Pristimantis ockendeni (Boulenger, 1912)

Monico, Alexander Tamanini, Ferrao, Miqueias, Chaparro, Juan Carlos, Fouquet, Antoine & Lima, Albertina Pimentel, 2022, A new species of rain frog (Anura: Strabomantidae: Pristimantis) from the Guiana Shield and amended diagnosis of P. ockendeni (Boulenger, 1912), Vertebrate Zoology 72, pp. 1035-1065 : 1035

publication ID	https://dx.doi.org/10.3897/vz.72.e90435
persistent identifier	https://treatment.plazi.org/id/62A5E4C0-7EBC-5352-833F-0A44B5FAD525
treatment provided by	Vertebrate Zoology by Pensoft
scientific name	Pristimantis ockendeni (Boulenger, 1912)
status

Treatment

Figs 3 View Figure 3 , 4 View Figure 4 , 5 View Figure 5

Hylodes ockendeni - Boulenger (1912)

Eleutherodactylus ockendeni - Lynch (1996)

Pristimantis ockendeni - Heinicke et al. (2007), Hedges et al. (2008), Padial et al. (2014), Ocampo et al. (2016), von May et al. (2017), Villacampa et al. (2017)

Syntypes.

BMNH 1947.2.16.88, BMNH 1947.2.16.89 and BMNH 1947.2.16.90 (Fig. 3 View Figure 3 ), collected on 07 May 1907 from La Union, Rio Huacamayo, Carabaya, southeastern Peru (13°31′58.3″S, 69°45′06.7″W; 783 m elevation).

Th e specimens are in a good state of preservation, allowing for an easy view, for example, of the dark bar between the eyes, of oblique black streak in front of and behind the eye, oblique brown bars on the tibia, supratympanic stripe and melanophores in ventral surface.

We compared the newly collected individuals with the morphological diagnosis described by Boulenger (1912) and the syntypes of species (through high quality images made available by the Natural History Museum - London) to ensure that we are dealing with the same taxon. Moreover, the molecular analyses confirmed that our material is conspecific to a specimen (voucher RvM5_12) collected near the type locality (112 km linear distance) and made available by von May et al. (2017). Our samples are distributed at altitudes ranging from 223 m (in Brazil) to 924 m (in Peru), range that includes the altitude of the type locality (783 m).

The characters available in the species description are relatively vague. Boulenger (1912) defined the size of the species as 34 mm from snout to vent, based on the two female syntypes (NHM 1947.2.16.88 and NHM 1947.2.16.89). Boulenger might not have included the third syntype (NHM 1947.2.16.90) because he believed it was a juvenile individual, but our examination showed this specimen to be an adult male.

We acknowledge that>100 km separates the type locality from the closest newly collected population that could corresponds to Pristimantis ockendeni . Therefore, on the sole basis of geographical distance and considering the megadiversity and the cryptic morphology of this clade we cannot completely reject the hypothesis that our populations in fact belong to another species than P. ockendeni .

Nevertheless, four other nominal species of the Pristimantis unistrigatus species group are known to occur in the area [i.e. P. altamazonicus (Barbour & Dunn, 1921), P. carvalhoi (Lutz, 1952), P. divnae Lehr & von May, 2009, and P. ventrimarmoratus (Boulenger, 1912); Duellman 2005; Villacampa et al. 2017] and one undescribed (" Pristimantis sp. 3" sensu Villacampa et al. 2017). However, all of them have distinctive features on groin (yellow, orange or red marks) or belly (black marble) that readily distinguishes them from P. ockendeni . Also, the main lineages of the P. unistrigatus species group related to P. ockendeni [i.e., P. matidiktyo , P. daquilemai Brito-Zabata, Reyes-Puig, Cisneros-Heredia, Zumel & Ron, 2021, and P. delius (Duellman & Mendelson, 1995)], and possible any undescribed species related to them, can be easily distinguished from P. ockendeni . For example, P. matidiktyo differs by the dorsum without any marks and the presence of a small tubercle on the upper eyelid, P. daquilemai differs by its flanks densely tuberculated and the presence of a prominent rostral papilla at tip of snout, and P. delius is distinguished by the snout acutely rounded in dorsal view and the absence of dentigerous processes of vomers. Finally, the characters described by Boulenger (1912), although vague, but completed by the examination of the syntypes (see below), strongly suggests that the newly collected specimens are conspecific to P. ockendeni .

Referred material.

Available in Appendix 1.

Amended diagnosis.

The species is characterized by the combination of the following characters, based on the description of Boulenger (1912) with additions to this study, obtained through examination of the syntypes (Fig. 3 View Figure 3 ) and to the material recently collected (Fig. 4 View Figure 4 ): (1) dorsal skin smooth to shagreen, frequently with distinctly enlarged tubercles, with or without W-shaped on scapular region, a narrow light vertebral line may be present; (2) tympanum very indistinct, its length 30 to 44% of eye length, tympanic membrane and tympanic annulus visible but sometimes inconspicuous; with or without supratympanic stripe; (3) snout short and subacuminate in dorsal view, with moderately strong, curved canthus and very oblique, concave loreal region; (4) upper eyelid usually bearing tubercles; dark bar between the eyes, and oblique black streak in front of and behind the eye; cranial crests absent; (5) nostril near the tip of the snout; interorbital space hardly as broad as the upper eyelid; (6) tongue oval, entire or indistinctly nicked behind; (7) dentigerous processes of vomers in two oblique oval groups just behind the level of the choanae, clearly visible; (8) vocal slits present, vocal sac median to subgular; nuptial pads absent; (9) Finger I shorter than Finger II; fingers moderate, discs ovoid to expanded (widest on fingers III and IV); (10) fingers without lateral fringes; (11) three to four ulnar tubercles, ill defined; (12) tibia length 49-56% of SVL; (13) heel bearing one rounded tubercle; small tarsal tubercles, ovoid, poorly visible in fixed specimens; inner edge of tarsus with a short fold; (14) two feebly prominent metatarsal tubercles, being thenar tubercle ovoid to elliptical and small palmar tubercle ovoid; subarticular tubercles well developed but small; (15) toes without lateral fringes; scarcely a rudiment of web between them; (16) Toe I clearly smaller than Toe II, toes moderate, discs ovoid to expanded (widest on toes III to V); (17) dorsal coloration very variable; two or three oblique brown bars on the tibia; ventral surface smooth to slightly areolate, variable concentration of melanophores; and ventral region of the femur externally slightly areolate; (18) absence of spot on groin; (19) dichromatic iris in life, the lower part being coppery and the upper part cream; (20) SVL in adult males of 18.3-22.8 mm (n = 9; Table 4 View Table 4 ) and females of 30.4-30.6 mm (n = 2; Table 4 View Table 4 ); and (21) advertisement call with average call duration of 541 ± 151 ms, inter-note interval of 44 ± 5 ms (68 ± 11 ms), minimum frequency of 2,046-2,610 Hz, maximum frequency of 2,964-3,641 Hz and dominant frequency of 2,519-3,143 Hz.

Additional comments about P. ockendeni morphology.

Dorsal pattern of males is highly variable, showing three main patterns (n = 8 specimens): (i) coloration strongly delimited and different from sideview, with or without dark transverse stripes on dorsum (n = 4 specimens; Fig. 4A View Figure 4 ; in life, Fig. 5A View Figure 5 , 5B View Figure 5 , 5D View Figure 5 and 5F View Figure 5 ); (ii) irregular dark markings present on dorsum (Fig. 4C View Figure 4 ); and (iii) similar to the previous pattern, but lighter (Fig. E); mostly, interorbital region is covered by a dark band (n = 5 specimens). In addition, in some individual, W-shaped mark on scapular region present, even if not very evident (Fig. 4C View Figure 4 ; Fig. 5E View Figure 5 and 5G View Figure 5 ). Ventral surface can have different shades, varying according to the concentration of melanophores: darker (Fig. 4B View Figure 4 ), intermediary (Fig. 4D View Figure 4 ) or lighter (Fig. 4F View Figure 4 ). Ventral skin texture is smooth (arms and chest region) to areolate (especially in belly and on femur region).

Dorsal coloration of males in life is highly variable, from brown reddish to yellow greenish (Fig. 5A-5G View Figure 5 ) with or without interorbital bars. We do not have information on females. Arm is light yellowish. The iris is dichromatic (dark copper metallic down part and cream to silver upper part. Ventral surface is cream translucent (in the belly) and gray translucent (in the pelvic region) with dark melanophores (Fig. 5H View Figure 5 ).

Advertisement call of Pristimantis ockendeni .

We recorded 40 calls of seven males from the underwood vegetation of about 2 m above the ground (air temperatures 23-25°C; relative humidity 82-99). Descriptive statistics of call parameters are presented in Table 5 View Table 5 and voucher in Appendix 4. The advertisement call of Pristimantis ockendeni has a call duration of 541 ± 151 ms (334-939 ms, n = 40 calls) and 4-8 notes (6.1 ± 0.9, n = 243 notes) with note duration of 24-158 ms (41 ± 31 ms). Calls with 6 and 7 notes were the most common arrangement (37.5% of recorded calls, n = 15, and 32.5% n = 13, respectively), followed by calls with 5 notes (25%, n = 10 calls), and rarely with 4 and 8 notes (2.5% each, n = 1). Inter-note intervals within calls average 62 ± 11 ms (38-87 ms). The spectral structure of the note has average minimum frequency of 2,389 ± 190 Hz (2,046-2,610 Hz), while average maximum frequency is 3,350 ± 201 Hz (2,964-3,641 Hz), and the dominant frequency is 2,864 ± 202 Hz (2,519-3,143 Hz) (Fig. 6A View Figure 6 and 6B View Figure 6 ).