taxonID	type	description	language	source
322387A7BB1DFFAD96681F927D334BFA.taxon	materials_examined	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. 10 th 1901 ” during the Coppename expedition (Anonymous, 1903). 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. Referred specimens. MNHN-RA- 2020.0118 (field no. AF 3435), an adult male collected by A. 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 A 1212, 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 A 1210 – 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). 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 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 specimen of P. grandoculis (MNHN-RA- 2020.0118) from Bakhuis, Suriname. c A photograph in life of another recently collected specimen P. grandoculis (NZCS A 1210) 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. 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; Fouquet et al., 2013; Kok et al., 2018). 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 (Means & Savage, 2007) 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 (Means & Savage, 2007) 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 (Schlüter & Rödder, 2007; see also Kok & Barrio-Amorós, 2013) 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 (Rivero, 1968) 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).	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB1DFFAD96681F927D334BFA.taxon	description	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 A 1210 has a dark band running along the canthus rostralis which is absent in the other specimens (Fig. 2 c). The W-shaped scapular mark is well-developed 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 exoccipital, fro — frontal, max — maxillary, men — mentomeckelian, nas — nasal, neo — neopalatine, par — parasphenoid, pre — premaxillary, pro — prootic, pte — pterygoid, qua — quadratojugal, sep — septomaxilla, sph — sphenethmoid, squ — squamosal, vom — vomer 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. 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.	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB1DFFAD96681F927D334BFA.taxon	distribution	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 (Vacher et al., 2017) and Pipa aspera (Vacher et al., 2020), both species displaying similar geographical boundaries and deep genetic structures across 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 (Fouquet et al., 2018). Both are pairs of sister species with similar ecology and body size.	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB12FFB7966819F47D524D83.taxon	description	Pristimantis sp. 1 Fouquet et al., 2019	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB12FFB7966819F47D524D83.taxon	materials_examined	Holotype. MNHN-RA- 2020.0102 (field no. AF 2786), 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). Paratypes. Twenty-seven specimens (23 males and four females) from French Guiana and the state of Amapá, Brazil: MNHN-RA- 2020.0092 – 0112; MPEG 41819 – 41825 (Appendix 2).	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB12FFB7966819F47D524D83.taxon	etymology	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 = 24) and adult females 23.4 ± 0.5 mm (range 22.9 – 23.9 mm, n = 4) (Table 1); (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. 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; Fouquet et al., 2013; Kok et al., 2018). 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 (Means & Savage, 2007) by the presence of a tympanum (absent in P. jester), from P. saltissimus (Means & Savage, 2007) 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 (Schlüter & Rödder, 2007) by its smaller body size (females 32.4 – 33.6 mm in P. guaiquinimensis [see Kok & Barrio-Amorós, 2013] vs. 22.9 – 23.9 mm in P. crepitaculus sp. nov.), a distinct tympanum (weakly distinct in Pristimantis crepitaculus sp. nov.), from P. sarisarinama (Barrio-Amorós & Brewer-Carías, 2008) by iris colour in life (black vertical streak running across the iris in P. crepitaculus sp. nov. vs. no such streak in P. sarisarinama), and distinct vocalization (1 – 2, rarely 3 notes in P. sarisarinama vs. 6 – 12 notes in P. crepitaculus sp. nov.), from P. pulvinatus (Rivero, 1968) 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).	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB12FFB7966819F47D524D83.taxon	description	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 well-defined 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. 5 a). 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 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 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. 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. col — columella, den — dentary, exo — exoccipital, fro — frontal, max — maxillary, men — mentomeckelian, nas — nasal, neo — neopalatine, par — parasphenoid, pre — premaxillary, pro — prootic, pte — pterygoid, qua — quadratojugal, sep — septomaxilla, sph — sphenethmoid, squ — squamosal, vom — vomer 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. Well-developed 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 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 2 nd 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.	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB12FFB7966819F47D524D83.taxon	distribution	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.	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
322387A7BB06FFB995D01FAD788A4FBA.taxon	description	PLPDR 128 (MPEG 41823), Paratype, Oiapoque – Amapá, Brazil MNHN-SO- 2022 - 563 track 59 Marty and Gaucher (NA), Kaw montain − French Guiana 160106 _ 0692 (AF 3530), Itoupé− French Guiana MNHN- SO- 2022 - 561 160106 _ 0693 (AF 3539), Holotype, Itoupé− French Guiana MNHN-SO- 2022 - 560 LS 110010 (AF 1606), Saul Limonade − French Guiana MNHN-SO- 2022 - 559	en	Fouquet, Antoine, Peloso, Pedro, Jairam, Rawien, Lima, Albertina P., Mônico, Alexander T., Ernst, Raffael, Kok, Philippe J. R. (2022): Back from the deaf: integrative taxonomy revalidates an earless and mute species, Hylodes grandoculis van Lidth de Jeude, 1904, and confirms a new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the Eastern Guiana Shield. Organisms Diversity & Evolution (New York, N. Y.) 22 (4): 1065-1098, DOI: 10.1007/s13127-022-00564-w, URL: http://dx.doi.org/10.1007/s13127-022-00564-w
