Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species Author Paez, Nadia B. Author Ron, Santiago R. text ZooKeys 2019 868 1 112 http://dx.doi.org/10.3897/zookeys.868.26766 journal article http://dx.doi.org/10.3897/zookeys.868.26766 1313-2970-868-1 ED7FD98F964D402FAB705FC9B4CA4851 679C0640C93B545BAD44B1013B0B12E5 Pristimantis lutzae sp. nov. Common name. English: Lutz's Rain Frog. Spanish: Cutin de Lutz. Holotype. QCAZ 37546, an adult female from Mazan Reserve, Azuay Province, Ecuador ( 2.8689S , 79.1148W , 3047 m), collected by Luis A. Coloma, Santiago R. Ron, Italo G. Tapia, Ernesto Arbelaez , and Robin Moore on September 2007. Figure 20A . Figure 20. Holotypes of Pristimantis lutzae sp. nov. and P multicolor sp. nov. Photographs of preserved holotypes of A P. lutzae (QCAZ 37546, female) and B P. multicolor (QCAZ 47213, male). Dorsal view on the left, ventral view on the right. Specimens are shown at the same scale. Paratypes (32: 14 males, 13 females, 5 juveniles). Ecuador: Azuay Province: QCAZ 32785, adult female, QCAZ 32786, QCAZ 32788, QCAZ 32791, adult males, QCAZ 32792, juvenile female, from Yanuncay-Irquis Protected Forest, Paramo de Quimsacocha ( 3.0398S , 79.2147W , 3758 m), collected by Andres Merino-Viteri, Lorena E. Falconi , David Salazar-V, Paula Pena , Monica Paez , Ernesto Arbelaez , and Juan Daniel Jaramillo in December 2006; QCAZ 37509, adult female, from Mazan Reserve ( 2.8748S , 79.1292W , 3115 m), collected by Luis A. Coloma, Santiago R. Ron, Italo G. Tapia, Ernesto Arbelaez and Robin Moore in September 2007; QCAZ 37545, QCAZ 37547, QCAZ 37566, adult females, QCAZ 37561, QCAZ 37564, adult males, QCAZ 37550-551, juvenile females, collected with the holotype; QCAZ 37571, adult female, QCAZ 37570, adult male, from Mazan Reserve ( 2.8752S , 79.1292W , 3189 m), collected by Luis A. Coloma, Santiago R. Ron, Italo G. Tapia, Ernesto Arbelaez and Robin Moore in September 2007; QCAZ 51736, adult female, from San Vicente ( 2.7953S , 78.6981W , 3044 m), collected by Omar Torres, Vanessa Aguirre, Simon Lobos, Fernando Ayala and Estefania Boada in March 2011; QCAZ 53728, juvenile, from Cajas National Park, El Capo, Laguna Toreadora ( 2.7785S , 79.2453W , 4100 m), collected by Santiago R. Ron, Andres Merino, Fernando Ayala, Teresa Camacho and Martin Cohen on July 19, 2011. Canar Province: QCAZ 27467, QCAZ 27469, QCAZ 27534, adult females, QCAZ 27470-471, adult males, QCAZ 27472, juvenile female, from Mazar Wildlife Reserve, Rumiloma ( 2.5746S , 78.7455W , 3400 m), collected by Martin R. Bustamante, Joseph Mendelson and Michelle Cummer in February 2004; QCAZ 27521, adult male, from Mazar Wildlife Reserve, Rumiloma ( 2,5612S , 78.7336W , 3550 m), collected by Martin R. Bustamante, Joseph Mendelson and Michelle Cummer in February 2004; QCAZ 27596-597, adult females, from La Libertad ( 2.5466S , 78.6984W , 2895 m), collected by Martin R. Bustamante, Joseph Mendelson and Michelle Cummer in February 2004; QCAZ 47211, adult male, from Guallicanga ravine ( 2.4321S , 78.9022W , 3960 m), collected by Paola Mafla-Endara, Silvia Aldas-Alarcon and Freddy Velasquez-Alomoto in December 2009; QCAZ 56182, adult female, QCAZ 56183-186, adult males, from Charon Ventanas Community Tourism Center ( 2.6471S , 78.8905W , 3300 m), collected by Andrea Manzano, Paulina Romero, and Leonardo Negrete in July 2013. Diagnosis. A species of Pristimantis having the following combination of characters: (1) skin on dorsum shagreen to tuberculate with scattered low tubercles; thin middorsal fold present or absent; dorsolateral folds absent; flanks tuberculate, tubercles larger than those on dorsum, with or without scattered warts; lateral fold present or absent; skin on venter coarsely areolate; discoidal fold present or absent; (2) tympanic membrane and tympanic annulus prominent, its upper and posterolateral margin covered by supratympanic fold; (3) snout moderately long, round to subacuminate in dorsal, rounded in lateral view, with or without a small papilla at the tip; (4) upper eyelid bearing a small, rounded tubercle, surrounded by several lower tubercles; cranial crests absent; (5) dentigerous processes of vomers prominent, oblique, moderately separated, posteromedial to choanae; (6) vocals slits, vocal sac, and nuptial pads present in adult males; (7) Finger I shorter than Finger II; discs of digits expanded, elliptical to truncate; (8) fingers with broad lateral fringes; (9) ulnar tubercles small, distinct; (10) heel bearing a low rounded tubercle, surrounded by several smaller tubercles; inner and outer edge of tarsus bearing a row of small tubercles; short inner tarsal fold; (11) inner metatarsal tubercle elevated, ovoid, three times the size of round outer metatarsal tubercle; supernumerary tubercles indistinct; (12) toes with broad lateral fringes; basal webbing present; Toe V longer or much longer than Toe III (disc on Toe III reaches the middle of penultimate subarticular tubercle on Toe IV or slightly exceeds its distal edge, disc on Toe V reaches the middle of distal subarticular tubercle on Toe IV or slightly exceeds its distal edge); toe discs smaller than those on fingers, truncate to elliptical ( Fig. 8A ); (13) in life, dorsum light, orangey or dark brown; head with black or brown supratympanic, canthal, and interorbital stripes; groins, anterior and posterior surfaces of thighs pinkish, purplish or reddish brown, suffused or not with orange, with cream or light brown spots; ventral surfaces of body white to cream with or without brown reticulations; vocal sac in males yellow; iris golden to creamy brown with a reddish brown medial streak ( Fig. 21 ); (14) average SVL in adult females: 35.3 +/- 3.5 mm (29.7-33.9 mm; n = 15); in adult males: 24.6 +/- 1.7 mm (21.4-27.0 mm; n = 14). Figure 21. Color variation in live individuals of Pristimantis lutzae sp. nov. A QCAZ 53728 (juvenile female, SVL 26.5 mm) B QCAZ 56183 (male, SVL 23.3 mm) C QCAZ 56184 (male, SVL 24.8 mm) D QCAZ 56185 (male, SVL 26.1 mm). Dorsolateral view on the left, ventral view on the right. Comparison with other species. Pristimantis lutzae is similar to P. balionotus , P. chomskyi , P. gloria , P. multicolor sp. nov., and P. philipi . The most similar is P. balionotus , which occurs at lower elevations and can be recognized by the absence of basal webbing between toes (present in P. lutzae ) and having smaller discs on fingers. Pristimantis lutzae can be distinguished from P. chomskyi by the golden to creamy brown iris with a reddish dark brown streak (orange with a faint reddish brown streak in P. chomskyi ), and having a bigger tympanum (males Z = 2.45677, p = 0.0140, TD/SVL = 4.5-4.7% in P. chomskyi , 5-5.4% in P. lutzae ). Pristimantis gloria differs from P. lutzae in having a wartier skin, wide black reticulations on iris (thin in P. lutzae ), and a larger ratio between the length and width of the head (males Z = -5.00826, p <0.0001, HL/HW = 96.8-114.5% in P. gloria , 90-97% in P. lutzae ; females Z = -3.77517, p = 0.0002, HL/HW = 92-105% in P. gloria , 90-96% in P. lutzae ). Pristimantis multicolor sp. nov. has a longer head (males Z = 3.67756, p = 0.0002, HL/SVL = 33.4-37% in P. lutzae , 34.1-40.4% in P. multicolor sp. nov.; females Z = 3.9524, p <0.0001, HL/SVL = 35-37.5% in P. lutzae , 36.5-40.6% in P. multicolor sp. nov.), larger tympanum (males Z = 3.57469, p = 0.0004, TD/SVL = 5-5.4% in P. lutzae , 4.9-6% in P. multicolor sp. nov.; females Z = 3.9524, p <0.0001, TD/SVL = 4.9-5.6% in P. lutzae , 5.5-6.7% in P. multicolor sp. nov.) and larger eyes (males Z = 2.75174, p = 0.0059, ED/SVL = 10.3-12.1% in P. lutzae , 10.3-13.2% in P. multicolor sp. nov.; females Z = 3.3083, p = 0.0009, ED/SVL = 9.9-12.1% in P. lutzae , 10.7-12.4% in P. multicolor sp. nov.), relative to the body length, than P. lutzae . Pristimantis lutzae is readily distinguished from P. philipi because it has a visible tympanic membrane and annulus, and its males have vocal slits (absent traits in P. philipi ). Description of the holotype. An adult female (QCAZ 37546, SC 21029). Measurements (in mm): SVL 31.7; TL 13.7; FL 14.7; HL 11.2; HW 12.1; ED 3.7; TD 1.7; IOD 3.8; EW 2.8; IND 2.3; EN 3.5; TED 1.3. Head wider than long, slightly narrower than body; snout moderately long, rounded with a small papilla in dorsal and lateral view; cranial crests absent; nostrils slightly protuberant, narrow, directed laterally with slight dorsal inclination; canthus rostralis slightly concave in dorsal view, rounded in cross section; loreal region slightly concave; upper eyelid bearing a small, low and rounded tubercle surrounded by inconspicuous tubercles; tympanic annulus prominent, its upper and posterior margins concealed by supratympanic fold; tympanic membrane distinct; two low and rounded postrictal tubercles surrounded by lower tubercles. Choanae large, round, not concealed by palatal shelf of maxillae; dentigerous processes of vomers prominent, oblique, moderately separated, positioned posteromedial to choanae; each vomer bearing several teeth; tongue longer than wide, posterior border notched, posterior half free. Dorsum shagreen with scattered low tubercles, larger posteriorly; dorsolateral folds absent; skin on flanks bearing low rounded tubercles, larger than those on dorsum; skin on chest and belly coarsely areolate, that on throat shagreen, ventral surfaces of limbs smooth, ventral surfaces of thighs coarsely areolate; discoidal fold present. Ulnar tubercles rounded and low; outer palmar tubercle bifid, three times the size of ovoid thenar tubercle; subarticular tubercles prominent, rounded; low supernumerary tubercles; fingers bearing broad lateral fringes; Finger I shorter than Finger II; discs on fingers expanded and truncate; pads on fingers surrounded by circumferential grooves on all fingers ( Fig. 8A ). Dorsal surfaces of hindlimbs shagreen with scattered small tubercles; posterior surfaces of thighs smooth, ventral surfaces of thighs coarsely areolate; heel bearing a median, low, rounded tubercle surrounded by smaller ones; outer and inner edge of tarsus bearing a row of small tubercles; small inner tarsal fold present; inner metatarsal tubercle ovoid, elevated, three times the size of round outer metatarsal tubercle; plantar surface with small, indistinct supernumerary tubercles; subarticular tubercles prominent, rounded; toes bearing broad lateral fringes; basal webbing between Toes IV and V present; discs on toes smaller than those on fingers, slightly expanded, truncate; all toes having pads surrounded by circumferential grooves; relative lengths of toes: I <II <III <V <IV; Toe V much longer than Toe III (disc on Toe III reaches the distal edge of penultimate subarticular tubercle on Toe IV, disc on Toe V reaches the distal edge of distal subarticular tubercle on Toe IV; Fig. 8A ). Coloration of the holotype in preservative is shown in Figure 20A ; coloration in life, unknown. Coloration of holotype in preservative . Dorsum grayish dark brown with lighter irregular reticulations and scattered black spots; dark brown supratympanic stripe, canthal, and interorbital bands; dorsal surfaces of limbs with the same background as dorsum and darker irregular transversal bands and scattered dark brown spots; groins, anterior, and posterior surfaces of thighs reddish brown with cream small spots; ventral surfaces of body cream; soles and palms dusty cream ( Fig. 20A ). Coloration of holotype in life . Unknown. Variation. Variation in preservative is based on 40 individuals of the type series and photographs from eight individuals. Variation in life and preservative is shown in Figures 21 , 22 . Coloration in life is provided in parenthesis. Dorsal coloration varies from light to dark gray or brown (light, orangey or dark brown); markings on dorsum (light brown, yellow or orange) are present or absent, most individuals have irregular reticulations, some have a series of parallel longitudinal stripes; dorsum and flanks may bear scattered black or white spots. All individuals bear supratympanic and canthal stripes (black, brown); an interorbital stripe or band is present except for individuals with the longitudinal pattern on dorsum. Flanks have the same background color as dorsum. Groins, posterior and anterior surfaces of thighs are cream to brown with small paler spots (pinkish, purplish or reddish brown, suffused or not with orange, with cream or light brown spots). Limbs bear dark transversal bands or scattered small dark spots. Ventral coloration varies from cream to dusty cream (white to cream); venter with or without brown reticulations. The iris is golden to creamy brown with a reddish dark brown medial horizontal streak. White to light-blue sclera. Figure 22. Color variation in preserved individuals of Pristimantis lutzae sp. nov. A Dorsal view of (from left to right): QCAZ 37545 (female), QCAZ 37547 (female), QCAZ 27534 (female), QCAZ 27597 (female) B Dorsal view of: QCAZ 32785 (female), QCAZ 56185 (male), QCAZ 53728 (juvenile female), QCAZ 27471 (male) C Ventral view of specimens in ( A ) D Ventral view of specimens in B. See Suppl. material 2 for locality data. All specimens are shown at the same scale. Distribution, natural history, and conservation status. Pristimantis lutzae is known from Paramo, Inter-Andean Shrub, Western and Eastern Montane Forest in the Andes of Azuay and Canar Provinces in Ecuador, between 2895-4100 m a.s.l ( Fig. 1 ). Individuals collected at night were found in bunch grasses, pastures, and low vegetation up to 80 cm above the ground. Individuals collected at day were found in pastures or underneath rocks. Calling males have been found on bunch grasses or low vegetation during February, July, and December at night. Despite the relatively small distribution range of this species (Extent of Occurrence = 2338 km2) we assign it to the Least Concern Red List category because its distribution overlaps with four protected areas, Cajas National Park, Mazan Reserve, Mazar Wildlife Reserve, and Yanuncay Irquis Protected Forest, and it is a common species in these places. Etymology. The specific epithet is a noun in the genitive case and is a patronym for Bertha Lutz, who was a Brazilian herpetologist. We name this species after her in recognition of her scientific career and her activism in the fight for gender equality. Remarks. Specimens of this species were previously referred as Pristimantis riveti (Despax 2011) based on Lynch (1979) characterization of the species (e.g., Almendariz and Orces 2006; Heinicke et al. 2007 ; Padial et al. 2014 ). Photographs of the holotype of P. riveti ( Fig. 23B ) as well as the location of its type locality indicate that P. lutzae is a different species. See Taxonomic status of P. riveti section for details. Through morphological and molecular evidence, we recognize P. lutzae as a different species and assign it to the P. phoxocephalus species group. Figure 23. Preserved holotypes of Pristimantis phoxocephalus and P. riveti . Dorsal, ventral and lateral views of: A Pristimantis phoxocephalus (KU 142075) B Pristimantis riveti (MNHNP 1902.357). Shown at the same scale.