identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
0B85AE9687215775A880D63DF730612D.text	0B85AE9687215775A880D63DF730612D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Niceforonia tempestuosa Ospina-Sarria & Rodríguez-Betancourt & Gómez-Sepúlveda & Morales-Vertel & Vélez-Franco & Gallego-Posada & Herrera-Rengifo & Grajales Escobar & Ceballos-Castro & García-Gómez & Velásquez-Trujillo & Ortiz Baez & Mendez Narvaez 2026	<div><p>Niceforonia tempestuosa sp. nov.</p><p>Figs 4 A – D, 5 A – C, 6 A, 6 C</p><p>Proposed standard Spanish name.</p><p>Rana de los bosques montanos nublados.</p><p>Proposed standard English name.</p><p>Montane cloud forest frog.</p><p>Type material.</p><p>Holotype. • ARUQ -1885 (field no. JJS 702), an adult male from an interior forest at a site named “ <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-76.28333&amp;materialsCitation.latitude=4.75" title="Search Plazi for locations around (long -76.28333/lat 4.75)">Corazón</a> ” within the Reserva Natural Comunitaria Cerro El Inglés (4°45'N, 76°17'W, 2261 m), Municipality of El Cairo, Valle del Cauca, Colombia; collected by Jhon Jairo Ospina-Sarria, Andrés Felipe Rodríguez-Betancourt, Ximena Gómez-Sepúlveda, Iván Darío Morales-Vertel, Valentina Vélez-Franco, Irene Ceballos-Castro, and Isabella García-Gómez on 25 October 2024 . Paratypes. A total of 11 specimens. • ARUQ -1882 (adult male) and ARUQ -1883 (juvenile male), collected on 24 October 2024; • ARUQ -1884 (adult male), collected with the holotype on 25 October 2024; • ARUQ -1889, 1891, 1893 (adult males), ARUQ -1892 (subadult male), ARUQ -1890 (subadult female), and ARUQ -1886 (juvenile male), collected on 25 April 2025; all from Corazón, Reserva Natural Comunitaria Cerro El Inglés . • ARUQ -1887 (adult female) and ARUQ -1888 (subadult female), collected on 20 April 2025 from Bosque Puente Quebrada Las Amarillas, Reserva Natural Comunitaria Cerro El Inglés .</p><p>Diagnosis.</p><p>Niceforonia tempestuosa is diagnosed by the following combination of characters: (1) skin of dorsum smooth with scattered small warts; dorsolateral and \ / - shaped scapular folds present; venter smooth, discoidal fold absent; (2) tympanic membrane differentiated, upright-oval to round; its length 33.3 – 44.8 % of eye length in six adult males and 47 % in one adult female; prominent tympanic annulus, its upper edge covered by supratympanic fold, extending only just posterior to tympanic annulus; (3) snout subacuminate in dorsal view, rounded in profile; a fleshy swelling of lip in adult males; canthus rostralis straight; (4) upper eyelid bearing minute tubercles, narrower than IOD (62.0–85.1 % IOD); interocular tubercle absent; cranial crest absent; (5) choanae small, ovoid; partially concealed by palatal shelf of maxillary arch; dentigerous processes of vomers prominent, triangular in outline, separated medially for a distance equal to half the width of the visible dentigerous process, positioned posterior to level of choanae, each dentigerous process of vomers bearing five to seven teeth; (6) males lacking vocal slits, small non-spinous nuptial pads on median surface of thumb above joint with metacarpal; (7) first finger equal or slightly longer than second; circumferential grooves present on all fingers; discs slightly wider than digits, disc on finger I equal to disc on finger II and these in turn smaller than discs on fingers III and IV; morphology of the ungual flap on fingers pointed; (8) fingers with lateral fringes, weakly defined on outer side of finger 4; palmar tubercle bifid (inner lobe largest); thenar tubercle oval, slightly smaller than inner lobe of palmar tubercle; supernumerary tubercles prominent, two on thumb and third finger and one on second and fourth; subarticular tubercles present; subarticular tubercles projecting, with elongate base and larger than supernumerary tubercles; (9) three low ulnar tubercles not coalesced; antebrachial tubercle present; (10) small tubercles on heel and outer edge of tarsus; inner tarsal fold absent; (11) elongate inner metatarsal tubercle, its length twice its width; protuberant, conical outer metatarsal tubercle one-half size of inner metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles present; subarticular tubercles projecting, with elongate base; (12) toes bearing prominent lateral fringes, in finger III and IV only visible in the distal part of digits; toe webbing absent; toe III much longer than toe V; toe III extending to penultimate subarticular tubercle of toe IV; toe V reaching two-thirds between basal and penultimate subarticular tubercles of toe IV; discs and circumferential grooves present on all toes; discs of toes II – IV larger than discs on fingers I and V; (13) color in life: dorsum reddish-brown to grayish, with or without two dark brown marks originating at the snout, extending through the loreal region and upper eyelids, and continuing along the \ / - shaped scapular folds to converge at the sacral region. Supratympanic fold with a dark mark extending to the upper edge of the cream-colored postrictal ridge. A broad dark mark, bordered by a cream coloration, extends along the outer edge of the dorsolateral folds; it becomes wider at the posterior ends, where the supra-inguinal spots are located. Black anal patch present. Diffuse brown labial bars and marked transverse brown bars on the arms and legs. The groin, and anterior and posterior surfaces of the thighs are reddish-orange, the anterior part of flanks is yellow, and the ventral surfaces of thighs, belly, and arms are yellow. The throat is brown with silvery-gray flecks, featuring a superficial yellow coloration restricted to its medial portion that does not extend to the margins. However, the intensity of this yellow and orange coloration may vary among individuals; specifically, it can be much less intense (paler) in juveniles, subadults, and some females. Ventral surfaces of hands and feet, dark. Iris bright copper with black flecks; (14) males smaller than females; SVL in adult males 21.5–25.1 mm (mean ± 1 SD = 23.7 ± 1.3 mm; n = 6) and 31.1 mm in one female.</p><p>Comparisons with congeners.</p><p>Niceforonia tempestuosa is distinctive in the genus by having dorsolateral folds, tympanic membrane, fleshy swelling of lip in adult males, nuptial pads, circumferential grooves on all fingers, black facemask on loreal region, dorsum reddish-brown to grayish with two dark brown marks, ventral surfaces of thighs, belly, and arms, yellow, black anal patch, and dark supra-inguinal spots. Niceforonia tempestuosa most closely resembles N. mantipus and N. nigrovittata by having dorsolateral folds and \ / - shaped scapular folds, tympanic membrane, snout subacuminate in dorsal view, fleshy swelling of lip in adult males, dark supra-inguinal spots, venter smooth and by lacking discoidal fold and vocal slits. Nevertheless, Niceforonia tempestuosa is distinguished by having a yellow belly and ventral surfaces of the arms and thighs (light gray background with silvery flecks in N. mantipus), and a reddish-orange coloration on the groin and the anterior and posterior surfaces of the thighs (dark brown with silvery and coppery-red flecks in N. mantipus). Furthermore, males of the new species exhibit a brown throat with silvery-gray flecks and a superficial yellow coloration restricted to its medial portion that does not extend to the margins, whereas males of N. mantipus have a light gray throat with silvery flecks and two well-defined, large dark blotches on the lateral base of the throat (Figs 4, 5). Additionally, Niceforonia tempestuosa is distinguished by having nuptial pads (absent in N. nigrovittata) and a reddish-orange posterior surface of the thighs (brown with cream flecks in N. nigrovittata) (Lynch and Duellman 1980).</p><p>Niceforonia tempestuosa can be distinguished from the remaining species of the genus by having circumferential grooves on all fingers (absent in fingers I and II in N. adenobrachia Ardila-Robayo, Ruiz-Carranza &amp; Barrera-Rodriguez, 1996, N. brunnea, N. latens Lynch, 1989, N. nana, and N. peraccai). Furthermore, Niceforonia tempestuosa is distinguished from N. adenobrachia, N. brunnea, and N. peraccai by having a black anal patch, and from N. latens and N. nana by the presence of a tympanic membrane. The development of dorsolateral folds distinguishes N. tempestuosa from N. aderca (Lynch, 2003), N. elassodiscus, N. fallaciosa (Duellman, 2000), and N. philippi . Additionally, N. tempestuosa lacks vocal slits (present in N. aderca, N. dracula, N. philippi, and N. quilloturo) and discoidal fold (present in N. dracula, N. elassodiscus, N. fallaciosa, N. quilloturo) (Reyes-Puig et al. 2026). The presence of dark supra-inguinal spots distinguishes N. tempestuosa from N. babax (Lynch, 1989) and N. lucida (Cannatella, 1984); these species further differ from N. tempestuosa by having vocal slits. Niceforonia tempestuosa differs from N. columbiana (Werner, 1899) by lacking basal webbing (Lynch 1975), and from N. araiodactyla (Duellman &amp; Pramuk, 1999) and N. elassodiscus by lacking a discoidal fold (Duellman and Pramuk 1999). Furthermore, the presence of scattered small warts on a smooth dorsum distinguishes N. tempestuosa from N. araiodactyla, which has uniformly smooth dorsal skin (Duellman and Pramuk 1999). Additionally, the reddish-orange coloration on the posterior surface of the thighs distinguishes N. tempestuosa from N. elassodiscus (which has a uniform gray to brown coloration on the posterior surface). Finally, Niceforonia tempestuosa is genetically highly divergent from all other species in the genus, with uncorrected p - distances in 16 S gene ranging from 7.44–18.33 % (Table 2).</p><p>Descriptions of the holotype.</p><p>An adult male with head slightly wider than body; HW 40.8 % of SVL; HL 35.1 % of SVL; snout long, subacuminate in dorsal view, rounded in profile; eye-nostril distance 82.7 % of diameter of eye; nostrils protuberant, directed slightly back. Canthus rostralis straight in dorsal view, not elevated; loreal region depressed; top of head flat; upper eyelid bearing minute tubercles, its width 62.1 % of IOD; supratympanic fold extending only just posterior to tympanic annulus; tympanic membrane present, upper edge of tympanic annulus covered by supratympanic fold; postrictal tubercles fused into ridge. Choanae small, ovoid; partially concealed by palatal shelf of maxillary arch; dentigerous processes of vomers prominent, each process bearing seven teeth; tongue longer than wide, its posterior border not notched, posterior fourth not adherent to floor of mouth; vocal slits absent.</p><p>Skin of dorsum smooth with scattered small warts; dorsolateral and \ / - shaped scapular folds present; skin on belly, and throat smooth (Figs 4 A – D, 5 A – C); discoidal fold absent, no tubercles in the cloacal region. Three low ulnar tubercles on the ventrolateral edge of the forearm, not coalesced; palmar tubercle bifid (inner lobe largest); thenar tubercle oval, slightly smaller than inner lobe of palmar tubercle; subarticular tubercles projecting, with elongate base, one on thumb and second finger and two on third and fourth fingers, and larger than supernumerary tubercles; supernumerary tubercles prominent, two on thumb and third finger and one on second and fourth. Small nuptial pads on median surface of thumb above joint with metacarpal.</p><p>Fingers having lateral fringes, weakly defined on outer side of finger 4; relative lengths of fingers I&gt; II &lt;IV &lt;III, all fingers having terminal ventral pads well defined by circumferential grooves; disc on thumb equal to disc on finger II; discs on fingers III and IV smaller than tympanic membrane; morphology of the ungual flap on fingers pointed. Short robust hind limbs; when hind limbs flexed against axis of body, heels touching or barely overlapping; tibia length 54.7 % of SVL; foot length 55.9 % of SVL; heel and outer edge of tarsus having small tubercles; inner tarsal fold absent; inner metatarsal tubercle elongate, its length twice its width; conical outer metatarsal tubercle one-half size of inner metatarsal tubercle; toes bearing lateral fringes and discs on expanded pads; webbing absent; relative lengths of toes I &lt;II &lt;V &lt;III &lt;IV, toe III much longer than toe V, toe III extending to penultimate subarticular tubercle of toe IV, toe V reaching two thirds between basal and penultimate subarticular tubercles of toe IV (Figs 5 B, 6 C), all toes having terminal ventral pads well defined by circumferential grooves; discs of toes II – IV larger than discs on fingers I and V; subarticular tubercles projecting, with elongate base, one on toes I and II, two on toes III and V, and three on toe IV; supernumerary plantar tubercles absent.</p><p>In life, dorsum brown with two dark brown marks originating at the snout and extending through the loreal region and upper eyelids, where they become diffuse, and continuing along the \ / - shaped scapular folds (without converging). A weak dark mark, bordered by a cream coloration, extends along the outer edge of the dorsolateral folds; it becomes wider at the posterior ends, where the supra-inguinal spots are located. Postrictal ridge cream-colored; labial bars diffuse and brown; arms and legs with marked transverse brown bars. Black anal patch bordered by a cream coloration. Groin, and anterior and posterior surfaces of thighs, reddish-orange; anterior part of flanks, yellow. Ventral surfaces of thighs, belly, and arms, yellow with uniformly distributed dark brown marks (more abundant on the flanks and anterior part of the belly). The throat is brown with silvery-gray flecks, featuring a superficial yellow coloration restricted to its medial portion that does not extend to the margins (Fig. 4 C, D). Iris bright copper with black flecks (Fig. 4 A, B).</p><p>Measurements of holotype (mm).</p><p>SVL 24.5, tibia length 13.4, foot length 13.7, HL 8.7, HW 9.9, IOD 2.9, tympanum diameter 1.3, internarial distance 3.0, width of upper eyelid 1.8, diameter of eye 2.9, eye-nostril distance 2.4.</p><p>Variation.</p><p>Sexual dimorphism in size, with adult females being larger than adult males (Table 3). Dorsal coloration also varies, ranging from reddish-brown to grayish-brown. The same applies to the longitudinal dark markings extending from the snout to the sacral region, which are present in some specimens. We also noted variation in the yellow and orange coloration of ventral surfaces can be much less intense (paler) in juveniles, subadults, and some females. In contrast, the dark supra-inguinal spots are consistently present in all specimens. The fleshy swelling of the lip is only evident in adult males.</p><p>Distribution and ecology.</p><p>The species is known only from two localities in humid montane forests of the central part of the Cordillera Occidental of Colombia, within the Key Biodiversity Area (KBA) Serranía de los Paraguas: Reserva Natural Comunitaria Cerro El Inglés (El Cairo, Valle del Cauca) and El Mirador (Balsal, Versalles, Valle del Cauca, 4°40'N, 76°17'W), between 2169–2294 m (Fig. 1). The species inhabits the leaf litter of well-preserved forest interiors, where it can be found by disturbing the substrate during the day or actively foraging at night. The species exhibits a preference for sloping over level microhabitats. Individuals have also been observed within crevices formed by the buttress roots of large trees and among the stilt roots of Wettinia sp. palms (Fig. 7). As a leaf-litter dweller, this species is syntopic with Strabomantis cerastes, another inhabitant of the forest floor in the Serranía de los Paraguas.</p><p>Etymology.</p><p>The specific epithet is the feminine adjective tempestuosa (meaning “ stormy ”), derived from the Latin noun tempestas (meaning “ storm ” or “ weather ”). This name was chosen for local communities as a metaphor for the global climate emergency, referring to the increased frequency and intensity of extreme weather events — symbolized by the “ storm ” — which represent a growing threat to global biodiversity. The discovery of this new species in the Key Biodiversity Area (KBA) Serranía de los Paraguas underscores the resilience of this biodiversity hotspot amidst the current climate crisis, safeguarded by the persistent efforts of local communities and biodiversity guardians.</p><p>Remarks.</p><p>Lynch and Duellman (1980), Lynch (1989), and Lynch (2003) reported the occurrence of a fleshy swelling of the upper lip in adult males of N. nigrovittata, N. mantipus, and N. aderca, respectively, followed more recently by Reyes-Puig et al. (2026) for N. quilloturo . We herein report this character in males of Niceforonia latens and Niceforonia tempestuosa . According to Lynch (2003), the presence of this character exclusively in males suggests an involvement in either courtship or the excavation of egg burrows. Among brachycephaloid frogs, a character resembling the fleshy swelling of the upper lip found in Niceforonia occurs in species of the genus Eleutherodactylus, subgenus Pelorius (Hedges et al. 2008 a) . As noted by Hedges et al. (2008 a), species possessing shovel-shaped snouts are specialized in constructing enclosed underground chambers for vocalization and, presumably, oviposition. To date, the courtship and nesting behaviors of Niceforonia species remain unknown; therefore, the functional implications of this labial swelling warrant further investigation.</p></div>	https://treatment.plazi.org/id/0B85AE9687215775A880D63DF730612D	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	Ospina-Sarria, Jhon Jairo;Rodríguez-Betancourt, Andrés Felipe;Gómez-Sepúlveda, Ximena;Morales-Vertel, Iván Darío;Vélez-Franco, Valentina;Gallego-Posada, Nayely;Herrera-Rengifo, Jose Daniel;Grajales Escobar, John Edison;Ceballos-Castro, Irene;García-Gómez, Isabella;Velásquez-Trujillo, David Andrés;Ortiz Baez, Ayda Susana;Mendez Narvaez, Javier	Ospina-Sarria, Jhon Jairo, Rodríguez-Betancourt, Andrés Felipe, Gómez-Sepúlveda, Ximena, Morales-Vertel, Iván Darío, Vélez-Franco, Valentina, Gallego-Posada, Nayely, Herrera-Rengifo, Jose Daniel, Grajales Escobar, John Edison, Ceballos-Castro, Irene, García-Gómez, Isabella, Velásquez-Trujillo, David Andrés, Ortiz Baez, Ayda Susana, Mendez Narvaez, Javier (2026): A new species of Niceforonia (Amphibia, Anura, Strabomantidae) from the Serranía de los Paraguas, Colombia, with the description of a new phenotypic synapomorphy for the Niceforonia nigrovittata species group. ZooKeys 1289: 159-184, DOI: 10.3897/zookeys.1289.198411
