cost of Global Biodiversity and Conservation Knowledge

Knowledge products comprise assessments of authoritative information supported by standards, governance, quality control, data, tools, and capacity building mechanisms. Considerable resources are dedicated to developing and maintaining knowledge products for biodiversity conservation, and they are widely used to inform policy and advise decision makers and practitioners. However, the financial cost of delivering this information is largely undocumented. We evaluated the costs and funding sources for developing and maintaining four global biodiversity and conservation knowledge products: The IUCN Red List of Threatened Species, the IUCN Red List of Ecosystems, Protected Planet, and the World Database of Key Biodiversity Areas. These are secondary data sets, built on primary data collected by extensive networks of expert contributors worldwide. We estimate that US$160 million (range: US$116–204 million), plus 293 person-years of volunteer time (range: 278–308 person-years) valued at US$14 million (range US$12–16 million), were invested in these four knowledge products between 1979 and 2013. More than half of this financing was provided through philanthropy, and nearly three-quarters was spent on personnel costs. The estimated annual cost of maintaining data and platforms for three of these knowledge products (excluding the IUCN Red List of Ecosystems for which annual costs were not possible to estimate for 2013) is US$6.5 million in total (range: US$6.2–6.7 million). We estimated that an additional US$114 million will be needed to reach pre-defined baselines of data coverage for all the four knowledge products, and that once achieved, annual maintenance costs will be approximately US$12 million. These costs are much lower than those to maintain many other, similarly important, global knowledge products. Ensuring that biodiversity and conservation knowledge products are sufficiently up to date, comprehensive and accurate is fundamental to inform decision-making for biodiversity conservation and sustainable development. Thus, the development and implementation of plans for sustainable long-term financing for them is critical

Shortfalls and Solutions for Meeting National and Global Conservation Area Targets

Governments have committed to conserving greater than or equal to 17% of terrestrial and greater than or equal to 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 km super(2) terrestrial PA network needs only 3.3 million km super(2) 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

Assessing the cost of global biodiversity and conservation knowledge

Knowledge products comprise assessments of authoritative information supported by stan-dards, governance, quality control, data, tools, and capacity building mechanisms. Considerable resources are dedicated to developing and maintaining knowledge productsfor biodiversity conservation, and they are widely used to inform policy and advise decisionmakers and practitioners. However, the financial cost of delivering this information is largelyundocumented. We evaluated the costs and funding sources for developing and maintain-ing four global biodiversity and conservation knowledge products: The IUCN Red List ofThreatened Species, the IUCN Red List of Ecosystems, Protected Planet, and the WorldDatabase of Key Biodiversity Areas. These are secondary data sets, built on primary datacollected by extensive networks of expert contributors worldwide. We estimate that US$160million (range: US$116–204 million), plus 293 person-years of volunteer time (range: 278–308 person-years) valued at US$14 million (range US$12–16 million), were invested inthese four knowledge products between 1979 and 2013. More than half of this financingwas provided through philanthropy, and nearly three-quarters was spent on personnelcosts. The estimated annual cost of maintaining data and platforms for three of these knowl-edge products (excluding the IUCN Red List of Ecosystems for which annual costs were notpossible to estimate for 2013) is US$6.5 million in total (range: US$6.2–6.7 million). We esti-mated that an additional US$114 million will be needed to reach pre-defined baselines ofdata coverage for all the four knowledge products, and that once achieved, annual mainte-nance costs will be approximately US$12 million. These costs are much lower than those tomaintain many other, similarly important, global knowledge products. Ensuring that biodi-versity and conservation knowledge products are sufficiently up to date, comprehensiveand accurate is fundamental to inform decision-making for biodiversity conservation andsustainable development. Thus, the development and implementation of plans for sustain-able long-term financing for them is critical

A flagship for Austral temperate forest conservation: an action plan for Darwin's frogs brings key stakeholders together

Darwin’s frogs Rhinoderma darwinii and Rhinoderma rufum are the only known species of amphibians in which males brood their offspring in their vocal sacs. We propose these frogs as flagship species for the conservation of the Austral temperate forests of Chile and Argentina. This recommendation forms part of the vision of the Binational Conservation Strategy for Darwin’s Frogs, which was launched in 2018. The strategy is a conservation initiative led by the IUCN SSC Amphibian Specialist Group, which in 2017 convened 30 governmental, non-profit and private organizations from Chile, Argentina and elsewhere. Darwin’s frogs are iconic examples of the global amphibian conservation crisis: R. rufum is categorized as Critically Endangered (Possibly Extinct) on the IUCN Red List, and R. darwinii as Endangered. Here we articulate the conservation planning process that led to the development of the conservation strategy for these species and present its main findings and recommendations. Using an evidence-based approach, the Binational Conservation Strategy for Darwin’s Frogs contains a comprehensive status review of Rhinoderma spp., including critical threat analyses, and proposes 39 prioritized conservation actions. Its goal is that by 2028, key information gaps on Rhinoderma spp. will be filled, the main threats to these species will be reduced, and financial, legal and societal support will have been achieved. The strategy is a multi-disciplinary, transnational endeavour aimed at ensuring the long-term viability of these unique frogs and their particular habitat

Leptodactylus coca, SP. NOV.

LEPTODACTYLUS COCA SP. NOV. (FIGS 1, 2) <p> <i>Holotype:</i> Adult male, NKA 3630, field number SR 99.134, collected near Carretera Antigua from Villa Tunari to Cochabamba (<i>c</i>. 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.</p> <p> <i>Paratypes:</i> 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.</p> <p>DIAGNOSIS</p> <p> <i>Leptodactylus coca</i> differs in its advertisement call from all other known members of the <i>L. marmoratus</i> group for which acoustic data are available. <i>Leptodactylus araucaria</i> (Kwet & Angulo, 2002) and <i>L. nanus</i> can be distinguished from <i>L. coca</i> by their small size [maximum SVL in <i>L. araucaria</i> males, 18.8 mm (Kwet & Angulo, 2002); maximum SVL in <i>L. nanus</i> males, 19.4 mm (Kwet, 2007)], whereas <i>Leptodactylus lutzi</i> (Heyer, 1975) differs from the new species by its large size [SVL in males, 25.7–33.5 mm (Kok <i>et al.</i>, 2007)], the presence of a dark triangular seat patch, and distinct spotting on the posterior face of the thigh (Heyer, 1975). Individuals of <i>Leptodactylus bokermanni</i> 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 <i>L. coca</i>. The new species differs from <i>Leptodactylus martinezi</i> Bokermann, 1956 in lacking four longitudinal series of dark glands on the dorsum (Bokermann, 1956). <i>L. andreae</i>, <i>L. diptyx</i>, <i>L. hylaedactylus</i>, <i>Leptodactylus marmoratus</i> Steindachner, 1867, and <i>Leptodactylus thomei</i> Almeida & Angulo, 2006 all have distinct, different advertisement calls to those of <i>L. coca</i>. There is a recently described species of <i>Leptodactylus</i> of the <i>marmoratus</i> group from French Guiana, <i>Leptodactylus heyeri</i> (Boistel, de Massary & Angulo, 2006), which has a similar call to that of <i>L. coca</i>. 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 <i>L. heyeri</i> is issued at a much lower call rate than that of <i>L. coca</i>, and the call of <i>L. heyeri</i> does not show evidence of pulse structure, whereas it is present in <i>L. coca</i>. The species also differ morphologically: <i>L. heyeri</i> 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.</p> <p>DESCRIPTION OF HOLOTYPE</p> <p> 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>></i> I = II <i>></i> 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 <i>></i> III <i>></i> V <i>></i> II <i>></i> 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.</p> <p>* Holotype.</p> <p>COLOUR IN PRESERVATIVE</p> <p>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.</p> <p>COLOUR IN LIFE</p> <p>The coloration in life does not differ much from the colour described in preservative, just more intense.</p> <p>MEASUREMENTS OF HOLOTYPE (IN MM)</p> <p>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.</p> <p>MORPHOLOGICAL VARIATION</p> <p> 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 <i>sensu</i> Heyer, 1973) and flattened.</p> <p>* Holotype.</p> <p> 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 <i>sensu</i> Heyer, 1973) and mildly flattened or not flattened.</p> <p>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.</p> <p>ADVERTISEMENT CALL</p> <p> 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 <i>L. coca</i> has been previously described in Angulo (2004) as <i>Adenomera</i> ‘Chapare’; it is here directly associated to the species name <i>L. coca</i>. 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 Márquez <i>et al</i>. (2002).</p> <p> In comparison with other species of the <i>L. marmoratus</i> group (see Kwet & Angulo, 2002; Angulo <i>et al</i>., 2003; and references within; Kok <i>et al.</i>, 2007; and this study, for acoustic parameters for other species) the advertisement call of <i>L. coca</i> 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).</p> <p>NATURAL HISTORY</p> <p> 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 <i>L. marmoratus</i>; Lutz, 1931, 1947).</p> <p>ETYMOLOGY</p> <p> The new species is named after the coca plant, <i>Erythroxylon coca</i>, given its occurrence near coca plantations of the Chapare region, in Bolivia.</p> <p>CONSERVATION STATUS</p> <p>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).</p>Published as part of <i>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)</i> on pages 62-67, DOI: 10.1111/j.1096-3642.2008.00447.x, <a href="http://zenodo.org/record/4687663">http://zenodo.org/record/4687663</a&gt

Notes on range extension and geographic variation of calls in Adenomera thomei (Anura: Leptodactylidae)

New localities are herein reported for Adenomera thomei in Minas Gerais state, increasing its known distribution towards the Brazilian northwest and expanding knowledge on its habitat use. The advertisement calls of Adenomera thomei here reported exhibit variation in the call parameters known for this species, but also reveal a degree of overlap with the calls of a closely related undescribed species, suggesting that further research is needed to elucidate geographic call variation and population identities

Leptodactylus thomei Almeida & Angulo, 2006, sp. nov.

Leptodactylus thomei sp. nov. (Figures 1–3) Holotype. MBML 2521, adult male, collected at Povoação, state of Espírito 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 Müller, 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 ºC 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 º C (n = 1 call), 19.9 ºC (n = 7 calls) and 21.5 ºC (n = 7 calls). Of these recordings, one set corresponds to what we believe may be “intermediate” calls (n = 7 recorded at 21.5 ºC) 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 João Carlos Alciati Thomé, in recognition for his conservation efforts on the Doce River Coastal Plain, northern Espírito 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 Espírito Santo (Povoação, municipality of Linhares) and it probably also occurs in Governador Lindemberg, Espírito 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 Espírito Santo, Brazil, with remarks on the systematics of associated populations, pp. 1-25 in Zootaxa 1334 on pages 4-17, DOI: 10.5281/zenodo.17427