Pristimantis normaewingae, Reyes-Puig & Yánez-Muñoz & Libke & Vinueza & Carrión-Olmedo, 2024

Reyes-Puig, Juan Pablo, Yánez-Muñoz, Mario H., Libke, Zane, Vinueza, Patricio & Carrión-Olmedo, Julio C., 2024, Phylogenetic diversity of the Pristimantis anaiae species group, with a description of a new species of Pristimantis (Anura, Strabomantidae) from Cerro Candelaria, Tungurahua, Ecuador, Evolutionary Systematics 8 (1), pp. 101-118 : 101-118

Pristimantis normaewingae sp. nov.

Figs 1 , 2 , 3 English Common name: Norma Ewing Rain Frog. Spanish Common name: Cutín de Norma Ewing .

Holotype.

Adult female. Ecuador; Tungurahua Province, Cerro Candelaria Private Protected Area ; 1 ° 27.410 ' S, 78 ° 18.192 ' W, 3042 m a. s. l.; collected by JPRP, PV, Darwin Recalde, Eduardo Peña, Paulette Benavides and Nantar Kuja on 23 November 2022; on a shrub branch covered with moss at 170 cm above the ground; GenBank: PP 102395 , PP 102423 ; DHMECN 18409 .

Paratypes.

Ecuador • (6 ♀, 23 ♂, 9 juveniles); four adult females ( DHMECN 18861 , DHMECN 18863 , DHMECN 18865 , DHMECN 18868 ); from the same protected area near holotype locality; 1 ° 26.301 ' S, 78 ° 18.105 ' W; alt. 2626 m; collected by JPRP, Evelyn Toapanta, Elizabeth Chicaiza Toa and Luis Recalde on 22–23 February 2023 ; Two adult females ( DHMECN 4849 , and DHMECN 4846 ); from the same locality as the holotype; 1 ° 26.399 ' S, 78 ° 17.991 ' W; alt. 2700 m; 5 May 2008; collected by JPRP and Luis Recalde .

Thirteen adult males ( DHMECN 18392 , DHMECN 18396 , DHMECN 18393 , DHMECN 18397 , DHMECN 18401 , DHMECN 18402 , DHMECN 18403 , DHMECN 18404 , DHMECN 18406 , DHMECN 18407 , DHMECN 18411 , DHMECN 18412 , DHMECN 18414 ), collected in the same locality and date as the holotype . Two adult males ( DHMECN 18862 , DHMECN 18896 ); collected in the same locality and date as DHMECN 18861 . Four adult males ( DHMECN 17188 , DHMECN 17189 , DHMECN 17191 , DHMECN 17192 ); collected in the same locality as the holotype by JPRP and ZL on 18 January 2021 . Two Adult males ( DHMECN 4847 , DHMECN 4975 ); collected in the same locality and data as DHMECN 4849 .

Female juvenile ( DHMECN 17190 ), collected in the same locality and data as DHMECN 17188 . Five male juveniles ( DHMECN 18398 , DHMECN 18400 , DHMECN 18405 , DHMECN 18408 , DHMECN 18413 ); collected in the same locality and data as the holotype . Male juvenile ( DHMECN 18864 ); collected in the same locality and data as DHMECN 18861 .

Diagnosis.

We assign the new species to the genus Pristimantis and the Pristimantis anaiae species group based on molecular phylogeny (Fig. 1 ). Pristimantis normaewingae sp. nov. is characterized by the following combination of characters: (1) skin of dorsum shagreen with small low tubercles, skin on throat areolate, skin on belly strongly areolate bearing scattered tubercles; (2) discoidal and thoracic fold present; (3) dorsolateral folds fragmented, instead a dorsolateral row of enlarged tubercles is present in the posterior back, more evident in males; (4) tympanic membrane absent, tympanic annulus present small (on average, 40.99 % of eye diameter); (5) supratympanic fold present with low tubercles, a pair of prootic and occipital tubercles aligned with several subconical tubercles aligned forming a “ W ” in the scapular region; (6) several subconical and rounded postrictal tubercle present; (7) snout short and subacuminate in dorsal and rounded in lateral view, with 4 tubercles aligned with canthal region forming a cross in dorsal view; (8) upper eyelid with one conical tubercle and few lower tubercles; (9) cranial crests absent; (10) Dentigerous processes of vomers present, oblique, separated; vocal slits and nuptial pads absent; (11) Finger I shorter than Finger II, discs of digits expanded to broadly expanded, truncate; (12) fingers with narrow lateral fringes, all fingers with elongated and thin hyperdistal tubercles; (13) low and rounded ulnar tubercles, more conspicuous in males, low ulnar fold evident ventrally; (14) heel bearing a low subconical or rounded tubercle; row of subconical or rounded tubercles along external tarsus; (15) toes with broad lateral fringes, basal webbing absent, all toes with elongated hyperdistal tubercles; (16) Toe V much longer than Toe III, Toe condition (Toe III surpasses the distal border of the distal subarticular tubercle of Toe V and reaches the distal border of the penultimate tubercle of Toe IV; Toe V reaches towards the outermost border of the distal tubercle of toe IV); (17) In life, dorsal coloration is composed by irregular marks mixed with banded pattern with different tones of brown, yellow and green; dark canthal, subocular and supratympanic stripes present, middorsal oval or irregular dark marks delineated by pale lines, surrounded by irregular contrasting marks, flanks with brown or greenish-brown tones, dark incomplete oval marks in sacral region with thin light borders are present on each side. Ventral surfaces of the belly, throat, and limbs are marmorated with dark brown and cream or gray marbling. In males, light reddish or pink flecks are slightly evident in palms and soles, and part of ventral portions of limbs, while in females reddish or pink tones are absent in such ventral surfaces; the iris is light blue with a horizontal wide copper medial band and black reticulations; light pale sclera; (18) SVL in females, 25.0– 30.6 mm (mean = 28.14, SD = 1.94, n = 7); SVL in adult males, 16.7–22.27 mm (mean = 19.68, SD = 2.49, n = 20); Tables 3 , 4 .

Comparison with other species.

(Fig. 4 , Table 4 ) Pristimantis normaewingae sp. nov. can be distinguished from all seven species of the Pristimantis anaiae species group by having incomplete oval marks in sacral region, a row of enlarged dorsolateral tubercles and a light blue iris with a copper horizontal wide medial band with black reticulations; while the other species present different iris patterns, complete oval marks in sacral region and lacks dorsolateral row of enlarged tubercles like P. anaiae , P. resistencia , P. glendae and P. venegasi , with the exception of P. kunam and P. maryanneae , that respectively present larger and smaller dorsolateral row of enlarged tubercles. P. kunam is the smallest species of the group (SVL: 14.66), however, Pristimantis normaewingae sp. nov. interspecific SVL variation overlaps with the other species of the group, and has less conspicuous ulnar tubercles and light brown venter in comparison with the conspicuous conical ulnar, and tarsal tubercles, dark greenish with dark flecks body coloration and bronze iris with black reticulations of P. kunam . P. maryanneae present less tuberculated skin, tympanum absent, small and inconspicuous row of dorsolateral tubercles, and pale ventral surfaces than P. normaewingae sp. nov. that has contrasting body and iris coloration, tympanum hidden beneath the skin with evident tympanic anulus and more tuberculated skin. Detailed comparisons of diagnostic characters between all species of Pristimantis anaiae group are shown in Table 4 .

Description of the holotype.

(Figs 2 , 3 ) Adult female ( DHMECN 18409 ). Head wider than long, slightly wider than the body, snout acuminate in dorsal view, short and rounded in profile; canthus rostralis slightly rounded in dorsal view, concave in lateral view, four rounded tubercles aligned in a cross pattern in dorsal view; loreal region slightly concave; cranial crests absent, upper eyelid with a subconical tubercle surrounded by few smaller tubercles, interorbital tubercle present; tympanic annulus oval distinct beneath the skin; tympanic membrane absent; several rounded low postrictal tubercles and irregular warts; dark supratympanic fold present, distinct, covered with rounded subconical tubercles.

Dentigerous processes of vomers present, oblique, separated, posteromedial to choanae; each vomer bearing several inconspicuous small teeth. Skin on dorsum shagreen with small low tubercles, skin on flanks shagreen with small low tubercles, dorsolateral folds fragmented, instead a dorsolateral rows of rounded low tubercles are present more evident in the posterior back, discoidal and thoracic fold present; skin on throat coarsely areolate with some scattered low warts, chest and belly areolate with some low rounded tubercles; ventral surfaces of thighs areolate; skin in upper cloacal region strongly areolate; row of low subconical ulnar tubercles present, a low ulnar fold is evident ventrally; palmar tubercles prominent and elevated, outer palmar tubercle large bifid “ horseshoe ” shape well differentiated, thenar tubercle is oval and half size than the outer; subarticular tubercles well-defined, pronounced round in ventral and lateral view; hyperdistal tubercles flattened in ventral and lateral view; indistinct supernumerary tubercles; lateral fringes on fingers wide; Finger I shorter than Finger II; disc on Finger I slightly expanded and oval, discs on Fingers, II, III and IV widely expanded truncate; pads on all fingers well defined and surrounded by circumferential grooves (Fig. 3 ).

Hindlimbs slender; upper surfaces of hindlimbs with scattered low tubercles; posterior and ventral surfaces of thighs rugose; heel bearing low subconical tubercle and smaller surrounding; outer surface of tarsus present a row of low subconical tubercles; tarsal fold not evident; well-defined inner metatarsal tubercle, large oval, elevated in lateral view, outer metatarsal prominent rounded about third size of the inner; plantar surface with subarticular tubercles rounded and well defined, subarticular tubercles well-defined, slightly prominent and rounded; hyperdistal tubercles flattened in ventral and lateral view; toes with wide lateral fringes, widened on Toes IV – III – II – I; basal webbing between toes absent; discs nearly as large as those on fingers, expanded in all toes specially on Toes III, IV and V, Toe I oval; all discs have pads surrounded by well-defined circumferential grooves; relative lengths of toes I < II < III < V < IV; Toe V much longer than Toe III (Toe III reach the hyper distal tubercles of Toe V and reaches the distal border of the medial subarticular tubercle of Toe IV; Toe V reaches the proximal border of the distal tubercle of Toe IV) (Fig. 3 )

Measurements of the holotype (in mm). SVL = 30.56; Tibia Length = 15.93; Foot Length = 14.91; Hand Length = 9.82; Head Length = 11.36; Head Width = 11.15; Eye Diameter = 3.69; Tympanum Diameter = 1.79; Forearm Length = 7.09; Snout Length = 4.75; Tarsus Length = 8.59; Thigh Length = 12.72; Upper Arm Length = 5.86; Interorbital Distance = 3.85; upper Eyelid Width = 2.59; Internarial Distance = 2.0; Eye – Nostril distance = 2.88; Finger III width = 1.88; Toe IV Width = 1.75.

Color of holotype in life.

(Fig. 5 ) Dorsal coloration dark brown with diagonal bands on flanks and limbs, anterior snout light brown, dark subocular and supratympanic marks, W-shaped dark blotch present on scapular region, underlined by white lines. Flanks with diagonal dark brown bands, surrounded by pale lines, hidden surfaces of the groin and thighs brown. Tip of the digits with pale color. All ventral surfaces are light brown marbled or marmorated with darker brown tones. Iris coloration pale gray with light blue at peripherical surfaces, extensive fine dark reticulations and medial horizontal copper stripe is present, white sclera.

Color of holotype in preservative.

(Fig. 6 ) Dorsal coloration of body and limbs composed of a dark brown with diagonal dark bands delineated with thin white lines, incomplete oval sacral marks delineated by white on the back, bold dark scapular chevron delineated by pale line, tips of the digits with pale marks. Ventral surfaces are light brown and marmorated with dark brown. Pale disc surfaces on hands and feet.

Variation.

(Figs 5 , 6 ) Morphometric variations are shown in Table 3 . In preservative, the new species presents a wide gradient of banded patterns in dorsal and limb coloration, with backgrounds ranging from pale cream tones, different dark brown tones, and even greenish with orange tones; dark marks on the flanks are delineated by thin pale lines. Ventrally specimens vary with different tones of light brown peppered with gray flecks, especially on the throat, heavily marmorated with dark brown irregular marbling; some specimens present white dots irregularly dispersed on the body.

In life, polychromatic, sexual dimorphism. In dorsal view, females dark brown to pale brown with scapular and sacral marks less conspicuous than males. Iris coloration pale gray with a medial horizontal line reddish brown finely dark reticulated, white sclera ( DHMECN 18409 , DHMECN 18863 ). Males with distinctive dorsal and flank coloration ranging from reddish brown to orange brown. P osterior surface of head and scapular region ranging from light brown to yellowish green l ines surrounding marks on the of flanks ranging from white to pale ochre. Sacral marks ranging from reddish, orange, and dark brown. Iris variation in males with blueish gray with a coppery ( DHMECN 18400 , 18392) or faint reddish brown medial horizontal line ( DHMECN 18404 ). Ventrally, males and females with a grayish brown background with dark brown blotches, yellow cream, or grayish cream. Juveniles with pale or dark and light brown, interorbital region dark brown ( DHMECN 18996 ) or with interorbital line orange brown ( DHMECN 17191 ).

In life, Pristimantis normaewingae sp. nov. shows some variation in tuberculation. Males show eyelids, interorbital and scapular tubercles elevated and conical ( DHMECN 18400 , 18392, 18996). Females only have small, rounded, and conical tubercles ( DHMECN 18409 , DHMECN 18863 ). Some individuals show plasticity in the state of tubercles texture under different environmental conditions, ranging from elevated spiny to relaxed low tuberculated state, additionally preservation could reduce protuberance of tubercles.

Etymology.

The specific epithet normaewingae is a noun in the genitive case and is a patronym in honor of Norma Ewing, a passionate conservationist and animal rescue volunteer from United States. Her belief that all species, and especially frogs, are interconnected, is rooted in her early childhood surrounded by nature living and working on a farm. She has always cared for animals of all shapes and sizes. As a volunteer with animal rescues, she was admired for her ability to nurture orphaned and injured animals.

Distribution, natural history observations, and conservation status.

(Figs 7 , 8 ) Pristimantis normaewingae sp. nov. is known only from the type locality in the Cerro Candelaria Protected Area at an elevation of 2550 to 3150 m. The collection site is eastern Andean cloud forest with a dominance of bromeliads in the forest floor and in the branches of the trees and shrubs, Clusia trees, palms, and Weinmannia , with a high density of epiphytes, mosses, ferns, lichens, and orchids. Sympatric species found with the new species were Pristimantis gladiator complex, P. tungurahua , P. marcoreyesi , and Niceforonia sp.

Because of the lack of records, evidence of population status, and geographic distribution, we recommend assigning Pristimantis normaewingae sp. nov. to the Data Deficient IUCN Red List Category (based on IUCN, 2019 guidelines), however, the only known population is protected within Cerro Candelaria Private Protected Area.

Figures

Figure 1. Phylogenetic relationships among the Pristimantis anaiae species group, with the inclusion of Pristimantis normaewingae sp. nov., Pristimantis maryanneae, and two candidate species (Pristimantis sp. 1, P. sp. 2).

Figure 2. Preserved holotype (DHMECN 18409, adult female) of Pristimantis normaewingae sp. nov, in dorsal, ventral, and lateral views. Photographs JPRP.

Figure 3. Ventral views of hand (left) and foot (right) of female holotype (DHMECN 18409) of Pristimantis normaewingae sp. nov. Photographs JPRP.

Figure 4. Life comparison between members of Pristimantis anaiae species group. A. P. normaewingae sp. nov. (DHMECN 18400, male paratype); B. P. normaewingae sp. nov. (DHMECN 18409, female holotype); C. P. kunam (QCAZ 56438, male holotype, ); D. P. resistencia (QCAZ 66519, female holotype); E. P. anaiae (DHMECN 16859, male); F. P. maryanneae (DHMECN 18227, male); G. P. glendae (QCAZ 56437, adult male); H. P. venegasi, (MZUTI 6571, female). Arrows show area of the oval marks in sacrum, proposed diagnostic of the group by Ortega et al. 2022. Photographs JPRP, MYM, JCC, Jorge Brito, and Bioweb.

Figure 5. Life coloration variation of the type series of Pristimantis normaewingae sp. nov. A 1 – A 3. DHMECN 18863, adult female; B 1 – B 3. DHMECN 18404, adult male; C 1 – C 3. DHMECN 17191, juvenile; D 1 – D 3. DHMECN 18996 male juvenile. Photographs JPRP and ZL.

Figure 6. Color pattern variation in preserved specimens of Pristimantis normaewingae sp. nov. A. DHMECN 18400, male; B. DHMECN 18393, male; C. DHMECN 18222, male; D. DHMECN 18223, female; E. DHMECN 18228, male; F. DHMECN 18212, male. Photographs JPRP.

Figure 7. Distribution map of the Pristimantis anaiae species group including a new species from the mountains south of the Pastaza River.

Figure 8. Pristimantis normaewingae sp. nov. in its habitat at Cerro Candelaria. A. DHMECN 18868, female; B. DHMECN 18401, male on Neurolepis aristata (Poaceae) at night; C. DHMECN 17188, juvenile on bromeliad at night.; D. DHMECN 18400, male on leave of shrub; E. habitat at the type locality of the new species in Cerro Candelaria. Photographs JPRP and ZL.