identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
85F826E58E0F5EDABF3AA1FF49A9BDDE.text	85F826E58E0F5EDABF3AA1FF49A9BDDE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pristimantis donnelsoni Juan Pablo Reyes-Puig, Miguel Urgiles-Merchan, Carolina Reyes-Puig, Daniela Franco-Mena, Diego Batallas & Juan M. Guayasamin 2023	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> 
Pristimantis donnelsoni Juan Pablo Reyes-Puig, Miguel 
Urgiles-Merchan
, Carolina Reyes-Puig, Daniela Franco-Mena, Diego Batallas &amp; Juan M. Guayasamin
 sp. nov.</p>
            <p>Figs 1, 2, 3, 4, 5, 6, 7, 8</p>
            <p>Type material.</p>
            <p> Holotype. DHMECN 14701, adult female (Figs 2 - 7), collected by CRP, JPRP, and DFM from Reserva Chamana, eastern slopes of Tungurahua Volcano, Ulba,  Baños , Tungurahua, Ecuador (1.4272°S, 78.3967°W; 3,028 m alt.) on 13 August 2018. </p>
            <p> Paratypes. (22, 13 ♂ / 9 ♀, Fig. 6): ZSFQ 1087, adult female collected by CRP, JPRP and DFM from Reserva Chamana, Ulba,  Baños , Tungurahua, Ecuador (1.421°S, 78.3866°W; 2909 m alt.) on 11 August 2018. ZSFQ 1090 and 1096, adult females collected by CRP, JPRP, and DFM from Reserva Chamana, Ulba,  Baños , Tungurahua, Ecuador (1.4256°S, 78.3854°W; 2977 m alt.) on 12 August 2018. ZSFQ 1100, adult male collected by CRP, JPRP, and DFM from Reserva Chamana, Ulba,  Baños , Tungurahua, Ecuador (1.4273°S, 78.3853°W; 3000 m alt.) on 12 August 2018. ZSFQ 1103 and 1105, adult males collected with the same data as ZSFQ 1090. DHMECN 16606, adult male collected by JPRP from Reserva Chamana, Ulba,  Baños de Agua Santa, Tungurahua, Ecuador (1.4343°S, 78.3942°W; 3125 m alt.) on 14 August 2018. DHMECN 16608, 16610 and 16613, adult males with the same data as DHMECN 16606. DHMECN 4772, 4777 and 4782 adult females collected by JPRP from  Nahuazo-Runtún ,  Baños ,  Baños de Agua Santa, Tungurahua, Ecuador (1.4258°S, 78.4257°W; 3100 m alt.) on 28 March 2008 DHMECN 4778, adult male and DHMECN 4779, adult female, collected by JPRP and Nelson Palacios from San Antonio,  Baños ,  Baños de Agua Santa, Tungurahua, Ecuador (1.4572°S, 78.3016°W; 2850 m alt.) on 30 April 2007. DHMECN 4769, adult male collected by JPRP and  Salomón Ramírez from Pondoa,  Baños ,  Baños de Agua Santa, Tungurahua, Ecuador (1.4370°S, 78.4426°W; 3000 m alt.) on 31 March 2007. DHMECN 4774 and DHMECN 4781 adult males collected by JPRP from Pondoa,  Baños ,  Baños de Agua Santa, Tungurahua, Ecuador (1.4370°S, - 78.4426°W; 3240 m alt.) on 21 March 2007. DHMECN 4807, adult male collected by JPRP,  Salomón Ramírez , and Stalin  Cáceres , and Luis Recalde from Bosque Protector Cerro Candelaria,  Río Verde,  Baños de Agua Santa, Tungurahua, Ecuador (1.4572° S, 78.3016°W; 2850 m alt.) on 11 May 2008. DHMECN 18159 and 18160, adult females collected by JPRP, Patricio Vinueza, Paulete Benavidez, and Nantar Kuja in Bosque Protector Guamag, Ulba,  Baños de Agua Santa, Tungurahua, Ecuador (1.4194°S, 78.3648°W; 2800 m alt.) on 28 May 2022. DHMECN 18185, adult male collected by JPRP, Patricio Vinueza and Eduardo  Peña in Finca Palmonte,  Río Negro,  Baños de Agua Santa, Tungurahua, Ecuador (1.4349°S, 78.2629°W; 2485 m alt.) on 14 May 2022. </p>
            <p>Generic placement.</p>
            <p> As defined by Lynch and Duellman (1997), Hedges et al. (2008) and Franco-Mena et al. (2023), the  Pristimantis myersi group (subgenus  Pristimantis Trachyphrynus ) contains species with the following combination of traits: (1) small body size (SVL in females &lt;34.6 mm; in males &lt;20.5 mm); (2) short snout; (3) robust body; (4) Toe V longer than Toe III, Finger I shorter than II; (5) digital discs narrow or slightly expanded (expanded in  P. floridus ); and (6) cranial crests absent. In addition, all species in the group are found on low vegetation or at ground level or even underground. The morphology of the new species agrees with all the aforementioned diagnostic traits; therefore, we place it in the genus  Pristimantis , subgenus  Pristimantis Trachyphrynus . </p>
            <p>Diagnosis.</p>
            <p>(1) Skin of dorsum shagreen, occipital fold evident, dorsolateral folds low in banded individuals or weakly defined in uniform or irregular dorsal colour patterns, skin on venter coarsely areolate (Fig. 2); (2) tympanic annulus and tympanic membrane present, tympanum prominent, sexually dimorphic, tympanum in males 7% SVL, tympanum in females 6% SVL; (3) snout subacuminate in dorsal view (tip of snout pointed), rounded in lateral view (Fig. 2); (4) upper eyelid with several small rounded to subconical tubercles; upper eyelid in males 56% IOD (in females 59%) IOD; cranial crests absent; (5) odontophore processes of the vomer partially concealed, oblique in outline, each processes bearing several small, poorly defined teeth; (6) sphenethmoides with acuminated anterior border dorsally, ventrally present an anteriorly rounded acuminated projected border, posterior border horizontal articulated with frontoparietals; posterior border of frontoparietal present enlarged quadrangular process posteriorly in dorsal view; zygomatic ramus of squamosal short and rounded in dorsal view; procesus cultriform of the parasphenoides reaching posterior level of the vomers, acuminated anterior border (Fig. 8); (7) males with vocal slits and subgular vocal sac; lacking nuptial pads; (8) finger I shorter than II, discs on fingers II-IV rounded (Fig. 3); (9) fingers with slightly visible lateral fringes; (10) ulnar tubercles small, low; (11) heel with small tubercles; outer edge of tarsus with small tubercles, more accentuated in males; (12) inner metatarsal tubercle oval elongated, about twice the size of outer metatarsal tubercle that is rounded; (13) toes lacking lateral fringes, webbing absent; Toe V slightly longer than III, discs slightly expanded (Fig. 3); (14) dorsum brown in several shades, with dark extreme yellow or green tones, females generally darker than males that have usually a banded pattern; banded limbs; venter light brown with dark brown flecks; hidden surfaces of the groin, thighs and armpits pinkish to reddish (in life), chest and throat present light brown tones (Figs 4 - 7); (15) SVL in males, 14.9-16.4 mm (mean = 13.8, SD = 3.9, n = 32), in females, 13.5-19.7 mm (mean = 17.1, SD = 4.41, n = 18); (16) advertisement call with two notes, duration note of 236-293 ms, with intervals of 522-620 ms; call is comprised by tonal sounds of constant frequency, with a slight upward modulation at end of call; (17) dominant frequency at 2.93 kHz, with two partial harmonics, first one with a range of 5.86-5.94 kHz and the second one with a range of 8.79-8.96 kHz; (18) call duration ranging from 1052-1136 ms.</p>
            <p>Comparison with other species</p>
            <p> (Figs 5, 8, 9).  Pristimantis donnelsoni sp. nov. is smaller than its closer congeners, externally is most similar to  P. gladiator (Lynch, 1976) and  P. festae (Peracca, 1904), (Kruskal-Wallis Chi2 = 25.39, p = &lt;0.001, Table 2, Fig. 9), males in  P. gladiator 15.8 mm vs.  P. donnelsoni sp. nov. 13.8 mm; females in  P. gladiator 20.2 vs.  P. donnelsoni sp. nov. 17.1 mm; males in  P. festae 16.4 mm vs.  P. donnelsoni sp. nov. 13.8 mm; females in  P. festae 20.7 vs.  P. donnelsoni sp. nov. 17.1 mm. Additionally, the new species differs in having reddish and pinkish marks on the groin and ventral surfaces in several tones of brown, unlike  P. gladiator which features dark orange marks on a dark brown venter.  Pristimantis donnelsoni sp. nov. has a subacuminated snout in comparison to  P. gladiator which is acuminate. On the other hand, the snout of  P. festae is rounded in dorsal view and subacuminated in  P. donnelsoni sp. nov. Another species, closely related and morphologically similar to  P. donnelsoni , is  P. kayi sp. nov. Although the two new species are extremely similar, differences in the tympanum diameter are evident, TD in  P. donnelsoni sp. nov. 0.9 mm vs.  P. kayi sp. nov. 1.0 mm (Kruskal-Wallis Chi2 = 10.28, p = &lt;0.05*, Table 2, Fig. 8.).  Pristimantis leoni and  P. sirnigeli are found at the Western Andean Cordillera, but they lack distinctive concealed reddish flash marks on the groin that are characteristic of  P. donnelsoni sp. nov. </p>
            <p>Legend: SVL (snout-vent length), HW (head width), HL (head length), EN (eye-nostril distance), IND (internarial distance), IOD (interorbital distance), EW (upper eyelid width), TD (tympanum diameter) eye-nostril, ED (eye diameter), TL (tibia length), HaL (Hand length), FL (foot length), FW (Finger III width), TW (Toe IV length).</p>
            <p> Differences in the skull morphology amongst closely-related species are summarised in Table 3 as follows:  Pristimantis donnelsoni sp. nov. has the posterior border of the frontoparietal with an elongated quadrangular crested process projected posteriorly; while in  P. gladiator , it is irregular and slightly projected posteriorly; in  P. kayi sp. nov. posterior border of frontoparietal, it is irregular and not projected; finally,  P. festae exhibit frontoparietal with rounded posterior border not projected. </p>
            <p> The length and shape of the zygomatic ramus in the squamosal, are short and rounded in  Pristimantis donnelsoni sp. nov.;  P. gladiator presents a short and blunt zygomatic ramus;  P. kayi sp. nov. shows an elongated and sharp zygomatic ramus, reaching level of the maxillae; whereas in  P. festae , zygomatic ramus is very short and blunt, not extending to the level of the maxillae (Fig. 8). </p>
            <p> In ventral view of the skull, differences are evident in  P. donnelsoni sp. nov., the anterior border of sphenethmoides is triangular and posterior border presents horizontal articulations with frontoparietals; in  P. gladiator , anterior border of sphenethmoides is rounded and projected anteriorly, posterior border presents a pair of posterior suboval processes, articulated with frontoparietals; in  P. kayi sp. nov., the anterior border of sphenethmoides is blunt and not projected and the posterior border presents oblique articulation with frontoparietals. </p>
            <p> In  P. donnelsoni sp. nov., the parasphenoid does not reach posterior border of the skull, lateral alary process of parasphenoid is short and does not reach level of maxilla, the cultriform process of the parasphenoid reaches posterior level of vomers and ends in a sharp anterior border, differing from  P. gladiator , in which parasphenoid does not reach the posterior region of vomers with a subacuminated anterior border; in  P. kayi sp. nov., the cultriform process of the parasphenoid reaches the posterior level of vomers and ends in a blunt anterior border, in  P. festae , the sphenethmoides presents an anterior border of cultriform process and does not reach vomers, its anterior border is short and very acute (Table 3, Fig. 8). </p>
            <p>Description of the holotype</p>
            <p>(Figs 2, 3, 4). Adult female (DHMECN 14701) with robust body and short limbs; head slightly longer than wide, not as wide as body, head width 40% of SVL (15.4 mm); head length 41% of SVL; snout short, with a fleshly terminal rounded tip, subacuminated in dorsal view and rounded in lateral view; eye-nostril distance 9.9% of SVL; nostrils narrow higher than long, laterally directed; canthus rostralis angular in dorsal and lateral view; loreal region slightly concave; lips rounded; upper eyelid bearing small low rounded tubercles; snout and interorbital region with scattered small low rounded tubercles; upper eyelid width 49.3% of IOD; tympanic annulus visible on its 3⁄4 part, with the dorsal margins obscured by supratympanic fold; tympanic membrane present, distinct; tympanum diameter 43.6% of eye diameter, one subconic postrictal tubercle surrounded by little low tubercles; rounded tubercles along the supratympanic fold and 2-3 rounded tubercles surrounded by small tubercles on the temporal region; Choanae small, rounded, not concealed by palatal shelf of maxilla; dentigerous processes of vomers low, distinct, oblique in outline, widely separated, positioned posteromedial to choanae; each vomer bearing 2 or 3 teeth; tongue twice as long as wide, free posteriorly on two-thirds of its length. Temporal folds are formed by a row of small low tubercles, extending from the posterior margin of the upper eyelid to the suprascapular region. Skin on the dorsum with scattered, low-rounded tubercles, small folds formed by little rounded tubercles on upper areas of flanks; lower flanks finely granular; skin on the upper surfaces of forelimbs and hind limbs with indistinct rows of low rounded tubercles.</p>
            <p>Skin of the throat, chest, and ventral surfaces and limbs is weakly areolate, venter coarsely areolate; discoidal fold weakly defined; cloacal sheath short; skin in the cloacal region smooth. Row of 2 or 3 rounded ulnar tubercles; palmar tubercles low, outer palmar tubercle bifid, V-shaped, approximately twice as large as elongate ovoid thenar tubercle; subarticular tubercles low, well-defined, rounded in ventral view and flattened in lateral view; supernumerary tubercle at the base of fingers present, indistinct; fingers bearing lateral fringes; Finger I shorter than Finger II; discs on Fingers I and II, rounded and slightly expanded; discs more expanded on Fingers III and IV; ventral pads on fingers well-defined by circumferential grooves on Fingers II, III and IV, not evident on Finger I (Fig. 3).</p>
            <p>Hind limbs robust, tibia length 47.54% of SVL; foot length 47.67% of SVL; upper surfaces of hind limbs with low rounded tubercles; heel bearing one to four low rounded tubercles; the outer surface of tarsus bearing a row of four low rounded tubercles; inner tarsal fold present; inner metatarsal tubercle evident, oval, elongate elliptical, three times size of outer metatarsal tubercle; inner plantar surface smooth; subarticular tubercles poorly defined, rounded in ventral view and flattened in lateral view; toes lacking lateral fringes; webbing between toes absent; discs on toes rounded, slightly expanded; all toes with ventral pads defined by circumferential grooves, less distinct on Toe I; relative lengths of toes: I &lt;II &lt;III &lt;V &lt;IV (Fig. 3); Toe V slightly longer than III, disc of Toe III not reaching medial subarticular tubercle of Toe IV; the distal end of the disc on Toe V reaches the medial subarticular tubercle of Toe IV.</p>
            <p>Measurements of the holotype</p>
            <p>(in mm). Adult female, DHMECN 14701, SVL = 15.4; Tibia Length = 7.3; Foot Length = 7.4; Hand Length = 3.8; Head Length = 6.3; Head Width = 6.2; Eye Diameter = 1.9; Tympanum Diameter = 0.8; Forearm Length = 4.2; Snout Length = 5.3; Tarsus Length = 4.2; Thigh Length = 5.5; Upper Arm Length = 3.4; Interorbital Distance = 2.1; upper Eyelid Width = 1.0; Internarial Distance = 1.5; Eye-Nostril distance = 1.5; Wide Finger III = 0.4; Toe IV Width = 0.4.</p>
            <p>Colour of holotype in life</p>
            <p>(Figs 4, 5, 7). Head, sides of the head, dorsum, flanks and limbs brown, with a large rounded white blotch on the middle of dorsum; light and dark brown banded lips, the broad dark brown supratympanic band extends from the tympanum to level of insertion of arm; flanks darker than dorsum with oblique, pale and diffuse bands; several pinkish-whitish markings scattered near the groin and axilla; limbs banded in light and dark brown tones. Throat and venter dark brown covered with scattered tiny white spots; ventral surfaces of limbs lighter brown with grey tones. Iris silver with a dark copper-coloured horizontal bar.</p>
            <p>Colour of holotype in ethanol</p>
            <p>(Figs 2, 3, 6). Dorsum dark brown with a mid-dorsal white spot; limbs banded with light brown; lips banded; dark supratympanic band delineated below with thin pale line; flanks dark brown to black with pale oblique bands; several white markings near groin and axilla; chin with brown marking outlined with pale shades; throat brown with pale spots; belly dark brown covered with minute white spots; ventral surfaces of limbs and palms light brown with pale spots.</p>
            <p>Osteology of the skull.</p>
            <p>The skull of the adult female paratype DHMECN 4808, is illustrated on its dorsal and ventral surfaces in Fig. 8. We describe main skull bones with diagnostic characters and the key differences are summarised in Table 3. Skull is slightly longer than wide. Dorsally paired nasals overlap the sphenethmoid, the sphenethmoid articulates posteriorly with the frontoparietal, posterior border of frontoparietal present quadrangular projected process posteriorly. A large frontoparietal fontanelle, connected to two parietal fontanelles, is delimited by the sphenethmoid and the frontoparietals.</p>
            <p>In ventral view, anterior margin of the sphenethmoid is acuminated. Palatines overlap with the sphenethmoid and vomers. The vomer is narrowly separated medially; each vomer has three distinctive rami dentigerous process of the vomer reaches the level of the palatine. The parasphenoid present an inverted cross shape; cultriform process with an acute anterior border that reaches level of vomers, the alary processes are directed transversely, partially covering the otic area; the posterior process of the parasphenoid does not reach the foramen magnum. In dorsal view, zygomatic ramus of squamosal presents a short and rounded anterior border extending backwards to the level of the maxilla (Fig. 8).</p>
            <p>Variation</p>
            <p> (Figs 5, 6, 7, 9).  Pristimantis donnelsoni is highly polymorphic and sexually dimorphic species (Suppl. material 1). Females show dark brown dorsal colouration with black to brown banded flanks and tiny white spots; brown with white spots on dorsum; white interorbital line from snout to cloaca, whitish interorbital band, broad brown dorsal mid-dorsal band delineated towards flanks with dark brown to yellowish to black. Some individuals with shades of green. Venter brown in various shades with small whitish spots; some individuals dark brown-grey with a cross pattern along the mid-line and shoulder girdle, white line delineating the outer edge of the tarsus, leg and cloaca. Males are smaller than females and characterised by dorsal banding patterns in shades of grey, brown, cream and yellowish, some individuals with pale dorsolateral lines along the dorsolateral folds; white interorbital bar; hidden groin surfaces and armpits with white or pale markings mottled with shades of grey or brown. Some individuals with shades of green on the dorsum and flanks. </p>
            <p>Call description</p>
            <p> (Fig. 10). The call description of  Pristimantis donnelsoni sp. nov. is based on recordings of a male paratype DHMECN 4774, on 21 March 2007, 19:00 h, made by JPRP, on the northern flank of Tungurahua Volcano, no environmental parameters were obtained. The calling male was sitting on herbaceous plants on the forest floor. The call is comprised by tonal sounds of constant frequency, with a slight upward modulation at the end of the song. It presents a dominant frequency at 2.93 kHz, with two partial harmonics, first one with a range of 5.86-5.94 kHz and the second one with a range of 8.79-8.96 kHz. It has a duration ranging from 1052-1136 ms, with intervals ranging from 4531-5985 ms, emitting at a rate ranging from 8.43-10.60 calls/minute. The calls are composed of two notes. The notes have a duration ranging from 236-293 ms, with intervals ranging from 522-620 ms. </p>
            <p>Distribution and natural history observations.</p>
            <p> Pristimantis donnelsoni sp. nov. is known from seven localities on the southern mountains of the Pastaza drainage, Tungurahua Province, Ecuador (Fig. 11). Three localities are located on the Tungurahua Volcano, whereas the others correspond to nearby protected areas (Chamana Reserve and Cerro Candelaria Reserve, Finca Palmonte and Bosque Protector Hacienda Guamag), all at elevations of 2950-3800 m. The collection localities range from high Andean montane forest, with trees of the genera  Ceroxylum ,  Clusia ,  Weinmannia , and others, between 10-20 m high, with abundant epiphytes, bromeliads orchids, and mosses, to paramo habitat which is mainly grassland with shrubs in the higher part (Fig. 12). Most of the individuals were found sitting on herbs or on leaf litter on the forest floor, with dense ferns and small herbaceous plants, always near ground level (&lt;30 cm above ground). Calling males are active during the day and sporadically call at night. Sympatric species include  Pristimantis aff. devillei ,  P. modipeplus in Tungurahua Volcano,  P. puruscafeum and.  P. marcoreyesi and  P. lacrimosus group sp. nov., in Cerro Candelaria and Chamana. </p>
            <p>Etymology.</p>
            <p> The specific epithet "  Pristimantis donnelsoni " is a noun in the genitive case and a patronym for Don Nelson Palacios (Fig. 12D). We named this new species after him as a tribute to his friendship and collaboration in the first collections of new species described during the last decade and several additional important localities within the upper basin of the Pastaza River. This is a special recognition to him for eternity. </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/85F826E58E0F5EDABF3AA1FF49A9BDDE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Reyes-Puig, Juan Pablo;Urgiles-Merchan, Miguel;Franco-Mena, Daniela;Guayasamin, Juan M.;Batallas, Diego;Reyes-Puig, Carolina	Reyes-Puig, Juan Pablo, Urgiles-Merchan, Miguel, Franco-Mena, Daniela, Guayasamin, Juan M., Batallas, Diego, Reyes-Puig, Carolina (2023): Two new species of terrestrial frogs of the Pristimantis gladiator complex (Anura, Strabomantidae) from the Ecuadorian Andes, with insights on their biogeography and skull morphology. ZooKeys 1180: 257-293, DOI: http://dx.doi.org/10.3897/zookeys.1180.107333, URL: http://dx.doi.org/10.3897/zookeys.1180.107333
10218311988457C8AE068B321C63C959.text	10218311988457C8AE068B321C63C959.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Pristimantis kayi Juan M. Guayasamin, Miguel Urgiles-Merchan, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas & Juan Pablo Reyes-Puig 2023	<html xmlns:mods="http://www.loc.gov/mods/v3">
    <body>
        <div>
            <p> 
Pristimantis kayi Juan M. Guayasamin, Miguel 
Urgiles-Merchan
, Daniela Franco-Mena, Carolina Reyes-Puig, Diego Batallas &amp; Juan Pablo Reyes-Puig
 sp. nov.</p>
            <p>Figs 1, 5, 8, 12, 13, 14, 15, 16, 17</p>
            <p>Type material.</p>
            <p> Holotype. ZSFQ 0775, adult female (Figs 5C, 13, 15 - 17) collected by  José Vieira, David Brito-Zapata, Jefferson Mora, and Carolina Meneses from  Volcán Sumaco, Sumaco National Park, Napo, Ecuador (0.5696°S, 77.5939°W; 2322 m alt.) on 11 September 2018. </p>
            <p> Paratypes (13, 4 ♂ / 9 ♀, Figs 14, 17). ZSFQ 0779, adult female. ZSFQ 773, adult male. ZSFQ 782, adult female with the same data as the holotype, ZSFQ 783, adult female collected by  José Vieira, David Brito-Zapata, Jefferson Mora, and Carolina Meneses from Sumaco National Park, Napo, Ecuador (0.5696°S, 77.5753°W; 2,476 m alt.) on 1 September 2018; ZSFQ 778, adult male, ZSFQ 780 and 781 with same data as ZSFQ 783. DHMECN 14446, adult female collected by Mario Humberto  Yánez Muñoz , JPRP, and DFM from Cerro Mayordomo, Reserva Machay,  Río Negro,  Baños de Agua Santa, Tungurahua, Ecuador (1.3686°S, 78.2693°W 8; 2972 m alt.) on 2 March 2018. DHMECN 15226 and 15237, adult females collected by Kelsey Huisman and Eduardo  Peña from Vizcaya, Reserva Naturetrek, Ulba,  Baños de Agua Santa, Tungurahua, Ecuador (1.396°S, 78.3942°W; 3150 m alt.) on 12 December 2019. DHMECN 15231 and 15233, adult females collected by Kelsey Huisman and Eduardo  Peña from Cerro Mayordomo, Reserva  Ecológica Machay,  Río Verde,  Baños de Agua Santa, Tungurahua, Ecuador (1.368°S, 78.2692°W; 2969 m alt.) on 24 November 2019. DHMECN 16212, adult female collected by JPRP fromVizcaya, Reserva Naturetrek, Ulba,  Baños de Agua Santa, Tungurahua, Ecuador (1.3962°S, 78.3941°W; 3062 m alt.) on May 2021. DHMECN 18440, adult male collected by JPRP, Eduardo  Peña , and Edgar  Martínez , from Los  Mortiños , Leito, Patate, Tungurahua, Ecuador (-1.315235, -78.452793; 3448 m alt.) on 6 July 2022. MZUTI 2209, adult female collected by Juan M. Guayasamin and Lucas Bustamante from Cordillera de los Guacamayos, Napo, Ecuador (0.3769°S, 77.5048°W; 2190-2247 m alt.) in March 2013. MZUTI 2010, 2011, 2013 and 2014 with the same data as MZUTI 2209. </p>
            <p>Generic placement.</p>
            <p> As defined by Lynch and Duellman (1997), Hedges et al. (2008), and Franco-Mena et al. (2023), the  Pristimantis myersi group (subgenus  Pristimantis Trachyphrynus ) contains frogs with the following combination of traits: (1) small body size (SVL in females &lt;34.6 mm; in males &lt;20.5 mm); (2) short snout; (3) robust body; (4) Toe V longer than Toe III, Finger I shorter than II; (5) digital discs narrow or slightly expanded (expanded in  P. floridus ); and (6) cranial crests absent. In addition, all species in the group are found on low vegetation or at ground level, or even underground. The morphology of the new species agrees with all the aforementioned diagnostic traits. </p>
            <p>Diagnosis.</p>
            <p> (1) Skin on dorsum shagreen with small scattered, rounded warts; upper flanks with numerous low warts; dorsolateral and w-shaped occipital folds present (Figs 13 - 15); (2) tympanic membrane and tympanic annulus well differentiated (Fig. 12); tympanic annulus not sexually dimorphic; tympanum in males 7% SVL, tympanum in females 7% SVL; (3) snout rounded in dorsal, protruding in lateral view (Fig. 15); (4) upper eyelid usually bearing one conical or subconical tubercle and many low tubercles; upper eyelid about in males 63% IOD, in females 59% IOD; cranial crests absent; (5) dentigerous processes of the vomer evident, each process bearing 2-3 teeth; (6) sphenethmoides on its dorsal view with short blunt anterior border, is short and blunt in ventral view, posterior border present oblique articulation with frontoparietals; posterior border of frontoparietal with an irregular border not projected in dorsal view; zygomatic ramus of squamosal elongated anteriorly, in dorsal view; procesus cultriform of the parasphenoides reaching posterior level of the vomers, rounded anterior border (Fig. 8); (7) males with vocal slits, nuptial pads absent; (8) finger I shorter than finger II; disc on Finger I not expanded; discs on Finger II-IV slightly expanded (Figs 6, 15); (9) fingers with thin lateral fringes; (10) two or three low ulnar tubercles present; (11) heel with low to conical tubercle; outer edge of tarsus with low conical tubercles; (12) inner metatarsal tubercle oval, about 1.5-2  × the length of round outer metatarsal tubercle; (13) toes with thin lateral fringes; webbing absent; Toe V slightly longer than Toe III; discs slightly expanded (Figs 15, 16); (14) in life, dorsum brown with darker and lighter markings; venter light grey to black, with or without orange to red spots of different sizes; groin with orange to red spots that are more conspicuous in females than males (see Colour variation; Figs 14, 17); (15) SVL in adult males, 11.8-15.6 mm (mean = 13.7, SD = 2.7, n = 30), SVL in adult females 12.3-20.2 mm (mean = 16.3, SD = 5.5, n = 13); (16) call composed of 1 to 7 notes, notes have a mean duration of 36.0  ± 18.3 ms, the mean interval between notes is 43.8  ± 36.2 ms, emitted at a mean rate of 15.7  ± 6.1 notes/second. The notes are composed of 2 to 13 pulses. Pulses have a mean duration of 3.9  ± 1.1 ms, with a mean interval between pulses of 4.2  ± 2.8 ms, emitted at a mean rate of 202.2  ± 119.7 pulses/second; (17) dominant frequency of 3.19  ± 0.05 kHz, with 2-6 partial harmonics in the spectrogram; (18) call duration of 165.1  ± 130.1 ms. </p>
            <p>Comparison with other species</p>
            <p> (Figs 5, 8, 9).  Pristimantis kayi sp. nov. is most similar to  P. gladiator (Lynch, 1976),  P. festae (Peracca, 1904), and  P. donnelsoni sp. nov.  Pristimantis kayi sp. nov. differs from  P. gladiator and  P. festae by being smaller (Kruskal-Wallis Chi2 = 25.39, p = &lt;0.001, Table 2, Fig. 9). Additionally,  P. kayi sp. nov. differs from  P. festae by having orange to red spots on the groin (white to reddish large spots in  P. festae ; Fig. 5) and slightly expanded discs on outer fingers (discs not expanded in  P. festae ).  Pristimantis kayi sp. nov. is similar to the allopatric  P. donnelsoni sp. nov.; however, the diameter of the tympanum in males is larger in  P. kayi sp. nov. compared to  P. donnelsoni sp. nov. and the other closely-related species (i.e.  P. festae and  P. gladiator ) (Kruskal-Wallis Chi2 = 10.28, p = &lt;0.05*, Table 2, Fig. 9). The most conspicuous differences in the skull morphology between the two new species are listed below (Fig. 8): in  P. kayi sp. nov., the parasphenoid is much more developed than in  P. donnelsoni sp. nov., with its lateral alary processes almost reaching the squamosal; also, the palatine bones are shorter in  P. kayi sp. nov. than in  P. donnelsoni sp. nov. Finally, the zygomatic ramus of the squamosal is much longer in  P. kayi sp. nov. than in  P. donnelsoni sp. nov. (Fig. 8). </p>
            <p>Description of the holotype</p>
            <p>(Figs 13, 15, 16). Adult female (ZSFQ 0775) robust body; head slightly longer than wide, not as wide as body, head width 35.27% of SVL (19.7); head length 38% of SVL; snout rounded in dorsal view, protruding in lateral view; eye-nostril distance 9% of SVL; with small papilla at tip (Fig. 1); in lateral view, distinct rostral ridge; loreal region slightly concave; nostrils protruding, laterally directed; interorbital area flat, wider than upper eyelid (upper eyelid width 62% of OID); cranial ridges absent; upper eyelid with one conical and several non-conical tubercles; tympanic membrane well defined, pigmented like surrounding skin; tympanic ring distinct, round; supratympanic fold present, obscuring anterodorsal and posterodorsal edges of the ring; tympanic diameter 46% of eye length; several low to conical tubercles situated in the area just posterior to tympanum. Choanae are small, with an oval to square shape, not concealed by the palatal shelf of maxillary; the dentigerous process of the vomer oblique, widely separated, posteromedial to choanae, each bearing two small teeth; tongue slightly longer than wide, granular, with a conspicuous notch along the posterior border.</p>
            <p>The skin of the head is greyish; the dorsum greyish, with scattered small tubercles, some of which are aligned over a W-shaped occipital marking, forming folds; upper flanks with numerous low warts; venter slightly areolate; discoidal fold absent; the cloacal sheath is absent; fingers with thin lateral bangs; length of fingers I &lt;II &lt;IV &lt;III; palmar tubercle round, thenar tubercle oval (Fig. 16); subarticular tubercles round, not prominent; supernumerary palmar tubercles not evident; disc sheath of Finger I not expanded; those of Fingers II-IV slightly expanded; all disc sheaths with nearly elliptical ventral pads defined by grooves (Fig. 16).</p>
            <p>Hind limbs relatively robust; tibia length 47% SVL; foot length slightly smaller than tibia length (foot length 44% SVL); tarsal tubercles present; small, conical tubercle on heel; toes with narrow lateral fringes (Fig. 16); subarticular tubercles round, not prominent; inner metatarsal tubercle oval, two times the size of outer tubercle; few low supernumerary plantar tubercles present (Fig. 16); disc covers of Toes slightly expanded; toes with defined pads; disc pads nearly elliptical; toe lengths I &lt;II &lt;III &lt;V &lt;IV (Fig. 16); the tip of Toe V reaches the middle level of penultimate subarticular tubercle of Toe IV; the tip of Toe III almost reaches the proximal border of penultimate subarticular tubercle of Toe IV.</p>
            <p>Measurements of the holotype</p>
            <p>(in mm). Adult female, ZSFQ 0775. SVL = 19.7; Tibia Length = 9.3; Foot Length = 8.6; Hand Length = 5.0; Head Length = 7.5; Head Width = 7.0; Eye Diameter = 2.4; Tympanum Diameter = 1.1; forearm length = 4.4; snout length = 5.6; Tarsus length = 5.3; Thigh Length = 8.3; Upper arm Length = 3.1; Interorbital Distance = 2.6; Upper Eyelid Width = 1.6; Internarial Distance = 2.1; Eye-Nostril distance = 1.8; Finger III Width = 0.8; Toe IV Width = 0.3.</p>
            <p>Colour of holotype in life</p>
            <p>(Fig. 13). Head, sides of the head, dorsum, flanks and limbs brown, with darker interorbital bar, lips banded with light and dark brown; broad dark brown supratympanic band extending from region anterior to tympanum to before arm insertion. Flanks lighter than dorsum with oblique, dark bands, several scattered whitish markings near the groin and a white spot in the axilla; limbs banded in shades of dark brown. The throat and belly are dark brown and covered with scattered small yellowish-white spots; the lower ventral part has orange spots; the ventral surfaces of the forelimbs are lighter brown. Iris golden with dark brown reticulations, with a reddish-copper horizontal bar.</p>
            <p>Colouration of holotype in ethanol</p>
            <p>(Figs 15, 16). The holotype has the following colour pattern: head pale brown with darker interorbital and labial bars; black supratympanic stripe; dorsum brown with several darker markings. Flanks grey to black with minute white spots and lighter diagonal stripes. Groin black with large white spots. The dorsal surfaces of limbs are pale brown with dark brown bars. Cloacal region pale brown, delimited by supracloacal black stripe. The anterior and posterior surfaces of the thighs are brown with several medium to large cream spots. Throat and chest cream-brown with darker marks on the centre (Fig. 15). Venter dark grey, with minute to large white spots. Palms and soles are mostly cream, with some pigmentation of Finger IV and Toe V (Fig. 16).</p>
            <p>Osteology of the skull.</p>
            <p>The skull of the adult female paratype DHMECN 14447 is illustrated on its dorsal and ventral surfaces in Fig. 8. We describe the main skull bones with diagnostic characters in Table 3. Skull is slightly longer than wide. Dorsally paired nasals overlap the sphenethmoid; the sphenethmoid articulates posteriorly with the frontoparietals, posterior border of frontoparietal is irregular and not projected. A large frontoparietal fontanelle, connected to two parietal fontanelles, is delimited by the sphenethmoid and the frontoparietals.</p>
            <p>In ventral view, anterior margin of the sphenethmoid is rounded. The palatines are relatively short and overlap the sphenethmoid on its anterior portion. Vomers are narrowly separated medially; each vomer has three distinctive rami; the dentigerous process of the vomer does not reach the level of the palatine and almost contacts the tip of the cultriform process of the parasphenoid. The large parasphenoid presents the shape of an inverted cross; the alary processes are directed transversely, partially covering the otic area and almost reaching the squamosal; the posterior process of the parasphenoid almost reaches the foramen magnum. In the dorsal view, the long zygomatic ramus of the squamosal presents an elongated and acuminated anterior border that extends towards the level of the maxilla (Fig. 8).</p>
            <p>Variation</p>
            <p> (Figs 14, 17). Dorsal surfaces with various shades of brown, with or without darker bands or bars. Darker facial markings such as labial, canthal and interorbital bars are usually present. Dorsal mark is an inverted  “V” in the coccygeal region; other patterns include arrangements of thin lines arranged longitudinally. Dorsal surface of hands and feet with irregular brown spots. The fore- and hind limbs are banded with dark brown, separated by light brown interspaces. Flanks with various shades of brown, usually lighter than dorsum. Hind surfaces of thighs with black or dark brown bars and cream interspaces. Groin with orange and sometimes yellowish-white spots on an orange background. Belly brown; sometimes mottled. The belly may have numerous orange to yellowish-white spots. In some specimens, there is a yellowish-white cross on the belly. Iris is golden to bronze with fine black reticulations and a reddish-copper horizontal middle stripe. </p>
            <p>Call description</p>
            <p> (Fig. 18). The description is based on the call of an adult male (MZUTI 852), recorded during the night (21:40 h) by Italo Tapia on 7 June 2012, at the Cordillera de los Guacamayos, Province of Napo, Ecuador. The male was calling from leaf litter at ground level. The air temperature was 12.2 °C. The recording consisted of 21 calls, 37 notes and 856 pulses. The call of  Pristimantis kayi sp. nov. is composed of the emission of several elements with different characteristics, ranging from single notes to pulsed notes. The call (Fig. 1) has a mean dominant frequency of 3.19  ± 0.05 kHz, with 2-6 partial harmonics in the spectrogram. It has a mean duration of 165.1  ± 130.1 ms. The mean interval between calls is 4565.6  ± 5742.3 ms, emitted at a mean rate of 32.6  ± 23.2 calls/minute. Calls are composed of 1 to 7 notes. Notes have a mean duration of 36.0  ± 18.3 ms. The mean interval between notes is 43.8  ± 36.2 ms, emitted at a mean rate of 15.7  ± 6.1 notes/second. The notes are composed of 2 to 13 pulses. Pulses have a mean duration of 3.9  ± 1.1 ms, with a mean interval between pulses of 4.2  ± 2.8 ms, emitted at a mean rate of 202.2  ± 119.7 pulses/second (Table 4). The call of  Pristimantis kayi sp. nov. does not present stereotyped notes. Due to their structural elements (i.e. notes-pulses), notes can be classified into three different types: Single note calls (CNS; Fig. 18A), Pulsed note calls (CNP; Fig. 18B) and Complex calls (CNP; Fig. 18C), which contain single (non-pulsed) and pulsed notes in its structure. Temporal and spectral measurements of the different call types are shown in Table 4. </p>
            <p>Distribution and natural history observations.</p>
            <p> Pristimantis kayi sp. nov. has been recorded from the Cordillera de Guacamayos,  Volcán Sumaco (Napo Province) and Cerro Mayordomo, Machay Reserve, Naturetrek Vizcaya Reserve, Leito Reserve (Tungurahua Province) at an elevational range of 2190-3600 m a.s.l. (Fig. 11). Calling males were heard during the day and night. During the night, frogs of  Pristimantis kayi sp. nov. were found on the forest floor or perched on low vegetation (always below 60 cm from ground level), including shrubs, grasses (  Neurolepis sp.), ferns and  Selaginella . The species has been found in primary and disturbed Andean forests, with trees of the genus  Weinmannia ,  Clusia ,  Alnus and others, of 10-15 metres (Fig. 12). </p>
            <p>Etymology.</p>
            <p> The specific epithet "  Pristimantis kayi " is a noun in the genitive case and a patronym for Andreas Kay, a German physicist, biologist and friend who spent much of his life documenting and exploring biodiversity, contributing substantially to the conservation of Ecuadorian forests (Fig. 12). Andreas was part of the team that created  Fundación EcoMinga’s Dracula Reserva in north-western Ecuador. Additionally, his work shed light on understanding the diversity and endemism patterns of orchids in the Llanganates-Sangay Ecological Corridor, where he passed away in 2019. Some of his amazing photographs can be seen here: https://www.flickr.com/photos/andreaskay/albums/. </p>
        </div>
    </body>
</html>
	https://treatment.plazi.org/id/10218311988457C8AE068B321C63C959	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Reyes-Puig, Juan Pablo;Urgiles-Merchan, Miguel;Franco-Mena, Daniela;Guayasamin, Juan M.;Batallas, Diego;Reyes-Puig, Carolina	Reyes-Puig, Juan Pablo, Urgiles-Merchan, Miguel, Franco-Mena, Daniela, Guayasamin, Juan M., Batallas, Diego, Reyes-Puig, Carolina (2023): Two new species of terrestrial frogs of the Pristimantis gladiator complex (Anura, Strabomantidae) from the Ecuadorian Andes, with insights on their biogeography and skull morphology. ZooKeys 1180: 257-293, DOI: http://dx.doi.org/10.3897/zookeys.1180.107333, URL: http://dx.doi.org/10.3897/zookeys.1180.107333
