Paulo Passos

Paulo Melo-Sampaio

Luciana Ramos

Felipe Grazziotin

Antoine Fouquet
email: fouquet.antoine@gmail.com

Omar Torres-Carvajal

Antoine Suffix

Albertina P Suffix

Alexander T Suffix

Raffael Suffix

Familyname Kok

Philippe J R Suffix

Pedro Peloso

Rawien Jairam

Albertina P Lima

Alexander T Mônico

Raffael Ernst

Philippe J R Kok

P Cf Ockendeni

P Grandoculis

P Cf Marmoratus

Particle Given Name

Keywords: separated by '-') Amazonia -Amphibia -Morphology -Tomography Amazonia, Amphibia, Morphology, Tomography

published or not. The documents may come   L'archive ouverte pluridisciplinaire

Introduction

Pristimantis Jiménez de la Espada, 1870 is the most speciesrich genus among all vertebrates, currently containing more than 570 species mostly distributed throughout Amazonia, the Andes, the Chocó, and Pantepui [START_REF] Frost | Amphibian species of the world: An online reference[END_REF], and, albeit more marginally, in Central America, the Caribbean, the Cerrado, and the Atlantic Forest. Species in this genus are being named at a fast pace, with over 133 species described 1 3 in the last decade, and over 220 species since 2000 [START_REF] Frost | Amphibian species of the world: An online reference[END_REF]. This, along with diversity estimates based on DNA sequence data (e.g. [START_REF] Trevisan | Cryptic diversity and ancient diversification in the northern Atlantic Forest Pristimantis (Amphibia, Anura, Craugastoridae)[END_REF][START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF], suggests a vast number of unnamed species in this genus.

Many Pristimantis species occur at very narrow altitudinal ranges and have small distributions [START_REF] Duellman | Terrestrial-breeding frogs (Strabomantidae) in Peru[END_REF]. This pattern is most striking not only along Andean slopes and in the Pantepui region, but also in some lowland areas (e.g. [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF][START_REF] Kok | Two new charismatic Pristimantis species (Anura: Craugastoridae) from the tepuis of the "Lost World"(Pantepui region, South America)[END_REF]. Direct mode of development (endotrophic larvae that hatch as froglets) and narrow bioclimatic niches have been suggested as possible causes of high micro-endemism in this group [START_REF] Gonzalez-Voyer | Correlates of species richness in the largest Neotropical amphibian radiation[END_REF][START_REF] Vences | Speciation, species boundaries and phylogeography of amphibians[END_REF]. As a corollary, half of the 445 Pristimantis species evaluated by the International Union for Conservation of Nature (IUCN) are considered endangered (IUCN, 2020). It is therefore important to improve our knowledge of the actual diversity and distribution of Pristimantis species.

Pristimantis marmoratus [START_REF] Boulenger | Batrachians[END_REF] and P. ockendeni [START_REF] Boulenger | Descriptions of new batrachians from the Andes of South America, preserved in the British Museum[END_REF] have for long been used to designate multiple populations of superficially similar, hard to identify, small brown Pristimantis species of the "unistrigatus group" (sensu [START_REF] Hedges | New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation[END_REF], see discussion in [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]. In the Guiana Shield lowlands more specifically, P. marmoratus has been reported from northern Brazil, in the state of Pará [START_REF] Ávila-Pires | Notes on the vertebrates of northern Pará, Brazil: A forgotten part of the Guianan Region, I. Herpetofauna[END_REF], Guyana [START_REF] Ernst | On the cutting edge-the anuran fauna of the Mabura Hill Forest Reserve, central Guyana[END_REF][START_REF] Ernst | Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities[END_REF][START_REF] Ernst | Amphibian communities in disturbed forests: Lessons from the Neo-and Afrotropics[END_REF], French Guiana [START_REF] Lescure | Atlas des amphibiens de Guyane[END_REF], and Suriname [START_REF] Ouboter | Amphibians of Suriname[END_REF], whereas P. ockendeni has been reported from the Reserva Florestal Adolpho Ducke (RFAD) in the Amazonas state of Brazil [START_REF] Lima | Guia de sapos da Reserva Adolpho Ducke, Amazônia Central[END_REF]. However, [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF] clearly demonstrated that the range of P. marmoratus (type locality "foot of Mount Roraima", Venezuela, ca. 1067 m) is restricted to the eastern Pantepui uplands and highlands between 600 and 1800 m above sea level (asl), and [START_REF] Elmer | Three new species of leaflitter frogs from the upper Amazon forests: Cryptic diversity within Pristimantis "ockendeni"(Anura: Strabomantidae) in Ecuador[END_REF] suggested that the range of P. ockendeni (type locality "La Union, Rio Huacamayo, Carabaya, [Departamento Puno,] Peru, ca. 600 m") is restricted to the Andean foothills of southeastern Peru. Therefore, the Guiana Shield lowland populations that were previously identified as P. marmoratus and P. ockendeni likely remain unnamed, except for at least some "P. marmoratus" populations from Suriname for which a name is available: Hylodes grandoculis van Lidth de Jeude, 1904. The species was described based on a single specimen from central Suriname ("the basin and the sources of the Coppename") and is considered a synonym of P. marmoratus since [START_REF] Frost | Amphibian Species of the World[END_REF]. Several species cooccur in the region, and the conspecificity of the holotype of H. grandoculis with recently collected material needs, therefore, to be clarified. [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF] delimited three major lineages corresponding to populations previously identified as P. marmoratus or P. ockendeni: (1) Pristimantis cf. marmoratus, occurring in the easternmost part of the region (state of Amapá in Brazil and French Guiana) and corresponding to Pristimantis sp. 1 of [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF] and [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]; (2) Pristimantis cf. ockendeni, occurring in southern Suriname and central Guyana (Eleutherodactylus marmoratus of [START_REF] Ernst | On the cutting edge-the anuran fauna of the Mabura Hill Forest Reserve, central Guyana[END_REF][START_REF] Ernst | Diversity erosion beyond the species level: Dramatic loss of functional diversity after selective logging in two tropical amphibian communities[END_REF][START_REF] Ernst | Amphibian communities in disturbed forests: Lessons from the Neo-and Afrotropics[END_REF]; and (3) Pristimantis sp. "Guianas", occurring in French Guiana, in southern Amapá state (Brazil), Suriname, and at the border between Guyana and the state of Pará, Brazil, corresponding to Pristimantis sp. 4 of [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF] and [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF].

To (1) disentangle this confusing situation and determine if one of these lineages would correspond to Hylodes grandoculis, and ( 2) facilitate the description of potential related species, we examined the external morphology and osteology (via µ-CT scans) of the holotype of H. grandoculis, of the holotype of Pristimantis marmoratus, and of recently collected material for which we analyzed genetic, acoustic, and morphological variation.

Material and methods

Field work and deposition of specimens

We undertook extensive field work in French Guiana, Guyana, the Brazilian states of Amapá and Amazonas, and in Suriname (Bakhuis, Tafleberg, and Sipaliwini), including areas nearby the type locality of Hylodes grandoculis, collecting specimens and tissue samples, and recording advertisement calls and ecological data of numerous Pristimantis populations (Fig. 1). Frogs were collected by hand and killed with an intraperitoneal injection of Xylocaine ® (lidocaine hydrochloride). All specimens were individually tagged and fixed in 10% formalin before being transferred to a solution of 70% ethanol for permanent storage. Newly collected specimens were deposited in the Museum National d'Histoire Naturelle (Paris, France) (MNHN-RA-2020.0092-118), Museu Paraense Emílio Goeldi (MPEG 41819-41825), and the National Zoological collection of Suriname (NZCS A1210-1212). Other specimens examined are deposited in various zoological collections (Appendices 1 and 2). Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

Vacher et al. (2020) (Appendix 1). We employed BLASTN to verify that no other 16S sequence deposited on GenBank after the work of [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF] clustered with the species used in this study. Twenty-eight sequences were newly generated while the other sequences were retrieved from GenBank (Appendix 1). We completed these data with 13 sequences of 12S, eight retrieved from GenBank and five newly generated. Genomic DNA was extracted from tissue samples (muscle or liver) using either the Wizard ® Genomic DNA Purification Kit (Promega; Madison, WI, USA) or the DNeasy ® Blood & Tissue Kit (Qiagen; Hilden, Germany) following the manufacturer's protocols. We amplified targeted loci by standard PCR protocols and then used Sanger sequencing (primers 16RF: TAT CCC TAG GGT AAC TTG; 16FR: TTA CCA AAA ACA TCG CCT from [START_REF] Salducci | Phylogenetic relationships and biodiversity in hylids (Anura: Hylidae) from French Guiana[END_REF]. These sequences were aligned with the mitogenome of P. thymelensis (JX564889, which served as the outgroup), on the MAFFT7 online server under default parameters except the E-INS-i strategy, which is designed for sequences with multiple conserved domains and long gaps [START_REF] Katoh | MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization[END_REF]. We trimmed the alignment to keep only regions where sequences overlap, resulting in a final alignment of 2548 base pairs (bp). We investigated the phylogenetic relationships among a total of 191 terminals under maximum likelihood (ML), applying a GTR CAT model with RAxML v.8.2.4 [START_REF] Stamatakis | RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies[END_REF]. All RAxML analyses were performed on the CIPRES Science Gateway online server (Miller et al., 2010). Non-parametric bootstrapping values [START_REF] Felsenstein | Confidence limits on phylogenies: An approach using the bootstrap[END_REF] were estimated using 1,000 pseudoreplicates.

Morphometric analysis

We examined 67 adult specimens (45 males, 22 females, among which 27 were included in our molecular dataset) of P. cf. marmoratus [P. sp. 1 of [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF] and [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF], 13 males, 3 females], P. cf. ockendeni (8 males, 6 females), and Pristimantis sp. "Guianas" from Suriname (4 males, no females) and from French Guiana (13 males, 13 females) [P. sp. 4 of [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF] and [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]], and the holotype of Hylodes grandoculis (Appendix 2). The other specimens were a priori assigned to one of these species when they were collected from the same or nearby populations and after morphological examination. Sex was determined either in the field via calling activity, or in the laboratory by examining the condition of vocal slits, nuptial pads, and gonads. We measured 18 variables; abbreviations for measurements are as follows: SVL-snout-vent length; TiL-tibia length; FeL-femur length; TaL-tarsus length; FL-foot length (from outer edge of metatarsal tubercle to the tip of toe IV); HeL-head length; HW-head width; Ind-internarial distance; IOD-interorbital distance; ENeye-nostril distance (straight line distance between anterior corner of eye and nostrils); ED-horizontal eye diameter; TD-horizontal tympanum length; FD-disc width of Finger III; 4TD-disc width of Toe IV; ETS-eye-tip of snout distance (straight line distance between the anterior corner of eye and tip of snout); 1FiL-length of Finger I (from proximal edge of palmar tubercle to the tip of Finger I); 2FiLlength of Finger II (from proximal edge of palmar tubercle to the tip of Finger II); HL-hand length (from distal edge of palmar tubercle to the tip of Finger III). Measurements were taken using a digital caliper to the nearest 0.01 mm. Mean and ranges were rounded to the nearest 0.1 mm (all measurements by AF, except for MPEG specimens, measured by PP). We examined the variation of morphometric data among adult males through a principal component analysis (PCA) via FactoMineR package in R v.3.2.4 [START_REF] Lê | FactoMineR: An R package for multivariate analysis[END_REF]R Development Core Team, 2016). To control for variation in body size among individuals, we performed additional analyses on a size-corrected dataset (residuals) obtained by linear-regressing the original morphometric measures of each variable on SVL [START_REF] Strauss | Evolutionary allometry and variation in body form in the South American catfish genus Corydoras (Callichthyidae)[END_REF].

3D microtomography

We investigated the osteological variation across adult males of P. cf. marmoratus, Pristimantis sp. "Guianas" and the holotypes of Hylodes grandoculis (RMNH 4467) and P. marmoratus (BMNH 1947.2.16.92). Specimens were scanned using an EasyTom 150 (at kV = 40-70, resolution < 20 µm) from the MRI platform of ISEM (Institute of Evolutionary Sciences of Montpellier), except the holotype of H. grandoculis which was scanned using a CT/Zeiss Xradia at the Leiden Naturalis Biodiversity Center, and the holotype of P. marmoratus that was scanned using a Metris X-Tek HMX ST 225 System at the Natural History Museum, London. Segmentation of the full skeleton and of the cranium was done using Avizo (FEI Visualization Sciences Group, Burlington, MA, USA) and Biomedisa (Lösel et al., 2000). µ-CT scans and surface renderings will be deposited at www. morph osour ce. org upon manuscript acceptance.

Acoustic analysis

We gathered six call recordings of males of Pristimantis cf. marmoratus (including one track file from [START_REF] Marty | Sound guide to the tailless amphibians of French Guiana[END_REF]. To our knowledge, Pristimantis sp. "Guianas" has never been observed calling and no recording is available. The external tympanum and pharyngeal ostia are absent in that species, and males lack vocal slits. The calls of P. cf. marmoratus consist in series of short notes, and we measured seven call variables from waveforms using Audacity v.2.1.1 following [START_REF] Köhler | The use of bioacoustics in anuran taxonomy: Theory, terminology, methods and recommendations for best practice[END_REF]: call length, silence between calls, number of notes, note length, silence between the two first notes, silence between the two last notes, dominant frequency (taken with a spectral slice over the entire note). Whenever possible, up to four measurements of each variable were taken per recorded male, and the average was considered as a single measurement. Call recordings will be deposited at www. sonot heque. mnhn upon manuscript acceptance.

Results

Phylogenetic relationships

The relationships among species inferred from the ML analysis are poorly supported at several nodes (Fig. 1). However, sequences from individuals corresponding to the different putative species form strongly supported clades. The populations previously identified as P. cf. ockendeni from the Guiana Shield (Suriname and Guyana) form a clade related to several populations south of the Amazon River, which we hereafter refer to as the "trans-amazon complex". The Guiana Shield populations are more closely related to a single specimen from São Sebastião, Rio Abacaxis, Amazonas (p distance = 5.4%, Table 3). The "trans-amazon complex" is highly divergent from other Guiana Shield species, and is more closely related to western Amazonian species, although probably not directly related to P. ockendeni [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF]. Pristimantis cf. marmoratus is circumscribed to the easternmost part of the Guiana Shield and forms a poorly supported clade with P. marmoratus sensu stricto. Pristimantis sp. "Guianas" samples form a strongly supported clade with uncertain relationships and are subdivided into two main lineages (p distance = 4.5%, Table 3), one occurring in the west (Suriname, the Amapá and Pará states in Brazil) and the other one in the east (French Guiana). The geographic range of the western lineage of Pristimantis sp. "Guianas" encompasses the type locality of P. grandoculis. Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

U

Morphometric analyses

The PCA based on raw morphometric measurements provided two first components accounting for 63.4% of the total variation (Fig. 2a). The first component explains 46.7% of the variation, and variable coefficients are all highly and positively correlated. Overall, the variation along the first principal component (PC) axis is related to body size; since the three species largely overlap on this variable, they also do along this axis. The variation along the second axis is notably related to differences in TiL and FL (negatively correlated with PC2) and ED (positively correlated with PC2) and segregates populations of Pristimantis sp. "Guianas", which largely overlap, from the other species that only partly overlap with each other. The holotype of P. grandoculis is positioned within the western populations of Pristimantis sp. "Guianas".

The PCA based on size-corrected morphometric measurements provided two first components accounting for 48.1% of the total variation (Fig. 2b). The variation along the first axis is notably related to differences in HL, FeL and FL (positively correlated with PC1), and ED (negatively correlated with PC1) and segregates Pristimantis cf. ockendeni from the other species that only partly overlap with each other (eastern and western populations of Pristimantis sp. "Guianas" completely overlap). The variation along the second axis is notably related to differences in HW (positively correlated with PC2) and TaL (negatively correlated with PC2) and partly segregates P. sp. "Guianas" from P. cf. marmoratus. The holotype of P. grandoculis is positioned within populations of P. sp. "Guianas" but also within P. cf. marmoratus.

The examination of the ratios between the aforementioned variables and SVL (Fig. 2c) shows limited overlap across species particularly considering HL/SVL and TiL/ SVL. Moreover, the ratios displayed by the holotype of P. grandoculis are always within the range of French Guiana and Suriname populations of P. sp. "Guianas" and only marginally overlaps with the range of the other species.

Taxonomic conclusion

External morphology and osteology (see below) strongly suggest that the western populations that have been identified as Pristimantis sp. "Guianas" [P. sp. 4 of [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF] and [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]] correspond to Hylodes grandoculis. This species is geographically disjunct and phylogenetically distinct from P. marmoratus sensu stricto based on molecular data (Fig. 1; see also [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]. Our morphological and osteological data also unambiguously confirm the non-conspecificity of P. marmoratus and H. grandoculis. Therefore, we formally rescue H. grandoculis from the synonymy of P. marmoratus, and herein provide an amended diagnosis of Pristimantis grandoculis n. comb. (van Lidth de Jeude, 1904) based on the examination of the holotype and four adult specimens from Suriname. It must be mentioned that the specimens of Pristimantis sp. "Guianas" from French Guiana, although morphologically very similar to P. grandoculis, form a distinct, closely related clade that upon the examination of a sufficient number of adult specimens (not currently available) could be either considered conspecific or a distinct species and will be hereafter referred to as P. sp. "Guianas" East. The easternmost species previously reported as Pristimantis sp. 1 [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF] and Pristimantis cf. marmoratus [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF] is also morphologically clearly distinct from P. grandoculis and all known congeners in the area and is formally described and named herein as P. crepitaculus sp. nov. The "trans-amazon complex", in which some populations have been erroneously identified as P. ockendeni, will be treated elsewhere. Holotype. RMNH 4467. Type locality: "the basin and the sources of the Coppename", interior of Surinam. van Lidth de Jeude stated that the specimen was collected "Sept. 10th 1901" during the Coppename expedition (Anonymous). The expedition took place between August and late November 1901 "we went up the Coppename then the two rivers that form it until they become unseaworthy even for the korjales (boats) of the natives (3°57 N)". These indications suggest that the specimen was collected at the farthest point reached by the expedition and that 3.9500°N 56.7543°W are plausible coordinates for the type locality.

Species accounts

Referred specimens. MNHN-RA-2020.0118 (field no. AF3435), an adult male collected by A. Definition. Pristimantis grandoculis is characterized by the following unique combination of characters: (1) SVL small, adult males 16.5 ± 1.1 mm (range 14.7-17.9 mm, n = 5) (Table 1), female unknown; (2) dorsal skin tuberculate, with two pairs of enlarged tubercles on the scapular region embedded in a W-shaped scapular fold, ventral skin granular particularly on the femoral region; (3) external tympanum absent, tympanic membrane not differentiated and obscured by supratympanic fold on the posterodorsal edge of tympanum, tympanic annulus indistinct; (4) pharyngeal ostia absent; (5) columella absent; (6) tibia length 55-59% of SVL; (7) snout broadly rounded in profile and dorsal view; (8) each upper eyelid with two prominent tubercles; (9) choanae round and small (0.3 mm for MNHN-RA-2020.0118), dentigerous processes of vomers oblique, narrowly separated, each bearing 3-5 odontophores; (10) vocal slits absent, vocal sac absent; (11) one unpigmented whitish nuptial pad located on the preaxial side of the thenar tubercle on each thumb in male; (12) FI much shorter than FII, reaching Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

Morphological comparisons with other Guiana Shield lowlands Pristimantis of the "unistrigatus group". Pristimantis grandoculis can mainly be distinguished from P. espedeus by its smaller body size (SVL range in males = 14.7-17.9 mm vs. 20.7-24.8 in P. espedeus); external tympanum absent (vs. visible in P. espedeus) and dark grey groin colouration in life (vs. reddish in P. espedeus).

Pristimantis grandoculis can mainly be distinguished from P. inguinalis by dark grey groin colouration in life (vs. bright yellow inguinal mark in P. inguinalis); external tympanum absent (vs. visible in P. inguinalis) and translucent brownish grey ventral colouration with small cream spots (vs. entirely black in P. inguinalis).

All the other known Guiana Shield species of the "unistrigatus group" occur in the Pantepui region and are generally associated with highlands (i.e. > 700 m elevation; [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF][START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]. Pristimantis grandoculis was previously confused (and considered a synonym) with P. marmoratus, one of these Pantepui species, from which it can immediately be distinguished by iris colour in life (black vertical streak running across the iris in P. marmoratus vs. no such streak in P. grandoculis) and absence of columella and vocal slits (vs. present in P. marmoratus). Among the other Pristimantis species found at mid-elevation in Pantepui, P. grandoculis can mainly be distinguished from P. jester [START_REF] Means | Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America[END_REF]) by having preaxial fringes on FII and III (absent in P. jester), toes basally webbed between TII-IV with preaxial fringes on TII-V (webbing and fringes on toes absent in P. jester) and by iris colour in life (golden to copper in P. grandoculis vs. upper 2/5 of iris blue-grey in P. jester); from P. saltissimus [START_REF] Means | Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America[END_REF] by tympanum condition (external tympanum absent, with indistinct tympanic annulus in P. grandoculis vs. distinct with distinct tympanic annulus in P. saltissimus), by having preaxial fringes on FII and III (absent in P. saltissimus) and toes basally webbed between TII-IV with preaxial fringes on TII-V (webbing and fringes on toes absent in P. saltissimus); from P. guaiquinimensis [START_REF] Schlüter | Three new frogs of the genus Eleutherodactylus (Amphibia, Leptodactylidae) from Guaiquinima table mountain, Bolívar. Venezuela[END_REF]; see also [START_REF] Kok | On the taxonomic validity of Pristimantis tepuiensis (Schlüter & Rödder, 2007) and P. stegolepis[END_REF] mostly by tympanum condition (external tympanum absent, with indistinct tympanic annulus in P. grandoculis vs. distinct with distinct tympanic annulus in P. guaiquinimensis); from P. sarisarinama (Barrio-Amorós & Brewer-Carías, 2008) mostly by the absence of vocal slits in males (present in P. sarisarinama); from P. pulvinatus [START_REF] Rivero | A new species of Eleutherodactylus (Amphibia, Salientia) from the Guayana Region, Edo. Bolívar. Venezuela[END_REF] mostly by smaller body size in males (14.7-17.9 mm in P. grandoculis vs. 23.0-26.1 mm in P. pulvinatus) and distinct vomerine teeth (indistinct or absent in P. pulvinatus); and from P. memorans (Myers & Donnelly, 1997) mostly by the absence of vocal slits in males (present in P. memorans), and tympanum condition (external tympanum absent, with indistinct tympanic annulus in P. grandoculis vs. distinct with distinct tympanic annulus in P. memorans).

Variation. The holotype is relatively well preserved. However, it is highly discoloured, and overall tuberculation is difficult to assess. Proportions are similar among the five specimens examined; the amount of tuberculation on the skin varies, especially on the dorsum and lateral surfaces of the body. Colour pattern is variable across the recently collected specimens. NZCS A1210 has a dark band running along the canthus rostralis which is absent in the other specimens (Fig. 2C). The W-shaped scapular mark is welldeveloped anteriorly but less so posteriorly compared to the other specimens. An interorbital band darker than the dominant background is often present as well as dark bands on the arms and legs but all these markings are variable in sharpness across specimens.

Osteology of the holotype (RMNH 4467). Cranium (Fig. 4). Shape and proportions. The skull is well ossified, widest posterior to the orbit at the level of the articulation of the maxilla with the quadratojugal. The rostrum is moderate, the braincase is broad.

Neurocranium and dorsal investing bones. The nasals (separated medially from one another and covering all the nasal capsules dorsally), frontoparietals, parasphenoid and neopalatines are co-ossified with the sphenethmoid. The frontoparietal and prootic are fused. Ventrally, the prootics are fused with the parasphenoid alae. The exoccipitals are fused. The dorsal surface of the otic capsule is ossified. Frontoparietal crests are absent and the frontoparietal fontanelle is exposed. The septomaxilla is roughly spiralled, the medial ramus extending posterodorsal to the posterior ramus; the anterior ramus is thick; the lateral ramus is oblique with a long acuminate posterolateral extension; the posterior ramus extends from the middle of the lateral ramus ventromedially. The columellae (stapes) are absent.

Ventral investing bones. The parasphenoid cultriform process extends anteriorly from the anterior edge of the otic capsule and is co-ossified with the sphenethmoid. The parasphenoid alae are moderately long (about half of the length of the cultriform process), perpendicular to the anteroposterior body axis, broadening slightly laterally. The vomers are fused with the sphenethmoid; each vomer is composed of an arcuate bone bordering the anteromedial, medial, and posterior margins of the choana. The prechoanal ramus is expanded medially and anteriorly and bears a ventral flange along its medial edge. The postchoanal process is narrow and acuminate, slightly posteriorly curved. Well-developed dentigerous processes extend posteromedially from the union of the pre-and postchoanal processes. Each dentigerous process bears three teeth and is broadly separated from its counterpart medially. The neopalatine is fused with the maxilla distally. This complex is fused with the parasphenoid medially.

Maxillary arcade. The maxillary arcade bears many small teeth on the premaxilla and maxilla. The arcade is complete and connected to the slender quadratojugal. The premaxillae are separated medially, and their anterodorsal alary process is weakly divergent from the midline. The pars palatina is broad, with two well-defined processes: the medial (palatine) process is relatively narrow and runs roughly parallel toward its contralateral; the lateral process is broader. The premaxilla and maxilla are in lateral contact via a simple juxtaposition. The maxilla is long, with a broad pars palatina along its lingual margin and a moderately developed pars facialis. Suspensory apparatus. The triradiate pterygoid bears a slightly curved anterior ramus with a sculpted ventrolateral face, oriented anterolaterally toward the maxilla, with which it articulates at approximately the mid-length of the orbit. Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

The pterygoid is fused to the maxilla. The medial and posterior face of the medial rami of the pterygoid are about equal in length. The medial ramus is broader than the posterior and its posterior face is strongly sculpted. The lateral end of the medial ramus overlaps the lateral edge of the prootic. The quadratojugal is long, laterally curved, and slender, articulating anteriorly with the maxilla. It has a bulbous posteroventral process and articulates dorsally with the ventral ramus of the squamosal. The squamosal is dorsally bifurcated, broad, and sculpted, extending anterodorsomedially from the quadratojugal to the level of the otic capsule; the zygomatic ramus is short, whereas the otic ramus is long, almost reaching the posterior end of the skull.

Mandible. The mandible is slim and edentate. The mentomeckelians are small and arcuate in ventral view, medially and laterally broadened, and in narrow contact medially. The dentary is short and thin, posteriorly acuminate, and overlaps the angulosplenial for about a quarter of its length. Dentary is in contact with the angulosplenial posteriorly. The angulosplenial is long and arcuate, laterally slightly grooved. The coronoid process is weak.

Hyoid. The bony posteromedial processes of the hyoid are expanded proximally and separated from one another. No ossified parahyoid is present.

Axial skeleton (Fig. 4).

Vertebral column. The vertebral column has eight procoelous, presacral vertebrae. The vertebrae have complete neural arches and low neural processes. Atlas (presacral I) and presacral II are fused medially and laterally. The transverse processes of presacrals II-III are slightly expanded distally, thicker, and broader than those of other presacrals. The transverse processes of presacrals II and III are oriented ventrolaterally (II anteriorly, III posteriorly), whereas those of presacrals IV-VIII extend dorsolaterally (IV-VII posteriorly, VIII anteriorly). The relative lengths of the transverse processes and sacral diapophyses are: III > IV > Sacrum > V ≈ VI ≈ VII ≈ VIII > II. The sacral diapophyses are slightly expanded distally. The urostyle has a well-developed dorsal ridge that extends along half its length.

Pectoral girdle. The zonal portion has well-ossified coracoids, clavicles, scapulae, and cleithra. The clavicles are long, slender, and oriented anteromedially; the medial tips are narrowly separated from one another and located anteriorly from the level of anterolateral end of the clavicle that articulates with the scapula; the coracoid is long and flared, with its sternal end slightly broader than its glenoid end. The scapula is long with a prominent pars acromialis that is not separated from the pars glenoidalis. The scapula is about one and a half the length of the clavicle. The cleithrum is ossified, well-developed, anteriorly thicker, thinning posteriorly. The suprascapula is unossified.

Pelvis girdle. The long, slender iliac shafts bear dorsolateral crests throughout their length. The overall length of the girdle is more than two times the width between the anterior ends of the iliac shafts. The iliac prominence is broad and low, and pubes mineralized.

Manus and pes. The phalangeal formulae for the hand and foot are standard, 2-2-3-3 and 2-2-3-4-3, respectively. Terminal phalanges strongly T-shaped. Ossified prepollex and prehallux visible.

Natural history. Our data suggest that Pristimantis grandoculis is terrestrial and scansorial. Males were often found perched on low vegetation (0.5-1.5 m above the ground), at dusk, in a position suggesting calling activity. However, acoustic activity has never been observed neither in Suriname nor in French Guiana for P. sp. "Guianas" East (despite being locally abundant). It is noteworthy that both groups of populations also lack a columella and that external tympanum is absent in all specimens examined. Juveniles have been found on the leaf litter during the day, and this is also where juveniles, females and amplectant pairs of P. sp. "Guianas" East have been found in French Guiana suggesting that the reproduction takes place there. The species inhabits pristine terra firme forests from 50 to 700 m asl, generally along the slopes of small massifs.

Distribution. Pristimantis grandoculis occurs throughout the interior of Suriname (not along the coastal band), on the southern border of Guyana with Brazil (Acari mountains), and along the border between the Brazilian states of Pará and Amapá. The species probably extends further in southern Suriname and northern Pará state, Brazil. Phylogenetically related populations occur in French Guiana (Pristimantis sp. "Guianas" East), the Maroni River separating these two groups of genetically divergent populations. The taxonomic status of this group of populations needs to be clarified when additional material becomes available. In French Guiana, P. sp. "Guianas" East occurs throughout the territory except the Approuague-Oyapock interfluvium and is probably absent from northern Amapá state in Brazil. This pattern of distribution strikingly mirrors that of Anomaloglossus surinamensis [START_REF] Vacher | Cryptic diversity in Amazonian frogs: Integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield[END_REF] and Pipa aspera [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF], both species displaying similar geographical boundaries and deep genetic structures along the Maroni River. Interestingly, Pristimantis sp. "Guianas" East only overlaps with P. cf. marmoratus (Pristimantis sp. 1) in the southern half of French Guiana. Likewise, the distribution of A. surinamensis only weakly overlaps with that of A. blanci [START_REF] Fouquet | On the brink of extinction: Two new species of Anomaloglossus from French Guiana and amended definition of Anomaloglossus degranvillei and A. surinamensis (Anura: Aromobatidae)[END_REF]. Both are pairs of sister species with similar ecology and body size. Paratypes. Twenty-eight specimens (18 males and three females) from French Guiana and the state of Amapá, Brazil: MNHN-RA-2020.0092-0112; MPEG 41819-41825 (Appendix 2).

Etymology. The specific epithet is a latinised adjective referring to the call structure of the new species, which sounds like a rattle (crepitaculum).

Definition. The new species is characterized by the following unique combination of characters: (1) SVL small, adult males 17.0 ± 0.8 mm (range 15.7-18.3 mm, n = 11) and adult females 23.4 ± 0.5 mm (range 22.9-23.9 mm, n = 3) (Table 1) Morphological comparisons with other Guiana Shield lowlands Pristimantis of the "unistrigatus group". Pristimantis crepitaculus sp. nov. can mainly be distinguished from P. espedeus by its smaller body size (SVL range in males = 15.7-18.3 mm vs. 20.7-24.8 in P. espedeus); dark grey groin colouration in life (vs. reddish in P. espedeus); and advertisement call composed of more notes and higher dominant frequency (> 6 notes and > 3.1 kHz).

Pristimantis crepitaculus sp. nov. can mainly be distinguished from P. inguinalis by dark grey groin colouration in life (vs. bright yellow inguinal mark in P. inguinalis); light grey ventral colouration with small black spots (vs. entirely black in P. inguinalis); and advertisement call composed of series of notes (vs a single note in P. inguinalis).

Pristimantis crepitaculus sp. nov. can mainly be distinguished from P. grandoculis by visible tympanum (external tympanum absent in P. grandoculis); more tuberculate dorsal skin; enlarged tubercles on eyelids being equally distant from the eye (vs. posterior tubercles closer to the eye in P. grandoculis); fingers and toes without fringes (fringed in P. grandoculis), smaller tibia length (48-54% of SVL vs. 58-60%); and presence of vocal slits in males (absent in P. grandoculis).

All the other species of the unistrigatus group known from the Guiana Shield occur in the Pantepui region and are generally associated with highlands (i.e. > 700 m elevation; [START_REF] Fouquet | A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately elevated massifs of French Guiana[END_REF][START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF]. One of these Pantepui species, P. marmoratus, is possibly its closest relative. Pristimantis crepitaculus sp. nov. can mainly be distinguished from P. marmoratus by its fingers and toes without fringes (fringes present in P. marmoratus) and its advertisement call composed of a series of notes (vs. a single note in P. marmoratus). Among the other Pristimantis species found at mid-elevation in Pantepui, P. crepitaculus sp. nov. can mainly be distinguished from P. jester [START_REF] Means | Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America[END_REF] by the presence of a tympanum (absent in P. jester), from P. saltissimus [START_REF] Means | Three new malodorous rainfrogs of the genus Pristimantis (Anura: Brachycephalidae) from the Wokomung Massif in west-central Guyana, South America[END_REF] by iris colour in life (black vertical streak running across the iris in P. crepitaculus vs. no such streak in P. saltissimus), from P. guaiquinimensis [START_REF] Schlüter | Three new frogs of the genus Eleutherodactylus (Amphibia, Leptodactylidae) from Guaiquinima table mountain, Bolívar. Venezuela[END_REF] vs. 6-12 notes in P. crepitaculus sp. nov.), from P. pulvinatus [START_REF] Rivero | A new species of Eleutherodactylus (Amphibia, Salientia) from the Guayana Region, Edo. Bolívar. Venezuela[END_REF]) by its smaller body size (males 23.0-26.1 mm in P. pulvinatus vs. 15.7-18.3 mm in P. crepitaculus sp. nov.) and distinct vomerine teeth (indistinct or absent in P. pulvinatus), from P. memorans (Myers & Donnelly, 1997) by iris colour in life (black vertical streak running across the iris in P. crepitaculus sp. nov. vs. no such streak in P. memorans), a shorter inter-note interval (0.05-0.12 s in P. crepitaculus sp. nov. vs. 0.20-0.29 s in P. memorans), and by the posterior thigh surface being dark grey in life (vs. blackish with yellow flecking in P. memorans). Description of the Holotype. An adult male, SVL 15.8 mm. Head only slightly longer than wide, widest at corner of the mouth; snout slightly acuminate in dorsal and round in lateral views; canthus rostralis distinctly concave; nostrils nearly elliptical, directed almost completely laterally; interorbital region flat, loreal area slightly concave. Eyes large and protuberant, directed laterally, diameter of eyes much larger than tympanum diameter (ED/TD = 3.0), pupil elliptical. Supratympanic fold slightly distinct; tympanum small, ovoid, tympanic membrane completely covered by skin, but tympanic annulus distinct anteriorly and ventrally, covered by the M. depressor mandibulae posteriorly and dorsally. Vocal sac present, single, subgular, extending towards pectoral region between forearms. Choanae small (0.5 mm), round, not concealed by palatal shelf, larger than vomerine odontophores; a pair of small vomerine odontophores present; tongue ovoid, posteriorly free; vocal slits present, extending diagonally from lateral base of tongue to almost to the angle of the jaw.

Arms slender, not hypertrophied; lateral margins of arm and forearm free of fringes, folds, but with three tubercles; finger discs elliptical, expanded in all fingers, disc on FI the smallest, on FIII the largest; relative lengths of fingers I < II < IV < III; subarticular tubercles round, narrower than finger width; fringes absent on all fingers. Subarticular tubercles present on all fingers, one on FI and FI, two on FIII and FIV; all subarticular tubercles round and about the same size; inner metacarpal (thenar) tubercle present, oval; outer metacarpal tubercle large, W-shaped, ill-defined. Webbing between fingers absent.

Legs relatively long and slender, with rows of large and flat tubercles on tarsus and postaxially on foot (but lacking fringes or folds). All toes well developed, with expanded, elliptical, medium sized discs; discs on TIV and TV the largest, those on TI and TII the smallest; relative lengths of toes I < II < III < V < IV. Toes lacking fringes. Subarticular tubercles present on all toes; one tubercle present on TI, TII, and TV, two tubercles on TIII, three on TIV; all subarticular tubercles round, similar in size. Inner metatarsal tubercle oval; outer metatarsal tubercle small, round.

Skin on dorsum, head, dorsal surfaces of limbs, flanks, and groin tuberculate with enlarged tubercles on the scapular region and on eyelids, posterior surface of the dorsum, and dorsal surface of hindlimbs. Skin on the gular region smooth; chest, belly and undersurfaces of limbs granular. Cloacal opening directed posteriorly; cloacal region lacking tubercles.

Colour of holotype in life. Dorsal colour dark brown with V-shaped black band on the scapular region, a large middorsal black blotch and an ill-defined black blotch near groin (Fig. 5). Black bands and blotches located in the interorbital region, on the loreal region, the nasal region, below the eyes and the tympanic region. Interorbital region and V-shaped scapular mark delimited anteriorly by reddish bands. Lower flank as dorsum with a black blotch. Axillary region and upper arm yellow. Lower arm as dorsum with two black transversal bands. Dorsal colouration of legs as dorsum with four black transversal bars on thigh, four on tibia and three on tarsus. Throat background grey covered with minute black melanophores; belly skin grey covered with small cream spots and by black melanophores that become sparse near groin; ventral surfaces of thighs and arms as throat. Iris with copper metallic pigmentation and pupil ring interrupted dorsally and ventrally by transversal pigmentation (Fig. 5).

Colour of holotype in preservative. After four years in 70% ethanol, colours of the specimen faded, notably glandular supracarpal pad and throat pigmentation (Fig. 5). The snout is dorsally dark brown and well-delimited by a transversal light grey interorbital band extending on half of the eyelids. Posteriorly, the interorbital region is covered by a dark brown band. A V-shaped dark brown mark is welldefined on the scapular region posteriorly followed by two oblique traversal dark brown bands. Similar, transversal dark brown bands are present on the arms, hands and legs and feet (for more details see Fig. 5a).

Variation in the type series. Proportions vary little among the 21 paratypes. Males are smaller than females (15.7-18.3 mm vs. 22.9-23.9). The amount of tuberculation on the skin varies, especially on the dorsum and flanks. Outer metatarsal tubercle is always present, but its size/visibility vary among specimens.

Colour pattern is highly variable among specimens and several patterns are Osteology of the holotype (MNHN-RA-2020.0102). Cranium (Fig. 6). Shape and proportions. The skull is widest posterior to the orbit at the level of the articulation of the maxilla with the quadratojugal. The rostrum is moderate, the braincase is broad. A. Fouquet et al.

3

Neurocranium and dorsal investing bones. The nasals are broadly separated from one another and from the sphenethmoid; they cover most of the nasal capsules dorsally. The frontoparietals are well developed, co-ossified with the sphenethmoid. Frontoparietal crests are absent and the frontoparietal fontanelle (T-shaped) is exposed. The parasphenoid and neopalatines are co-ossified with the sphenethmoid. The frontoparietal and prootic are fused. Ventrally, the prootics are fused with the parasphenoid alae. The exoccipitals are fused. The dorsal surface of the otic capsule is mostly ossified. The septomaxilla is roughly spiralled, the medial ramus extending posterodorsal to the posterior ramus; the anterior ramus is thick; the lateral ramus is oblique with a long acuminate posterolateral extension; the posterior ramus extends from the middle of the lateral ramus ventromedially. The columellae (stapes) are well ossified, formed by the synostotic fusion of the long, thin pars media plectri (stylus) and the pars interna plectri (baseplate), which is curved.

Ventral investing bones. The parasphenoid cultriform process extends anteriorly from the anterior edge of the otic capsule and is co-ossified with the sphenethmoid. The parasphenoid alae are moderately long (about half of the length of the cultriform process), perpendicular to the anteroposterior body axis, broadening slightly laterally. The vomers are in narrow contact with the sphenethmoid; each vomer is composed of an arcuate bone bordering the anteromedial, medial, and posterior margins of the choana. The prechoanal ramus is expanded medially and anteriorly and bears a ventral flange along its medial edge. The postchoanal process is narrow and acuminate, slightly anteriorly curved. Welldeveloped dentigerous processes extend posteromedially from the union of the pre-and postchoanal processes, left process bears two small odontophores. Each dentigerous process is broadly separated from its counterpart medially. The neopalatine is narrowly in contact with the maxilla distally. This complex is fused with the parasphenoid medially.

Maxillary arcade. The maxillary arcade bears many small teeth on the premaxilla and maxilla. The arcade is complete and connected to the slender quadratojugal. The premaxillae are separated medially, and their anterodorsal alary process is weakly divergent from the midline. The pars palatina is broad, with two well-defined processes: the medial (palatine) process is relatively narrow and runs roughly parallel toward its contralateral; the lateral process is broader. The premaxilla and maxilla are in lateral contact via a simple juxtaposition. The maxilla is long, with a broad pars palatina along its lingual margin and a moderately developed pars facialis.

Suspensory apparatus. The triradiate pterygoid bears a slightly curved anterior ramus with a sculpted ventrolateral face, oriented anterolaterally toward the maxilla, with which it articulates at approximately the mid-length of the orbit. This pterygoid is fused to the maxilla. The medial and posterior face of the medial rami of the pterygoid are about equal in length. The medial ramus is broader than the posterior and its posterior face is strongly sculpted. The lateral end of the medial ramus overlaps the lateral edge of the prootic. The quadratojugal is long, laterally curved, and slender, articulating anteriorly with the maxilla. It has a bulbous posteroventral process and articulates dorsally with the ventral ramus of the squamosal. The squamosal is dorsally bifurcated, broad, and sculpted, extending anterodorsomedially from the quadratojugal to the level of the otic capsule; the zygomatic ramus is very short, whereas the otic ramus is long, almost reaching the posterior end of the skull.

Mandible. The mandible is slim and edentate. The mentomeckelians are small and arcuate in ventral view, medially and laterally broadened, and medially separated. The dentary is short and thin, posteriorly acuminate, and overlaps the angulosplenial for about a quarter of its length. Dentary is not in contact with the angulosplenial posteriorly. The angulosplenial is long and arcuate, laterally slightly grooved. The coronoid process is weak.

Hyoid. The bony posteromedial processes of the hyoid are expanded proximally and separated from one another. No ossified parahyoid is present.

Axial skeleton (Fig. 6).

Vertebral column. The vertebral column has eight procoelous, presacral vertebrae. The vertebrae have complete neural arches and low neural processes. Atlas (presacral I) and presacral II are fused medially and laterally. The transverse processes of presacrals II-III are slightly expanded distally, thicker, and broader than those of other presacrals. The transverse processes of presacrals II and III are oriented ventrolaterally (II anteriorly, III slightly posteriorly), whereas those of presacrals IV-VIII extend dorsolaterally (IV-V posteriorly, VI-VIII anteriorly). The relative lengths of the transverse processes and sacral diapophyses are: Sacrum > III > IV > V ≈ VI ≈ VII ≈ VIII > II. The sacral diapophyses are slightly expanded distally. The urostyle has a well-developed dorsal ridge that extends along most of its length.

Pectoral girdle. The zonal portion has well-ossified coracoids, clavicles, scapulae, and cleithra. The clavicles are long, slender, and oriented anteromedially; the medial tips are in contact and located anteriorly from the level of anterolateral end of the clavicle that articulates with the scapula; the coracoid is long and flared, with its sternal end slightly broader than its glenoid end. The coracoids overlap medially. The scapula is long with a prominent pars acromialis that is not separated from the pars glenoidalis. The scapula is about one and a half the length of the clavicle. The cleithrum is ossified, well-developed, anteriorly thicker, thinning posteriorly. The suprascapula is unossified.

Pelvis girdle. The long, slender iliac shafts bear dorsolateral crests throughout their length. The overall length of the Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

girdle is more than two times the width between the anterior ends of the iliac shafts. The iliac prominence is broad and low, and pubes mineralized.

Manus and pes. The phalangeal formulae for the hand and foot are standard, 2-2-3-3 and 2-2-3-4-3, respectively. Terminal phalanges strongly T-shaped. Ossified prepollex and prehallux visible.

Advertisement call. Six specimens calling from the underwood vegetation were recorded from about 2 m away at air temperatures between 23-25 °C and 90-100% relative humidity. Descriptive statistics of call parameters are presented in Table 2. Pristimantis crepitaculus sp. nov. emits every 7.04 s on average (range 5.44-8.62) series (call length mean = 0.59, range 0.50-0.75 s) of 6-12 short notes (note length mean = 0.004 s; range 0.003-0.006 s). These notes are emitted with increasing inter-note intervals within the call (mean 1 st interval = 0.068 s; range 0.054-0.085 s; mean 2nd interval = 0.102 s; range 0.078-0.125 s). The notes have a clear harmonic structure (i.e. with extensive amount of energy in the harmonics). The dominant frequency is 3.50 kHz on average (range 3.16-3.91 kHz) with a slight upward modulation within the call (ca. 0.2 kHz) (Fig. 7, Table 2).

Natural history. Pristimantis crepitaculus sp. nov. is a scansorial species. Males were found calling perched on low vegetation (0.5-2.0 m above the ground). Calling activity was limited to a short period (about 1 h) at dawn and early night. Calling, males often adopt a position head down along the trunks, small twigs, or leaves. They form groups of 2-10 males usually spaced by at least 2 m between individuals. Juveniles and females are very rarely found, but at least one female (MPEG 41825) was collected very close to a calling male (MPEG 41824, calling from a leaf at about 2 m above ground). The species inhabits terra firme forests from 50 to 700 m asl and seems to particularly thrive near ecotones, such as tree falls, clear-cuts, roads, rivers, maybe because the penetrating light allows denser understory cover and thus more abundant calling sites.

Distribution. The species is apparently endemic to the easternmost part of the Guiana Shield lowlands, i.e. throughout the Amapá state of Brazil and French Guiana (at the exception of the northwest part of the territory). It occurs in the upper Maroni region, which corresponds to the border with Suriname, and probably extends at least in southeastern Suriname, like other species that are also endemic to the easternmost part of the Guiana Shield (e.g. Amazophrynella teko, Boana dentei, Pristimantis gutturalis). However, it is unlikely that the range of P. crepitaculus sp. nov. extends to the Coppename river.

Discussion

Species diversity, endemism, and biogeography of the Eastern Guiana Shield

As a further step from [START_REF] Kok | Amended diagnosis and new description of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with description of its advertisement call and notes on its breeding ecology and phylogenetic relationships[END_REF], the present study clarifies the understanding of the megadiverse genus Pristimantis in the Guiana Shield, and of overall anuran diversity pattern in the region. Many anuran species remain to be described and named in Amazonia and notably in the Guiana Shield region, as suggested by almost all studies exploring particular groups (e.g. [START_REF] Gehara | High levels of diversity uncovered in a widespread nominal taxon: Continental phylogeography of the Neotropical tree frog Dendropsophus minutus[END_REF]Fouquet et al., 2015;[START_REF] Kok | Evolution in the South American "Lost World": Insights from multilocus phylogeography of stefanias (Anura, Hemiphractidae, Stefania)[END_REF][START_REF] Jaramillo | Vastly underestimated species richness of Amazonian salamanders (Plethodontidae: Bolitoglossa) and implications about plethodontid diversification[END_REF] and molecular diversity [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF]. The fact that many Amazonian frog species are circumscribed to particular and often relatively small subregions within Amazonia is progressively unveiling as taxonomic progresses are being made. [START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF] notably identified three bioregions within the Guiana Shield, one in the easternmost part, one in the southern part, and one in the western part. The distributions of P. espedeus, P. crepitaculus, P. sp. "Guianas" East, and P. grandoculis in the eastern portion, of P. cf. ockendeni in the southern, and of P. marmoratus, P. pulvinatus, and P. saltissimus in the western portions strongly corroborate this pattern. The determining factors of this pattern remain ambiguous, but probably originate from both historical climatic and landscape changes and current bioclimatic heterogeneity, notably the fact that the easternmost part of the Guiana notably receives more precipitation seasonally than other parts of the region (Silva et al., 2018;[START_REF] Vacher | Large scale DNA-based survey of frogs in Amazonia suggests a vast underestimation of species richness and endemism[END_REF] (Table 3).

A few lineages remain to be investigated for a better understanding of their taxonomic status, such as Pristimantis sp. "Guianas" East and the five lineages forming what we call a "trans-amazon complex" (P. cf. ockendeni). These candidate species require a thorough and integrative systematic review. Although P. grandoculis and P. sp. "Guianas" East are morphologically very similar (the latter also lacking a columella and external tympanum, see Fig. 8), the degree of divergence on the 16S fragment that we used is > 4% suggesting that these groups of populations may correspond to distinct species [START_REF] Fouquet | Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses[END_REF][START_REF] Vieites | Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory[END_REF]. However, the lack of acoustic data and the low number of available specimens from Suriname so far hamper any taxonomic decision about P. sp. "Guianas" East. Pristimantis diversity in the Pantepui region also requires further investigation, as notably illustrated by candidate species from La Neblina, Mount Ayanganna, Mount Wokomung (B. Means, pers. comm.). Probably many more yet undocumented isolated lineages occur in poorly explored areas of Pantepui, including isolated tepui summits. A larger-scaled phylogenetic investigation is needed to test whether a Guiana Shield clade of Pristimantis might have diversified in situ. A few groups have been documented to have extensively diversified throughout the Guiana Shield lowlands and highlands such as Anomaloglossus [START_REF] Vacher | Cryptic diversity in Amazonian frogs: Integrative taxonomy of the genus Anomaloglossus (Amphibia: Anura: Aromobatidae) reveals a unique case of diversification within the Guiana Shield[END_REF], and Otophryne [START_REF] Fouquet | Species diversity and biogeography of an ancient frog clade from the Guiana Shield (Anura: Microhylidae: Adelastes, Otophryne, Synapturanus) exhibiting spectacular phenotypic diversification[END_REF]. These cases are nonetheless relatively rare since most lowland amphibians have diversified throughout Amazonia. The Pristimantis species belonging to the "unistrigatus group" could represent an additional case of Guiana Shield restricted clade, whose monophyly and timing of diversification would be worth investigating.

The case of the "trans-amazon complex" (P. cf. ockendeni) is definitively no less complex. It is most certainly a lineage apart from the other species of the Guiana Shield that most likely originated from western Amazonian ancestors and secondarily dispersed to the Guiana Shield across the Amazon River. Similar and possibly concomitant trans-Amazon dispersals are documented in Allobates (A. aff. tapajos [START_REF] Réjaud | Historical biogeography identifies a possible role of the Pebas system in the diversification of the Amazonian rocket frogs (Aromobatidae: Allobates)[END_REF] and Boana (Boana gr. albopunctata [START_REF] Fouquet | Species diversity and biogeography of an ancient frog clade from the Guiana Shield (Anura: Microhylidae: Adelastes, Otophryne, Synapturanus) exhibiting spectacular phenotypic diversification[END_REF], Synapturanus [START_REF] Fouquet | Species diversity and biogeography of an ancient frog clade from the Guiana Shield (Anura: Microhylidae: Adelastes, Otophryne, Synapturanus) exhibiting spectacular phenotypic diversification[END_REF], Scinax [START_REF] Ferrão | High species richness of Scinax treefrogs (Hylidae) in a threatened Amazonian landscape revealed by an integrative approach[END_REF]. From 9 million years ago (Ma) onwards, the Amazon basin has been draining eastward into the Atlantic Ocean and the Amazon River was forming a barrier for South-North dispersal around 1 3

Osteological comparisons

At this stage, comprehensive osteological comparisons among species in the "unistrigatus group" are impossible due to the paucity of data, as the osteology of most of these species has not been described or illustrated. However, our osteological investigation of the holotypes of P. marmoratus and P. grandoculis unambiguously confirms the nonconspecificity of these taxa. The main diagnostic osteological characters are the lack of a columella and a more ossified skull in P. grandoculis (Figs. 4,6 and 8). The skull of MNHN-RA-2020.0118 is, however, poorly ossified, which could be an artefact of preservation as suggested by the lack of ossification of the brain case for instance. The skull of the holotype of P. grandoculis, although that specimen was collected in 1904, is still strongly ossified. Unfortunately, further comparisons with the holotype of P. marmoratus are made difficult by the poor quality of the µ-CT scans, and we refrain to describe its osteology until more µ-CT scans of additional specimens of P. marmoratus sensu stricto are available. Spatial distribution, external morphology, and osteological data converge in suggesting the conspecificity of the populations previously identified as P. sp. "Guianas" from Suriname and P. grandoculis. The absence of a columella in P. grandoculis is in line with the absence of vocal slit, pharyngeal ostia, and calling activity in that species (see below). The sister species (population from French Guiana) also lacks a columella and external tympanum. A complete investigation of the osteology of the "unistrigatus group" would certainly be of taxonomic/evolutionary interest.

Vertical niche partitioning among species and related traits on limbs

Juveniles, females, and amplectant pairs of Pristimantis sp. "Guianas" East (sister to P. grandoculis) are often found on the leaf litter, suggesting that the reproduction takes place directly either on the ground or at low height in the vegetation. In contrast, females of P. crepitaculus, P. espedeus, P. inguinalis and P. cf. ockendeni are rarely seen. Males of Pristimantis sp. "Guianas" East are also found on the ground but more frequently perched at low height in the underwood at night. However, juveniles of P. espedeus are also frequently seen on the ground as are juveniles of Pristimantis sp. "Guianas" East, but during the day. Small-bodied, terrestrial, and nocturnal anurans are very rare [START_REF] Pough | Morphology, physiology, and foraging behaviour[END_REF], possibly because predators such as spiders are very abundant in the leaf litter at night [START_REF] Magnusson | The fish and the frogs[END_REF]. Among the species of the "unistrigatus group" occurring in the Guiana Shield lowlands, Pristimantis grandoculis and P. sp. "Guianas" East can be considered as the lowest in terms of vertical partitioning. Vertical niche partitioning in Pristimantis has already been described notably in western Ecuador [START_REF] Lynch | Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador: Systematic, ecology, and biogeography[END_REF]Guyasamin & Funk, 2009) and in Pristimantis, Craugastor, and Diasporus in Costa Rica [START_REF] Miyamoto | Vertical habitat use by Eleutherodactylus frogs (Leptodactylidae) at two Costa Rican localities[END_REF]. These converging observations strengthen the idea that vertical partitioning among Pristimantis species is frequent, even at low elevations such as in the Amazonian lowlands, and may even play a direct role in speciation.

It is noteworthy that Pristimantis grandoculis and P. sp. "Guianas" East have longer limbs (TL, TiL, FL) and more terrestrial habits than scansorial P. crepitaculus, P. cf. ockendeni and arboreal P. inguinalis and P. espedeus, species. These differences in limb proportions may be related to jumping ability [START_REF] Citadini | Evolution of morphology and locomotor performance in anurans: Relationships with microhabitat diversification[END_REF][START_REF] Emerson | Skull shape in frogs: Correlations with diet[END_REF][START_REF] Zug | Anuran locomotion: Structure and function. I. Preliminary observations on relation between jumping and osteometrics of appendicular and postaxial skeleton[END_REF]. We hypothesize that long distance jumping performance may be selected in a leaf litter and scansorial environment which is relatively free of obstacles and that may be risky because predators are abundant in the leaf litter [START_REF] Pough | Morphology, physiology, and foraging behaviour[END_REF], whereas scansorial and arboreal species require less power but more agility in dense vegetation.

Lack of ear and of calling activity

Males of P. grandoculis and P. sp. "Guianas" East were often found perched on the vegetation in a position suggesting calling activity. However, acoustic activity has never been heard in that clade, neither in Suriname nor in French Guiana (despite being locally abundant), and these are the only species in the Guiana Shield lowlands with unknown call. Moreover, tympanum, columella, pharyngeal ostia, and vocal slits are all absent in that clade. These observations suggest that P. grandoculis may be both deaf and mute. The loss of hearing structures has occurred multiple times throughout the evolutionary history of anurans (Jaslow et al., 1988;[START_REF] Boistel | How minute sooglossid frogs hear without a middle ear[END_REF]Pereyra et al., 2016). It is particularly common in Bufonidae with 30% of the species being earless (Pereyra et al., 2016). It is also common in Terrarana [START_REF] Hedges | New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation[END_REF]Padial et al., 2014[START_REF] Goutte | Evidence of auditory insensitivity to vocalization frequencies in two frogs[END_REF][START_REF] Von May | Evolutionary radiation of earless frogs in the Andes: Molecular phylogenetics and habitat shifts in high-elevation terrestrial breeding frogs[END_REF] and is particularly prevalent in several clades distributed at high elevations (e.g. Bryophryne, Phrynopus; [START_REF] Duellman | Terrestrial-breeding frogs (Strabomantidae) in Peru[END_REF]. Only a few Pristimantis of the "unistrigatus group" have been reported to lack tympanum and vocal slits (e.g. P. imthurni and P. jamescameroni, see Kok, 2013; P. yaviensis, see Myers & Donnelly, 1996). However, these Pristimantis do call. As a matter of fact, most earless frogs, although anatomically deaf, still call and can perceive acoustic signals via other organs (e.g. [START_REF] Boistel | How minute sooglossid frogs hear without a middle ear[END_REF][START_REF] Goutte | Evidence of auditory insensitivity to vocalization frequencies in two frogs[END_REF]. But only a few frog species have been reported to be mute in addition to be deaf. Most of these frogs are living near noisy environments such as streams and advertise via other cues such as visual or chemical signalling [START_REF] Toledo | The anuran calling repertoire in the light of social context[END_REF]. The cases of a dull-coloured Amazonian lowland species such as P. grandoculis being deaf and mute are remarkable. Hypotheses regarding the loss of ear and of acoustic activity in this species remain open. Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3 Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

Fig. 1 a

 1 Fig. 1 a Maximum Likelihood phylogenetic tree obtained from the analysis of 191 sequences of 12S-16S (2548 bp) of Pristimantis. Bootstraps > 50% are indicated on the left side of the nodes. For sake of clarity, branches within species are collapsed. Only the species occurring in the lowlands are coloured in the tree according to the map. b Distribution of the different species included in the molecular dataset. The range of the species occurring in the lowlands is depicted by filled polygons (except P. espedeus and P. inguinalis that are not

  Fouquet, S. Cally and R. Jairam on 30 April 2015 at Bakhuis Mountains, Suriname (4.7246°N 56.7638°W, ~ 200 m asl; Fig. 3). NZCS A1212, an adult male collected by P. Ouboter and V. Kadosoe on 20 August 2013 at Tafelberg, Suriname (3.8049°N 56.1539°W ~ 500 m asl); NZCS A1210-11, two adult males collected by R. Jairam and D. Baêta on 23 June 2016 at Lely Mountain, Suriname (4.4158°N 54.6498°W, ~ 600 m asl).

  deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3 subarticular tubercle of FII; (13) fingers with preaxial fringes on FII and III; (14) finger discs broadly expanded, elliptical, thenar tubercle oval, palmar often broken in three distinct tubercles; (15) three enlarged ulnar tubercles often flat and barely visible; (16) axillary tubercles (sensu Myers & Donnelly, 2001) absent; (17) tarsal fold absent but tubercles present; (18) toes basally webbed between TII-IV with preaxial fringes on TII-V; (19) calcars absent, inner metatarsal tubercle oval, outer metatarsal tubercle round; (20) dorsal colouration mostly chestnut brown, but highly variable in colour and pattern, ventral colouration translucent brownish grey with small cream spots, mostly on chest, throat suffused with black melanophores and scattered with small irregular cream blotches; (21) iris golden with a copper horizontal band, absence of vertical streak; (22) posterior surface of thighs and groin dark grey in life and brown in preservative, absence of yellow spot on groin.

Fig. 3 a

 3 Fig. 3 a Dorsal, ventral, and lateral (head) views of the holotype (RMNH 4467) of P. grandoculis from Coppename River, Suriname. b Dorsal and ventral views and hand and foot of a recently collected

Fig. 4

 4 Fig. 4 Volumetric renderings of µ-CT scans of the skeleton of the holotype of Pristimantis grandoculis (RMNH 4467). a Full skeleton in dorsal (left), ventral (middle), and lateral (right) views. b Skull in dorsal (left), ventral (middle), and lateral (right) views. Abbreviations: ang-angulosplenial, den-dentary, dp-dentigerous process, exo-

  Pristimantis crepitaculus sp. nov. Pristimantis marmoratus Lescure & Marty, 2000 Pristimantis marmoratus Fouquet et al., 2007 Pristimantis marmoratus Dewynter et al., 2008 Pristimantis marmoratus Fouquet et al., 2012 Pristimantis sp. 1 Fouquet et al., 2013 Pristimantis sp. 1 Kok et al., 2018 Pristimantis sp. 1 Fouquet et al., 2019 Pristimantis cf. marmoratus Vacher et al., 2020 Holotype. MNHN-RA-2020.0102 (field no. AF2786), an adult male, collected by Antoine Fouquet, Maël Dewynter & Nicolas Vidal on 26 February 2015 at Mitaraka, French Guiana (2.2358 N 54.4493 W, ~ 200 m asl; Fig. 5).

  ;(2) dorsal skin tuberculate, two pairs of enlarged tubercles on the scapular region embedded in a W-shaped scapular fold, ventral skin granular particularly on the femoral region; (3) tympanum visible, tympanic membrane not or only poorly differentiated, tympanic annulus partially visible externally obscured by supratympanic fold on the posterodorsal edge of tympanum, tympanum c.a. 1/3 of eye length; (4) pharyngeal ostia present; (5) columella present; (6) tibia length 48-54% of SVL; (7) snout broadly rounded in profile and slightly acuminate in dorsal view; (8) each upper eyelid with two prominent tubercles; (9) choanae round and small (0.5 mm for the holotype), dentigerous processes of vomers oblique, narrowly separated, each bearing 2-4 small odontophores; (10) vocal slits present, vocal sac median, subgular; (11) one unpigmented whitish nuptial pad located on the preaxial side of the thenar tubercle on each thumb in male; (12) FI slightly shorter than FII, reaching disc of FII; (13) fingers without lateral fringes; (14) finger discs broadly expanded, elliptical, thenar tubercle ovoid, palmar tubercle ill-defined; (15) three enlarged ulnar tubercles often flat and barely visible; (16) axillary tubercles (sensu Myers & Donnelly, 2001) absent; (17) small tarsal tubercles present; (18) toes without lateral fringes but with weak lateral keels, webbing rudimentary basal between TII-V; (19) calcars absent, inner metatarsal tubercle oval, much larger than the round outer tubercle; (20) dorsal colouration highly variable from greenish to brownish with various patterns, ventral colouration light grey with small black spots denser on throat and legs; (21) iris reddish to copper with a darker horizontal band and a black vertical streak; (22) posterior surface of thighs and groin dark grey in life and brown in preservative, lack of yellow circumscribed spot on groin; (23) anterior surface of arms, posterior surface of flanks and dorsal surface of thighs often yellow in reproductive males; (24) advertisement call characterized by series (0.50-0.75 s long) of 6-12 very short notes (0.02-0.05 s long) with a dominant frequency ranging between 3.2-3.6 kHz and emitted every 5.4-8.6 s; (25) male calling activity exclusively crepuscular, usually upside down on tree trunks of low diameter.

3 Fig. 5 a

 35 Fig. 5 a Holotype of Pristimantis crepitaculus sp. nov. in dorsal, ventral views. b Hand, foot and lateral view of the head. c Photographs of the holotype in life. d Several photographs of additional specimens in life

  observed. A pattern similar to the holotype (middorsal and interobital bands) is present in female MNHN-RA-2020.0104 and in males MNHN-RA-2020.0110 and MPEG 41821. A white band between the anterior corner of the eyes is present in most individuals but some specimens have less conspicuous markings (e.g. MNHN-RA-2020.0102, MNHN-RA-2020.0107). The scapular V-shaped mark is light grey in a few specimens (MNHN-RA-2020.0101, MNHN-RA-2020.094) or reddish (MNHN-RA-2020.0102, MNHN-RA-2020.0109) and display brightly coloured upper arms and posterior portions of flanks in life. Two specimens (MNHN-RA-2020.0098 and deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3 MPEG 41825) have a large well-defined cream spot on top of the snout and another one (MNHN-RA-2020.0100) has a light brown scapular mark posteriorly prolonged along the back forming a Y-shaped pattern delimited by darker flanks. Two specimens, one male (MPEG 41823) and one female (MPEG 41821) have large cream blotches on the dorsum.

Fig. 6

 6 Fig. 6 Volumetric renderings of µ-CT scans of the skeleton of the holotype of Pristimantis crepitaculus (MNHN-RA-2020.0102). a Full skeleton in dorsal (left), ventral (middle), and lateral (right) views. b Skull in dorsal (left), ventral (middle), and lateral (right) views. Columellae are highlighted in red. Abbreviations: ang-angulosplenial,

Fig. 7

 7 Fig.7Sonograms and oscillograms on a 0.7-s window of a characteristic call of Pristimantis crepitaculus sp. nov. from Oiapoque,Amapá, Brazil (paratype, MPEG 41823)

Pristimantis grandoculis (van Lidth de Jeude, 1904)

	Hylodes grandoculis van Lidth de Jeude, 1904
	Eleutherodactylus marmoratus Hoogmoed, in Frost, 1985
	Eleutherodactylus marmoratus Avila-Pires et al., 2010
	Pristimantis marmoratus Ouboter & Jairam, 2012
	Pristimantis sp. 4 Jairam, 2019
	Pristimantis sp. "Guianas" Vacher et al., 2020

Table 1 Morphometric variables SVL TiL FeL TaL FL HeL HW Ind IOD EN ED TD FD 4TD ETS 1FiL 2FiL HL

 1

	4.3	3.9	4.7	5.5	5.1	5.8	3.8	3.6	4.1	5.1	4.7	5.4	4.2	3.7	4.8	5.9	5.3	6.4	4.4	3.8	4.9
	2.4	2.1	2.7	3.5	3.1	4.0	2.2	2.0	2.4	3.0	2.6	3.6	2.4	1.9	2.9	3.2	2.7	3.9	2.5	1.9	3.0
	2.1	1.8	2.5	3.0	2.8	3.2	2.0	1.8	2.2	2.6	2.2	2.9	2.0	1.7	2.4	2.8	2.5	3.2	2.0	1.8	2.4
	3.2	2.9	3.6	4.3	4.0	4.5	3.0	2.8	3.2	4.1	3.9	4.5	3.1	2.8	3.4	4.3	4.1	4.7	3.1	2.8	3.4
	0.9	0.8	1.1	1.3	1.2	1.3	0.8	0.8	1.0	1.2	1.0	1.3	0.8	0.6	0.9	1.1	0.9	1.4	0.8	0.7	0.9
	0.9	0.7	1.0	1.2	1.1	1.2	0.8	0.7	1.0	1.1	1.0	1.2	0.8	0.7	0.9	1.1	0.9	1.4	0.8	0.8	0.9
	0.9	0.7	1.1	1.1	0.9	1.3	0.8	0.7	0.9	1.0	0.8	1.2	NA	NA	NA	NA	NA	NA	NA	NA	NA
	2.5	2.3	2.9	3.1	2.7	3.3	2.6	2.4	2.8	3.2	3.1	3.4	2.3	2.1	2.5	3.2	2.7	3.5	2.5	2.3	2.7
	2.2	1.9	2.6	2.9	2.9	3.0	2.2	2.1	2.3	3.0	2.8	3.2	2.0	1.7	2.5	2.9	2.5	3.4	2.0	1.8	2.2
	2.0	1.8	2.3	2.4	2.4	2.5	2.0	1.8	2.1	2.7	2.6	3.1	2.0	1.7	2.2	2.5	2.1	2.8	1.9	1.9	2.0
	1.6	1.5	1.8	2.0	1.8	2.2	1.7	1.6	1.8	2.1	1.9	2.3	1.7	1.4	1.9	2.1	1.8	2.3	1.8	1.7	1.9
	6.4	5.7	6.9	8.4	7.7	9.0	6.2	5.7	6.7	8.6	7.5	9.6	5.5	4.7	6.5	8.2	7.2	9.0	6.1	5.6	6.4
	6.5	6.0	7.1	8.5	8.3	8.6	6.4	5.9	6.8	8.4	7.6	9.6	5.8	5.3	6.5	8.1	7.4	9.3	6.1	5.5	6.5
	6.8	6.1	7.3	9.1	8.9	9.5	6.3	5.8	6.8	8.2	7.9	8.6	7.1	5.9	8.0	9.6	8.7	10.6	7.0	6.1	7.7
	5.3	4.9	5.9	6.7	6.4	7.0	5.4	5.1	5.9	6.9	6.5	7.3	5.2	4.6	5.9	6.6	6.0	7.3	5.4	4.9	6.4
	8.3	7.5	9.2	10.8	10.3	11.0	7.8	7.1	8.3	10.4	10.0	11.5	8.4	7.2	9.4	11.1	10.5	12.7	8.2	7.3	9.0
	8.8	8.0	9.4	11.4	10.9	11.8	8.6	8.2	9.1	11.4	10.7	12.2	9.1	8.2	10.4	12.0	11.1	13.1	9.5	8.7	10.5
	17.0	15.7	18.3	23.4	22.9	23.9	17.4	16.2	18.4	24.1	21.4	25.7	16.0	14.7	17.5	22.5	20.6	25.0	16.5	14.7	17.9
	M (n = 19) Mean	min	max	F (n = 3) Mean	min	max	M (n = 8) Mean	min	max	F (n = 6) Mean	min	max	M (n = 13) Mean	min	max	F (n = 13) Mean	min	max	M (n = 5) Mean	min	max	F (n = 0)
	P. crepitaculus sp. nov						P. cf. ockendeni						P. sp "Guianas" East						P. grandoculis

Journal : Large 13127 Article No : 564 Pages : 34 MS Code : 564 Dispatch : 18

-5-2022

Table 2

 2 Acoustic variables

			Call length	Inter-call	Note length	Inter-note 1	Inter-note 2	Dom. Freq	Nb notes
	P. crepitaculus sp. nov	Mean (n = 6)	0.586	7.038	0.004	0.068	0.109	3497	7.9
		Min	0.504	5.440	0.003	0.054	0.078	3163	6
		Max	0.752	8.623	0.006	0.085	0.125	3908	12

  Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes…

	lon lon lon	-52.9017 -52.700 -56.7865	-56.7721	-52.1519 -53.2726 -56.7721	-52.1005 -52.3496 -56.7721 -59.9505	-53.2072 -52.34301 -52.3496 -58.6997	-52.1490 -53.4128 -53.9128 -58.6997	-51.8896 -53.7728 -53.9128 -58.6997	-54.3534 -53.2160 -54.4493 -56.1256 -58.6350	-53.2726 -52.8746 -53.2298 -51.9366	-53.2886 -54.6000 -53.2459	-52.5902 -51.6149 -53.7728 -61.1092 -61.1092 -52.0221 -52.5902 -51.1913 -53.7728 -55.7986 -58.6364 -52.6806 -51.1913 -54.6000 -52.0221 -56.1251 -53.1985 -51.6453 -52.3667 -51.6453	-53.4107 -51.6109 -53.5548	-53.4121 -51.7710 -51.8934 -54.4193	-54.4493 -53.4775 -53.2285	-54.4493 -66.0878 -53.2285	-52.1804 -66.0878 -52.5540 -54.6498 -54.6498	-52.5419 -66.0878 -58.6959 -58.6959 -53.0955 -52.7000 -55.1948	-53.2086 -52.7000 -56.1539 -56.1539
	lat lat lat	2.1969 4.0917 4.6562	4.6837	4.5481 3.6359 4.6837	4.5161 4.2927 4.6837 5.3995	3.6256 4.4088 4.2927 5.1553	4.1959 3.7193 3.5444 5.1553	3.8112 2.3709 3.5444 5.1553	2.2534 2.3491 2.2358 2.02682 -4.3442	3.6359 2.2480 3.4696 -2.9506	3.6112 2.3333 4.4723	3.9909 0.9793 2.3709 -4.9136 -4.9136 0.4493 3.9909 2.1271 2.3709 -6.3173 -4.3089 4.0848 2.1271 2.3333 0.4493 2.0244 3.5732 2.32361 4.1833 2.32361	3.0840 2.3216 5.4074	3.0836 3.8794 3.8061 4.4520	2.2358 3.18341 0.9091	2.2358 0.7195 0.9091	3.3129 0.7195 2.6284 4.4158 4.4158	2.6276 0.7195 1.2854 1.2854 3.0230 4.0917 4.9366	3.47363 4.0917 3.8049 3.8049
	12S access 16S access Genus Species Voucher Voucher2 Locality Country 12S access 16S access Genus 12S access 16S access Genus Species Voucher Voucher2 Locality Country Country Voucher2 Locality Voucher Species	NA KR811102 Pristimantis crepitaculus AF0563 Savane Virginie French N C Guiana NA KR811105 Pristimantis crepitaculus AF0757 St Georges French U O Guiana NA KDQF01000316 Pristimantis crepitaculus AF0910 MNHN-RA-2020.0093 Pic Coudreau du Sud French R Guiana NA KDQF01000366 Pristimantis crepitaculus AF1014 Saul French Guiana R NA KDQF01000382 Pristimantis crepitaculus AF1046 Saul French Guiana E C NA XXXXXXXX Pristimantis crepitaculus AF1387 MNHN-RA-2020.0094 Aratai French Guiana NA XXXXXXXX Pristimantis crepitaculus AF1388 MNHN-RA-2020.0095 Aratai French Guiana T E D NA XXXXXXXX Pristimantis crepitaculus AF1598 MNHN-RA-2020.0097 Nouragues French Guiana NA KDQF01000601 Pristimantis crepitaculus AF1607 MNHN-RA-2020.0100 Saul French Guiana NA KDQF01000749 Pristimantis crepitaculus AF1921 MNHN-RA-2020.0101 Flat de la Waki French Guiana NA KDQF01000754 Pristimantis crepitaculus AF1941 Flat de la Waki French P R O Guiana NA KDQF01001103 Pristimantis crepitaculus AF2786 MNHN-RA-2020.0102 Mitaraka French Guiana NA XXXXXXXX Pristimantis crepitaculus AF2799 MNHN-RA-2020.0103 Mitaraka French Guiana NA KDQF01001186 Pristimantis crepitaculus AF3002 Mémora D AP NA KDQF01001205 Pristimantis crepitaculus AF3047 Mitan G French O Guiana NA KDQF01001276 Pristimantis crepitaculus AF3161 MNHN-RA-2020.0108 Mitan D AP F NA KDQF01001437 Pristimantis crepitaculus AF3539 MNHN-RA-2020.0092 Itoupe French Guiana NA XXXXXXXX Pristimantis crepitaculus AF5440 Trois-Saut French Guiana JN690707 EU201063 Pristimantis crepitaculus AG_BM110 Kaw3 French Guiana NA KDQF01001542 Pristimantis crepitaculus AG321 Kaw French Guiana NA KDQF01001568 Pristimantis crepitaculus AG378 Montagne Maripa French Guiana NA KDQF01001645 Pristimantis crepitaculus AG494 Saul French Guiana NA KDQF01001886 Pristimantis crepitaculus BOAM048 Borne 4 French Guiana NA KDQF01002129 Pristimantis crepitaculus CAAM15 Montagne Cacao French Guiana NA EU201062 Pristimantis crepitaculus CM077 Trois-Sauts French Guiana NA KDQF01002243 Pristimantis crepitaculus CM171 Trijonction French Guiana NA KDQF01002504 Pristimantis crepitaculus FL04 Plote PPBIO AP NA KDQF01002521 Pristimantis crepitaculus FTA128 Rio Vila Nova AP NA KDQF01002528 Pristimantis crepitaculus FTA165 Rio Amapá Grande AP NA KDQF01002543 Pristimantis crepitaculus FTA239 Rio Amapá Grande AP NA XXXXXXXX Pristimantis crepitaculus FTA96 Rio Vila Nova AP NA KDQF01003282 Pristimantis crepitaculus MTR13884 Lourenço AP NA KDQF01003293 Pristimantis crepitaculus MTR13930 Lourenço AP NA KR811104 Pristimantis crepitaculus MTR24285 Lourenço AP NA KR811103 Pristimantis crepitaculus MTR24300 Oiapoque AP NA KDQF01003952 Pristimantis crepitaculus PG748 St Georges French Guiana NA KDQF01004462 Pristimantis crepitaculus WAAM011 Flat de la Waki French Guiana NA KDQF01003322 Pristimantis sp. Neblina 1 MTR15532 São Gabriel da Cachoeira AM NA KDQF01003323 Pristimantis sp. Neblina 1 MTR15534 São Gabriel da Cachoeira AM NA KDQF01003325 Pristimantis sp. Neblina 2 MTR15536 São Gabriel da Cachoeira AM NA KDQF01000102 Pristimantis sp. "Guianas" East AF0266 Nouragues French Guiana U N C NA KDQF01000104 Pristimantis NA KDQF01000354 Pristimantis NA XXXXXXXX Pristimantis NA KDQF01000558 Pristimantis NA KDQF01001024 Pristimantis NA XXXXXXXX Pristimantis NA XXXXXXXX Pristimantis NA XXXXXXXX Pristimantis NA XXXXXXXX Pristimantis NA KDQF01001850 Pristimantis NA KDQF01001860 Pristimantis NA KDQF01002249 Pristimantis JN690709 JN691316 Pristimantis NA XXXXXXXX Pristimantis NA XXXXXXXX Pristimantis XXXXXXXX KDQF01004305 Pristimantis NA KDQF01004306 Pristimantis NA XXXXXXXX Pristimantis NA XXXXXXXX Pristimantis NA KDQF01002880 Pristimantis NA KDQF01002881 Pristimantis NA KDQF01003777 Pristimantis NA NA XXXXXXXX Pristimantis crepitaculus AF5399 Camp Saut Richard French Guiana NA KDQF01000103 Pristimantis sp. "Guianas" East AF0268 Nouragues French Guiana KDQF01003795 Pristimantis NA KDQF01003796 Pristimantis NA KDQF01001388 NA KDQF01001400 NA KDQF01001403 NA KDQF01001417 EU186676 EU186676 XXXXXXXX KDQF01004221 XXXXXXXX KDQF01004222 XXXXXXXX KDQF01004223 NA NA NA NA NA NA NA NA XXXXXXXX KDQF01000865 KDQF01003210 KDQF01001829 KDQF01004256 KDQF01004257 KDQF01002705 KDQF01003224 KDQF01000889 KDQF01002605 U Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis Pristimantis N C O O R R E C T E D P R O O F sp. "Guianas" East AF0272 Nouragues French Guiana sp. "Guianas" East AF0978 Saul French Guiana sp. "Guianas" East AF1360 Grande Montagne Tortue French Guiana sp. "Guianas" East AF1505 Grande Montagne Tortue French Guiana sp. "Guianas" East AF2625 Atachi Bakka French Guiana sp. "Guianas" East AF2626 Atachi Bakka French Guiana sp. "Guianas" East AF2818 Mitaraka French Guiana sp. "Guianas" East AF5414 Savane Roche Dachine French Guiana sp. "Guianas" East AF5525 Haute Courcibo French Guiana sp. "Guianas" East BOAM001 Borne 4 French Guiana sp. "Guianas" East BOAM012 Borne 4 French Guiana sp. "Guianas" East CM184 Trijonction French Guiana sp. "Guianas" East CM317 CisameG French Guiana sp. "Guianas" East QM0423 Montagne de fer French Guiana sp. "Guianas" East QM1073 Gaa Kaba French Guiana sp. "Guianas" East ST310 Parna Tumucumaque AP sp. "Guianas" East ST311 Parna Tumucumaque AP grandoculis UFRJA024 NZCS A1211 Lelygebergte Suriname grandoculis UFRJA023 NZCS A1210 Lelygebergte Suriname grandoculis MPEG30085 Serra do Acari PA grandoculis MPEG30088 Serra do Acari PA grandoculis NZCS50 Brownsberg Suriname grandoculis NZCS A1019 Tafelberg Suriname grandoculis NZCS A1212 Tafelberg Suriname R P R O O Suriname Suriname Suriname Suriname Guyana Guyana Guyana Guyana Suriname AM PA AM AM PA AM Suriname F Bakhuis Bakhuis Bakhuis Mt. Ayanganna Mabura Hill forest reserve Mabura Hill forest reserve Mabura Hill forest reserve Sipaliwini Igarapé Açu, Rio Abacaxis UHE Belo Monte, Vitória do Xingu, Rio Xingu Campo Catuquira Campo Catuquira MINA PALITO São Sebastião, Rio Abacaxis Sipaliwini MNHN-RA-2020.0118 Bakhuis ROM40164 SMNS11989 SMNS11990 SMNS11994 T E D AF3365 AF3402 AF3408 AF3435 AF2155 MTR12690 BM153 SMS155 SMS156 JOG847 MTR12855 AF2257 E C R grandoculis grandoculis grandoculis grandoculis sp. Ayangana cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni cf. ockendeni
	1 3 1 3															1 3

Journal : Large 13127 Article No : 564 Pages : 34 MS Code : 564 Dispatch : 18-5-2022 A. Fouquet et al. Journal : Large 13127 Article No : 564 Pages : 34 MS Code : 564 Dispatch : 18-5-2022 Journal : Large 13127 Article No : 564 Pages : 34 MS Code : 564 Dispatch : 18-5-2022 A. Fouquet et al.

Acknowledgements

We thank Ate Alma-Cohen and Esther Dondorp from the Leiden Naturalis Biodiversity Center (National Research Institute for Biodiversity, The Netherlands), and Mark Wilkinson from the Natural History Museum (UK) who provided the µ-CT scans of the holotype of Hylodes grandoculis and the holotype of Pristimantis marmoratus, respectively. Renaud Lebrun and Quentin Martinez (ISEM) helped with data acquisition with the µ-CT facilities of the MRI platform member of the National Infrastructure France-BioImaging (supported by the French National Research Agency (ANR-10-INBS-04, «Investments for the future»), the Labex CEMEB (ANR-10-LABX-0004), and NUMEV (ANR-10-LABX-0020)). We also thank E. Courtois, L. Brandão, M. Dewynter, N. Vidal, J. Devillechabrole, J.-P. Vacher, P. Nunes, B. Villette, S. Cally, D. Baêta, P. Ouboter, V. Kadosoe, M. Blanc, J. Dias Lima, P. Gaucher, M.T. Rodrigues, and V. Rufray who helped to collect some of the specimens and tissue samples used in this work. Permission to conduct biodiversity research in Guyana was provided by the Environmental Protection Agency of Guyana under research permit number 180609 BR 112, and fieldwork was made possible through the Forestry Department of the Guyana Forestry Commission (GFC-PRDD), particularly R. Thomas. Collection permits for Brazil were provided by Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio).

Funding This study benefited from an "Investissement d'Avenir" grant managed by the Agence Nationale de la Recherche (CEBA, rANR-10-LABX-25-01). PP and ATM were supported by fellowships from Conselho Nacional de Desenvolvimento Científico e Tecnológico (302501/2019-3 and 300822/2021-9 to PP; 142153/2019-2 to ATM). PP was also supported by an award from the Maxwell/Hanrahan Foundation. The work of RE was funded by the German Academic Exchange Service (DAAD) and Deutsche Forschungsgemeinschaft (DFG; ER 589/2-1). The work of PJRK was funded by the Fonds voor Wetenschappelijk Onderzoek (FWO12A7614N and FWO12A7617N).

Availability of data and material

All new DNA sequences, call recordings, and µ-CT scans will be deposited upon manuscript acceptance.

Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes… 1 3

5 Ma. The processes that may have fostered multiple trans-Amazon dispersals around 5 Ma remain highly speculative.

Considerable uncertainty remains about the timing and amplitude of historical topographic, hydrological, and vegetational changes in Amazonia [START_REF] Albert | The changing course of the Amazon River in the Neogene: Center stage for Neotropical diversification[END_REF][START_REF] Bicudo | Andean tectonics and mantle dynamics as a pervasive influence on Amazonian ecosystem[END_REF][START_REF] Hoorn | The Amazon at sea: Onset and stages of the Amazon River from a marine record, with special reference to Neogene plant turnover in the drainage basin[END_REF][START_REF] Latrubesse | The Late Miocene paleogeography of the Amazon Basin and the evolution of the Amazon River system[END_REF]. The sediment discharge in the Amazon fan was relatively modest until 5 Ma and vastly increased in the Pliocene-Pleistocene [START_REF] Hoorn | The Amazon at sea: Onset and stages of the Amazon River from a marine record, with special reference to Neogene plant turnover in the drainage basin[END_REF]Alberts et al., 2018). The lower course of the Amazon River may have become an impassable barrier for these taxa only from the Miocene-Pliocene boundary onward. Moreover, this period also coincides with vegetational and climatic changes, notably the expansion of grasslands not only in the Andes and the Cerrado but also within Amazonia [START_REF] Kirschner | The onset of grasses in the Amazon drainage basin, evidence from the fossil record[END_REF].

Declarations

Ethics approval Experiments have been conducted in accordance with relevant national legislation on the use of animals for research.

Competing interests

The authors declare no competing interests. Author Query Form

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References (Miller et al., 2010;R Development Core Team, 2016;Lösel et al., 2000;[START_REF] Myers | Herpetofauna of the Yutajé-Corocoro massif, Venezuela: Second report from the Robert G. Goelet American Museum-Terramar expedition to the Northwestern tepuis[END_REF]Donnelly, 1996, 1997;Fouquet et al., 2015;Silva et al., 2018;Jaslow et al., 1988;Pereyra et al., 2016;Padial et al., 2014) was mentioned in the manuscript; however, this was not included in the reference list. As a rule, all mentioned references should be present in the reference list. Please provide the reference details to be inserted in the reference list and ensure that all references are cited in ascending numerical order AQ6 References A Fouquet (2016), [START_REF] Fouquet | Cryptic diversity in the Hypsiboas semilineatus species group (Amphibia, Anura) with the description of a new species from the eastern Guiana Shield[END_REF], PD Lösel (2020) and SM Silva (2019) were provided in the reference list; however, these were not mentioned or cited in the manuscript. As a rule, if a citation is present in the text, then it should be present in the list. Please provide the location of where to insert the reference citation in the main body text. Kindly ensure that all references are cited in ascending order.

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