Pristimantis espedeus, Fouquet, Antoine, Martinez, Quentin, Courtois, Elodie A., Dewynter, Maël, Pineau, Kévin, Gaucher, Philippe, Blanc, Michel, Marty, Christian & Kok, Philippe J. R., 2013

Fouquet, Antoine, Martinez, Quentin, Courtois, Elodie A., Dewynter, Maël, Pineau, Kévin, Gaucher, Philippe, Blanc, Michel, Marty, Christian & Kok, Philippe J. R., 2013, A new species of the genus Pristimantis (Amphibia, Craugastoridae) associated with the moderately evelated massifs of French Guiana, Zootaxa 3750 (5), pp. 569-586 : 572-581

Pristimantis espedeus sp. nov.

( Figs 2 , 4–5 , Table 1)

Eleutherodactylus sp. 1 Marty & Gaucher (1999)

Eleutherodactylus sp. 2 Lescure & Marty (2001)

Pristimantis sp. 2 Dewynter et al. (2008); Fouquet et al. (2012)

Holotype. R121 (AF1520), an adult male collected by EC and AF in Montagne Grande Tortue, (4.30901, - 52.357163; 370 m elevation), municipality of Régina, French Guiana, on 13 June 2013 at 18h40. The specimen was recorded and photographed prior to capture ( Fig. 2 ).

Paratypes (n=17). R116–117 (AF0277–0278), two males collected on 10 May 2007 near the Nouragues station in the Réserve Naturelle des Nouragues (4.091667, -52.7; 390 m elevation), municipality of Régina, French Guiana; R118–120 (AF1156, 1190, 1203), three males collected between 24 April and 3 May 2013 near the Aya camp in the Réserve Naturelle de la Trinité (4.619673, -53.408797; 320 m elevation), municipality of St Elie, French Guiana; R122–123 (AF 1521–1522), two males collected with the holotype; R124 (AG508) a male collected on 15 December 2012 in Monts Belvédère (3.719347, -53.412809; 350 m elevation), municipality of Saül, French Guiana; R125–131 (PG452–455, 475–477), seven males collected on 1 February 2005 in Montagne Kotika (3.933333, -54.1972222; 370 m elevation), municipality of Papaïchton, French Guiana; R132 (PG524), a male collected on 27 March 2006 on Pic Coudreau (3.306667, -53.1472222; 400 m elevation), municipality of Saül, French Guiana; and R133 (AG270), a female collected on 12 November 2005 in Mont Lucifer (4.766667, - 53.9166667; 370 m elevation), municipality of St Laurent du Maroni, French Guiana ( Fig. 1 ).

Etymology. The specific name is considered a noun in apposition and derives from the old Norman language espedeus meaning “special” referring to the peculiar ecology and call of the new species (see Advertisement call, and Distribution and ecology sections below). Moreover, espedeus phonetically resembles “sp. 2” in French, which is how the species has been named for almost two decades.

Definition and Diagnosis. The new species is characterized by the following unique combination of characters: (1) SVL small, adult males 23.0 ± 1.1 mm (range 20.7–24.8 mm, n=17), adult female 29.4 mm (n=1) ( Table 1); (2) dorsal skin granular, slightly more granular posteriorly, with a few enlarged tubercles sometimes forming a W-shaped scapular fold and/or dorsal dermal folds, dorsal skin less tuberculate in females, ventral skin granular; (3) tympanic membrane differentiated, tympanic annulus visible externally, supratympanic fold obscuring upper and posterodorsal edges of tympanum, tympanum 40% of eye length; (4) tibia length 1.9 X hand length (5) snout broadly rounded in profile and dorsal view; (6) each upper eyelid with two prominent tubercles; (7) choanae round and large (0.7 mm for the holotype), dentigerous processes of vomers oblique, narrowly separated, each bearing four teeth; (8) vocal slits present, vocal sac median, subgular, nuptial pads rugous translucid, barely visible, located on the dorsal surface of the proximal segment of Finger I; (9) Finger I shorter than the second, 72% of Finger II; (10) fingers and toes with lateral fringes; (11) axillary tubercle absent; (12) ulnar tubercles barely visible or absent; (13) calcars absent, heel tubercles absent, inner metatarsal tubercle oval, much larger than the round outer tubercle; (14) discs broadly expanded, elliptical with broad lateral fringes, outer tubercle round, palmar tubercle heart-shaped; (15) in preservative, dorsum tan, gray and brown, three colour patterns exist: dorsally banded, dorsolaterally banded, and streaked, ventral colouration creamish white with small dark spots becoming denser on throat; (16) iris golden or silver with a dark copper horizontal band; (17) anterior surface of thighs and groin reddish in life and brown in preservative, fingers and toes with red spots in life, lack of yellow circumscribed spot on groin; (18) advertisement call characterized by clusters (0.2– 0.5 s long) of 3–5 very short notes (0.02– 0.03 s long) with a dominant frequency ranging between 2.7–2.8 kHz and emitted every 1.6– 5.3 s; (19) calling activity exclusively crepuscular; (20) occurring at elevations between 200–700 m elevation on massifs reaching> 400 m.

SVL TiL FeL TaL FL HeL HW Ind IOD

P. espedeus sp. nov. M (17) 22.95 12.46 11.69 6.32 9.72 8.56 8.23 2.11 2.51 sd 1.07 0.68 0.53 0.46 0.49 0.41 0.33 0.21 0.23 F (1) 29.40 16.30 16. 0 0 8.30 13.40 11.20 11.00 2.50 3.50 sd NA NA NA NA NA NA NA NA NA

P. inguinalis M (11) 20.23 10.34 9.90 5.48 8.12 7.60 7.07 1.74 2.04 sd 1.09 0.66 0.67 0.45 0.49 0.36 0.51 0.17 0.17

P. sp. 1 M (13) 17.66 9.51 9.01 5.09 7.35 6.69 5.95 1.62 2.06 sd 1.08 0.46 0.58 0.18 0.52 0.59 0.50 0.20 0.17 F (2) 22.20 11.50 11. 0 0 6.40 8.95 8.4 7.45 1.80 2.3

sd 0.42 0.42 0.28 0. 0 0 0.07 0.14 0.49 0.14 0. 0 0

P. sp. 4 M (5) 16.36 9.36 8.68 5.32 7.38 6.28 5.30 1.74 1.96 sd 0.85 0.57 0.54 0.43 0.37 0.33 0.46 0.17 0.22 F (6) 22.69 12.20 11.47 7.10 9.57 8.43 7.85 2.05 2.52 sd 1.57 0.51 0.67 1.24 1.43 0.58 0.47 0.21 0.17 EN ED TD FD 4TD ETS 1FiL 2FiL HL

P. espedeus sp. nov. M (17) 2.74 3.06 1.19 1.39 1.29 4.05 3.01 4.19 6.56 sd 0.26 0.22 0.25 0.09 0.11 0.24 0.49 0.49 0.41 F (1) 4.30 4.01 2.30 1.70 1.60 5.20 4.50 6.10 8.60 sd NA NA NA NA NA NA NA NA NA

P. inguinalis M (11) 2.44 2.83 0.97 1.10 1.02 3.49 2.47 3.31 5.62 sd 0.27 0.12 0.09 0.12 0.10 0.25 0.15 0.15 0.34

P. sp. 1 M (13) 2.18 2.52 0.85 0.78 0.75 3.24 2.21 2.94 4.63 sd 0.32 0.21 0.14 0.12 0.13 0.19 0.24 0.34 0.24 F (2) 3.00 2.80 0.95 1.00 1.00 4.15 2.60 3.40 5.25 sd 0.00 0.00 0.21 0.14 0.14 0.35 0.00 0.28 0.49

P. sp. 4 M (5) 2.10 2.32 0.62 0.84 0.80 3.18 2.00 2.58 4.48 sd 0.32 0.25 0.04 0.11 0.12 0.23 0.23 0.28 0.33 F (6) 3.05 3.33 0.73 1.15 1.20 4.34 2.85 3.65 6.02 sd 0.22 0.14 0.10 0.18 0.14 0.21 0.33 0.58 0.34 Morphological comparisons with congeneric species. Using the raw data, MDFA clearly segregates well the four species ( Fig. 3 a). Three variables notably contribute to the discrimination in opposite ways: HL and HW on the one hand, and TiL on the other. When the MDFA is based on the residuals, discrimination is less clear, especially between Pristimantis espedeus sp. nov. and P. sp. 1 ( Fig. 3 b), suggesting that most of the morphological variation is size-based between these species ( Fig. 3 c). Nevertheless, the variables HL and TiL still contribute appreciably and in opposite ways. Therefore, we plotted the ratio between these two variables. In P. espedeus sp.

nov. and P. inguinalis , TiL is less than 50% of HL, while it accounts for> 50% in most specimens of P. sp. 1 and P. sp. 4 ( Fig. 3 d, Table 2 ).

Compared to the three other species of the unistrigatus group occurring in French Guiana (i.e. Pristimantis inguinalis , P. sp. 1 and P. sp. 4, the last two were previously partly misidentified with P. marmoratus in French Guiana e.g. in Lescure & Marty 2001), Pristimantis espedeus sp. nov. differs mainly by its larger SVL in males: mean of 23.0 mm (n=17) in P. espedeus sp. nov. vs. mean of 20.2 mm (n=11) in P. inguinalis , mean of 17.7 mm (n=13) in P. sp. 1, and mean of 16.4 mm (n=5) in P. sp. 4 ( Table 1; Fig. 3 ). Pristimantis espedeus sp. nov. further differs from P. sp. 1 and P. sp. 4 by its red-orange inner thighs and groin in life (dark grey in P. sp. 1 and P. sp. 4), a more granular skin, a less clear W-shaped scapular fold, a proportionally larger head ( Fig. 4 ), a less acuminate snout, more prominent tubercles on arms, larger finger discs, and a heart-shaped palmar tubercle (elliptical in P. sp. 1 and P. sp. 4). From P. inguinalis the new species mainly differs by its red-orange inner thighs and groin in life (a bright, well circumscribed yellow spot in P. inguinalis ), and a cream ventral colouration (black in P. inguinalis ).

From the species of the unistrigatus group reported to occur below ca. 700 m elevation in the Guiana Shield [i.e. P. ockendeni (Boulenger, 1912) and P. marmoratus (Boulenger, 1900) ], P. espedeus sp. nov. is readily diagnosable by its body size: P. ockendeni is larger (adult female holotype = 34.0 mm vs. 29.4 mm in P. espedeus sp. nov. female), and P. marmoratus is obviously smaller (adult male holotype [with large vocal slits present] = 18.0 mm vs. min. 20.7 mm in P. espedeus sp. nov. males). Pristimantis espedeus sp. nov. further differs from P. ockendeni by its reddish groin and thighs (absent in P. ockendeni ), and from P. marmoratus by its cream venter (the preserved holotype of P. marmoratus has a dark grey venter suggestive of a dark ventral colouration in life). Pristimantis ockendeni from Manaus (Lima et al. 2006) apparently corresponds to a still undescribed species given its smaller body size compared to the holotype of P. ockendeni , and can be easily distinguished from P. espedeus sp. nov. by call characteristics (see below). Three other species associated with the name P. ockendeni have been described by Elmer and Cannatella (2008) from the lowlands of Ecuador and do not correspond to P. espedeus sp. nov. given their smaller SVL and the absence of reddish colour on groin and thighs.

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). Among the Pristimantis species found at midelevation in Pantepui, P. espedeus sp. nov. can mainly be distinguished from P. jester (Means & Savage, 2007) by the presence of a tympanum (absent in P. j e s t e r) and presence of fringes on fingers (absent in P. jester ), from P. saltissimus (Means & Savage, 2007) by a slightly larger body size (16 mm – 27.1 mm in P. saltissimus vs. 20.7–29.4 mm in P. espedeus sp. nov.), a large, prominent tympanum (present, but indistinct in P. saltissimus ), and presence of fringes on fingers (absent 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 2013] vs. 29.4 mm in P. espedeus sp. nov. female), a distinct tympanum (weakly distinct), and presence of fringes on fingers (absent), from P. sarisarinama (Barrio-Amorós & Brewer-Carias, 2008) by the presence of nuptial pads in males (reported to be absent in P. sarisarinama ), tubercles on eyelids (absent in P. sarisarinama ), and distinct vocalisation (1–2, rarely 3 notes in P. sarisarinama vs. 3–5 notes in P. espedeus sp. nov.), from P. pulvinatus (Rivero, 1968) by its slightly smaller body size (males 23.0– 26.1 mm in P. pulvinatus vs. 20.7–24.8 mm in P. espedeus sp. nov.) and distinct vomerine teeth (indistinct or absent in P. pulvinatus ), from P. memorans (Myers & Donnelly, 1997) by the presence of fringes on fingers (absent in P. memorans ) and a higher internote interval (0.20– 0.29 s in P. memorans vs. 0.066– 0.124 s in P. espedeus sp. nov.).

Pristimantis espedeus sp. nov. can also be distinguished from the upper Amazonian species of the unistrigatus group Pristimantis lirellus (Dwyer, 1995) , P. croceoinguinis (Lynch, 1968) , and P. imitatrix (Duellman, 1978) , with which it shares phylogenetic affinities according to previous analyses (Fouquet et al. 2012) as well as from P. carvalhoi (Lutz & Kloss, 1952) and P. variabilis (Lynch, 1968) that are related to the previous species according to Dwyer (1995), by the absence of a bright yellow spot in the groin region [present in the aforementioned species according to Dwyer (1995)], and in having a reddish groin region instead.

Description of Holotype ( Figs. 2 , 4–5 ). An adult male 23.8 mm SVL in excellent condition. Head longer than wide (HW 8.4 mm, HeL 9.3 mm; HL 90.3% of HeL), cranial crests not detectable. In dorsal view, snout longer than eye length (ETS 130% of ED), broadly rounded, canthus rostralis straight and loreal region slightly concave, eyenaris distance slightly shorter than eye length (EN 94% of ED). Nares slightly protuberant, directed posterolaterally, visible from frontal and dorsal views. Upper eyelid tubercles slightly prominent on the granular skin. Tympanum distinct, 1.6 mm long, ca. 50% of eye length (3.3 mm). Supratympanic fold conspicuous in life, originating at posterior corner of eye, failing to reach shoulder; post-rictal tubercles absent. Vocal sac present. Throat and under surface of thighs smooth, belly granular; posteroventral surface of thigh and cloacal region granular; cloacal sheath absent. Dentigerous processes of the vomers oblique, narrowly separated, each bearing four teeth. Vocal slits visible on the floor of the mouth. Dorsal skin granular (more granular posteriorly) with enlarged tubercles, including dermal folds on dorsal portion of the flanks.

Hand length 29% of SVL. Finger I 79 % of II. Relative length of fingers III> IV> II> I; adpressed Finger I fails to reach proximal edge of digital pad of Finger II; adpressed Finger IV reaches the intercalary cartilage of Finger III on the left side, the base of the disc of Finger III on the right side. Large, barely visible rugous translucid nuptial pad on the dorsal surface of the proximal segment of Finger I. Lateral fringes on all fingers, best developed pre-axially on Fingers II and III ( Fig. 3 ). Finger discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded; disc of Finger III (1.4 mm) 2.1 times wider than t distal end of adjacent phalanx. Palmar tubercle large, well defined, not fully pigmented, deeply bifid; thenar tubercle large, protuberant, ovoid; supernumerary tubercles few, large (almost equal in size to subarticular tubercles), slightly protuberant; subarticular tubercles large, round and protuberant, one each on FI and FII, two each on FIII and FIV.

Hind limbs moderate in length, heels slightly overlap when held at right angles to sagittal plane; TiL 54% of SVL; FL 44% of SVL. Relative length of toes IV> V> III> II> I; tip of Toe V extends to the distal edge of the distal subarticular tubercle on Toe IV; tip of Toe III extends to the proximal half of the penultimate subarticular tubercle on Toe IV. Lateral fringes on all toes, best developed pre-axially on Toes III–IV; webbing basal between Toe IV–V ( Fig. 2 c). Toe discs mostly equal in size to finger discs, WTD/WFD = 1; toe discs broadly expanded, elliptical, broader than long, circumferential groove conspicuous, distal edge of disc rounded. Inner metatarsal tubercle elongate, oval, about four times the size of the projecting, round outer metatarsal tubercle; subarticular tubercles round, large and protuberant; supernumerary plantar tubercles small, low and round, increasing in size distally. Calcars absent; no outer tarsal tubercles detectable in preservative; inner tarsal fold not detectable.

Colour of the Holotype in preservative ( Fig. 2 a–f). Dorsal background colour light brown, with a broad dark brown vertebral stripe extending from snout to vent, and two large laterodorsal dark brown bands extending from anterior corner of eye to cloaca. Two dark brown bands on the forearms and three well-defined brown bands on shanks and tarsi, ill-defined bands on thighs. Cloacal area dark brown. Five ill-defined dark bands on the upper lip and the loreal region; black supratympanic ridge obscuring upper part of tympanum; black band extending from tip of snout, tapering to anterior corner of eye. Dark stripe on lower edge of orbit. Ventral colouration creamish white with small dark spots denser on throat and thighs.

Colour of the Holotype in life ( Fig. 2 g). Dorsal background colour chestnut brown, with a broad dark brown stripe extending from snout to vent flanked by two large laterodorsal dark brown bands extending from anterior corner of eye to cloaca. Dorsal half of flanks yellowish brown with ill-defined oblique dark brown reticulations. Loreal region yellowish brown with a series of ill-defined greyish brown markings on the upper lip. Arms and legs yellowish brown with transversal dark bands. Groin and anterior surface of thighs bright red. Red markings also visible on fingers and toes. Belly creamish white with yellowish granular tubercles; throat similarly coloured, but with small dark brown spots.

Variation. Pristimantis espedeus is highly polychromatic dorsally. Three main patterns exist: a dorsally banded morph (cf. holotype), a dorsolaterally banded morph (two dorsolateral reddish brown bands in life; R 120, 124, 133 in Fig. 5 ), and a streaked morph (with more or less well-defined chevron-like markings; R 118, 122, 123 in Fig. 5 ). A W-shaped dark brown scapular fold is sometimes visible particularly in the streaked morph.

Sexual dimorphism can be evaluated based on the only female collected. The female is much larger than the largest male, the skin is less granular, and thumbs lack nuptial pads. Female colouration is similar to the dorsolaterally banded morph observed in some males. No additional significant difference was detected.

Advertisement call ( Fig. 6 , Table 3 ). The following description is based on three recordings, one of the holotype (Montagne Grande Tortue) and two from uncollected individuals from the Réserve Naturelle des Nouragues and the Réserve Naturelle de la Trinité, respectively.

The advertisement call consists of short clusters (X= 0.33 s; range=0.24– 0.5 s) of generally 4 (3 to 5) short notes (X= 0.02 s; range=0.019– 0.025 s) having an internote interval of 0.09 s (range=0.066– 0.124 s) emitted between silent intervals of 3.21 s (range=1.58– 5.29 s) ( Fig. 6 ).

The spectral structure consists of three harmonics, with the fundamental frequency dominating (X=2.7 kHz, range=2.68–2.84 kHz, Fig. 6 ). Two secondary harmonics are visible (but not shown in Fig. 6 ), having a frequency of ca. 5.4 and 8.2 kHz, respectively.

The vocalization of Pristimantis espedeus sp. nov. is easily distinguished from that of the two sympatric species with known advertisement calls (the call of P. sp. 4 remains undocumented): P. inguinalis emits a single note of ca. 3 kHz every 3–5 s, and P. sp. 1 emits clusters of 7–10 very short notes of ca. 4 kHz. The species of the unistrigatus group occurring in Manaus referred to as P. ockendeni by Lima et al. (2006) emits shorter clusters (0.20– 0.25 s) of 4–5 notes with higher dominant frequency (3.8 kHz).

Distribution and ecology. The nine known populations of Pristimantis espedeus sp. nov. are located in the Réserve Naturelle des Nouragues, the Réserve Naturelle de la Trinité, the Réserve Biologique Lucifer Dékou- Dékou (Mont Lucifer), Mont Itoupé, Montagne Kotika, Monts Atachi-Bacca, Monts Belvédère, Pic Coudreau and Montagne Grande Tortue, respectively ( Fig. 1 ). Interestingly, the new species has not been detected from other massifs harbouring apparently suitable habitats such as Pic Matécho, Mont Galbao and Massif du Mitaraka. Pristimantis espedeus sp. nov. may occur on other massifs such as Montagne Bellevue de l’Inini, Montagne Française and possibly Lely and Nassau mountains in Suriname. All the documented populations seem completely isolated from one another by a very sharp altitudinal limit, (ca. 200 m elevation minimum).

Pristimantis espedeus sp. nov. is arboreal and only found in forests from 200 to 700 m elevation on massifs> 400 m. Calling activity is limited to dusk and dawn, from ca. 18h00 to 19h00 and from 5h30 to 6h30; males call head down on the trunks, between 0.5 and 2 m above the ground ( Fig. 2 g). Although the species seems predominantly arboreal, an amplectant pair and several juveniles were observed in the leaf litter by two of us (PG and AF, respectively). Calling activity is more intense at the beginning of the rainy season (January-March), significantly decreasing afterwards and apparently ends in June. The peculiar distribution of the new species suggests possible endemism to the Eastern Guiana Shield. Pristimantis espedeus sp. nov. occurs syntopically with P. sp. 4, and rarely with P. inguinalis as well as with species belonging to other species groups [ P. chiastonotus , P. zeuctotylus , P. gutturalis , P. aff . zimmermanae (P. sp. 3 in Lescure & Marty 2001)]

Figures

FIGURE 1. Map showing documented populations of Pristimantis espedeus sp. nov. (a) The Guiana Shield is delimited by a white line and the eastern part is separated from the western by a dotted line. A white convex polygon delimits the documented range of Pristimantis espedeus sp. nov. (b) A focus on French Guiana with white dots indicating populations for which vouchers exist (1: Réserve Naturelle de la Trinité, 2: Mont Lucifer, 3: Mont Kotika, 4: Montagne Grande Tortue (type locality), 5: Réserve Naturelle des Nouragues, 6: Pic Coudreau, 7: Monts Belvédère); black dots indicate populations without collected material (8: Monts Atachi-Bacca, 9: Mont Itoupé).

FIGURE 2. Holotype (R 121) of Pristimantis espedeus sp. nov. (a) dorsal view, (b) ventral view, (c) right foot, (d) right hand, (e) head in lateral view, (f) posterior face of thigh in preservative, (g) holotype calling.

FIGURE 3. Plots of discriminant scores from MDFA based on (a) raw morphological measurements (b) residual from the different measurement and SVL taken on the four Pristimantis spp. of the unistrigatus group occurring in French Guiana. Eigen value of each axis is indicated on the right and on the top. We plotted (c) the distribution of SVL for each species and given the results from the two MDFA (d) the ratio between TiL and HL.

FIGURE 4. Comparison between the four Pristimantis species of the unistrigatus group occurring with certainty in French Guiana. The four specimens are adult males. P. espedeus sp. nov. holotype: R 121 (AF 1520), P. inguinalis: R 179 (AF 804), P. sp. 1: R 189 (AF 757) and P. sp. 4: R 211 (AF 1030). (a) Photographs of living specimens (unreferenced); (b) dorsal view; (c) ventral view; (d) lateral view of head; (e) ventral view of hand and foot. Some characters of the diagnosis are indicated by arrows and corresponding numbers.

FIGURE 5. Variation among the type series of Pristimantis espedeus sp. nov. in preservative. R 133 (AG 270) is the only female collected and R 121 (AF 1520) is the holotype.

FIGURE 6. Spectrograms (above) and oscillograms (below) of the calls of three species of the unistrigatus group occurring in French Guiana; the call of P. sp. 4 is unknown. For ease of comparison spectrograms show dominant frequencies only.

Tables

TABLE 2. Canonical weights of each variable on the first two axes using raw data and residuals. Abbreviations are explained in the text.

	DS1	DS2	DS1	DS2
SVL	0.4043	0.5580	NA	NA
TiL	-0.7746	-1.9648	-0.6692	0.4986
FeL	-0.1695	-0.1890	-0.0789	0.0382
TaL	-0.1515	-0.5637	-0.3078	-0.4620
FL	0.3052	-0.6559	-0.2620	0.5919
HeL	0.1286	0.0694	0.0285	-0.1716
HW	0.4113	1.2443	0.3068	0.1673
Ind	-0.1468	-0.2853	-0.2139	-0.2204
IOD	0.2258	0.1345	0.1168	0.5189
EN	0.0329	0.5226	0.3617	-0.3499
ED	0.3033	0.0341	0.0474	0.6010
TD	-0.2631	0.4032	0.2167	0.0198
FD	0.2103	-0.5731	-0.2493	-0.3794
4TD	-0.1623	0.0055	-0.0085	-0.4549
ETS	0.0228	-0.1668	-0.0766	-0.2485
1FiL	-0.1890	-0.2406	-0.1557	-0.0419
2FiL	0.4760	0.1926	0.1273	0.3304
HL	0.3386	1.2663	0.4022	-0.6587

TABLE 3. Bioacoustic variables (mean and sd) in populations of Pristimantis espedeus sp. nov. from three localities. The poor quality of the recording of the holotype (Montagne Grande Tortue) does not allow measurements of the note duration, internote intervals, and frequency of harmonics.

	Nouragues	Trinité	Montagne Grande Tortue
Call duration (s)	0.34 (0.12)	0.39 (0.02)	0.27 (0.01)
Note duration (s)	0.02 (0.00)	0.02 (0.00)	NA
Notes/call	3.83 (0.98)	4 (0.00)	3 (0)
Silent interval (s)	3.61 (1.12)	3.45 (1.32)	2.58 (0.25)
Internote interval (s)	0.09 (0.02)	0.10 (0.02)	NA
Calls/min	12	20	20
Dominant Frequency (Hz)	2803 (19.26)	2696 (8.52)	2798 (12.74)
2nd harmonic (Hz)	5566 (50.73)	5348 (9.09)	NA
3rd harmonic (Hz)	8430 (41.56)	8063 (19.99)	NA