Acta Herpetologica 13(1): 51-64, 2018 ISSN 1827-9635 (print) © Firenze University Press ISSN 1827-9643 (online) www.fupress.com/ah DOI: 10.13128/Acta_Herpetol-20693 Natural history and conservation of the Nurse Frog of the Serranía del Perijá Allobates ignotus (Dendrobatoidea: Aromobatidae) in northeastern Colombia Hernán Darío Granda-Rodríguez1,2, Andrés Camilo Montes-Correa3,*, Juan David Jiménez-Bolaño3, Mar- vin Anganoy-Criollo4,5 1 Programa de postgrado en Áreas Silvestres y Conservación de la Naturaleza, Facultad de Ciencias Forestales y de la Conservación de la Naturaleza, Universidad de Chile, Santiago, Chile 2 Programa de Biología, Universidad del Magdalena, Santa Marta, Colombia 3 Grupo de Investigación en Manejo y Conservación de Fauna, Flora y Ecosistemas Estratégicos Neotropicales (MIKU), Universidad del Magdalena, Santa Marta, Colombia. *Corresponding author. E-mail: andresc.montes@gmail.com 4 Grupo de Cladística Profunda y Biogeografía Histórica, Laboratorio de Anfibios, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá D.C., Colombia 5 Laboratorio de Anfibios, Instituto de Biociencias, Universidad de Sao Paulo, Brasil Submitted on: 2017, 25th May; revised on: 2017, 6th September; accepted on: 2018, 1st March Editor: Emilio Sperone Abstract. We describe new findings on the geographic distribution, habitat uses, relative abundance, tadpoles, and advertisement call of Allobates ignotus. Data of habitat uses and relative abundance were collected during five annual fieldtrips of seven days each one, from April 2010 to June 2014, in a mining zone of Canime Creek Basin, La Jagua de Ibirico, Cesar Department, northeast Colombia. Tadpoles and advertisement call were obtained in La Veguita Creek Basin, Manaure Balcón del Cesar, Cesar. We report six new localities with presence of A. ignotus, all between 7-96 km in straight line to the type locality, from 194-1236 m a.s.l. This species is recorded for the first time in the Tropi- cal Dry Forest Life Zone and in La Guajira Department. Four substrate categories used by A. ignotus were detected (leaf-litter, rocks, naked floor, and lower branches), and differential use was highlighted. The larval morphology of A. ignotus was described, based on 11 specimens between 25-29 stages. Like other Allobates tadpoles, the gap of the second teeth row is “wide”. The advertisement call of A. ignotus is composed by a series of high frequency ascending pulsed notes. Possibly, the high frequency of the call is due to the acoustic disturbance generated by the streamflow noise of the creeks. In 2014, a decrease of relative abundance of A. ignotus in Canime Creek was detected. This, cou- pled to restricted distribution and loss of habitat quality are sufficient criteria to suggest the category of vulnerable for A. ignotus. Keywords. Advertisement call, Colombian Andes, Colombian Caribbean, geographic distribution, habitat uses, rela- tive abundance, tadpoles, tropical dry forest. INTRODUCTION The Serranía del Perijá is a mountain system of the northern South America, whose highest points make up the northern boundary between Colombia and Ven- ezuela. Because its geographic position, the Serranía del Perijá functioned in the past as a biological corridor between Andes and Caribe, showing biogeographical affinities with the Sierra Nevada de Santa Marta (= Santa Marta Mountains) and the Cordillera Oriental (= Colom- 52 Hernán Darío Granda-Rodríguez et alii bian Eastern Andes) (Hernández-Camacho et al., 1992). Currently, the Serranía del Perijá is considered of great importance from an amphibian biodiversity approach by the high endemicity, deserving to constitute a distinct ecogeographic unit (Romero-Martínez and Lynch, 2012). In Colombia, 14 species of the genus Allobates Zim- merman and Zimmerman, 1988 (Aromobatidae) are rec- ognized, broadly distributed in the wet and dry forests of the western and eastern lowlands of the country (Acosta- Galvis, 2017). However, there are gaps in the knowledge of the Colombian Allobates about of their taxonomy and natural history (Anganoy-Criollo, 2012). For the species of this country, there are some notes on the abundance, habitat use, conservation and feeding ecology of Allobates juanii (Cáceres-Andrade and Urbina-Cardona, 2009; Ast- wood-Romero et al., 2016), and a bit more information for Allobates femoralis and Allobates talamancae (v. gr. Junca and Eterovick, 2007; Ringler et al., 2009; Hopkins and Lahanas, 2011; Montanarin et al., 2011; Keafer et al., 2012; Lechelt et al., 2014), even if these data do not come from Colombian populations. For all other Colombian Allobates, the information on their geographic distribu- tion, habitat use, and calls, is mostly provided only by the original (re-)descriptions. Allobates ignotus Anganoy-Criollo, 2012 (Fig. 1) is an aromobatid frog, only known from its original descrip- tion, endangered, and endemic to the Serranía del Peri- já (IUCN SSC Amphibian Specialist Group, 2017). In Fig. 1. Specimens of Allobates ignotus from Canime Creek (A), La Frontera’s way (B), La Veguita Creek (C), Cesar department, Colombia; and Quebrada Nicaragua (D), La Guajira department, Colombia. Photographs by H.D. Granda-Rodríguez (A), A. del Portillo-Mozo (B), and J.D. Jiménez-Bolaño (C, D). 53Natural history and conservation of Allobates ignotus the distribution of this species, only three localities are known (Anganoy-Criollo, 2012; Acosta-Galvis, 2017) with very few available information. As a consequence, being many aspects of the biology, ecology, and natural history of A. ignotus unknown, it is hard to determine the conservation status, threats, and measures to species protection. In this paper, we report new data on several aspects of the natural history (geographic distribution, free-swimming tadpole, advertisement call, habitat use, and abundance) and conservation status of A. ignotus. MATERIALS AND METHODS Study area All records of Allobates ignotus were collected on the foot- hills and midlands on the western flank of the Serranía del Perijá. These localities have a bimodal rainfall regime, with high to moderate water deficit in dry periods (Rangel-Ch. and Carvajal-Cogollo, 2012). The first rainy peaks is from April to May and second from September to October. The two periods of low rains are from December to March and in July (Rangel- Ch. and Carvajal-Cogollo, 2009). The foothills of the Serranía correspond to the “Zonobioma Tropical Alternohígrico” (follow to Hernández-Camacho and Sánchez 1992) or to the Tropical Dry Forest Life Zone (sensu Holdridge, 1967). The midlands of the Serrania is a biome known as “Orobioma de la Selva Sub- andina” (Hernández-Camacho and Sánchez, 1992) or as Moist Forest, Wet Forest and Rain Forest life zones (Holdridge, 1967). The latter has a low water deficit in dry periods (Rangel-Ch. and Carvajal-Cogollo, 2009). Habitat use and abundance We obtained the relative abundance, habitat and micro- habitat uses data in three sections of the Canime Creek (Sector 1: 09°33’16.10’’N, 73°16’00.90’’W, 250 m a.s.l.; Sec- tor 2: 09°36’10.20’’N, 73°16’53.70’’W, 200 m a.s.l.; Sector 3: 09°34’22.42”N, 73°15’52.24”W, 194 m a.s.l.), a quaternary tribu- tary of the Cesar River, in the municipality of La Jagua de Ibir- ico, Cesar Department, northeast Colombia. Surveys were con- ducted in a mining area, where most of the land corresponds to coal exploitation zones and deposits of waste material. Grass- lands, wooded savannas, and cattle pastures occupy another larger percentage of the land use. The tropical dry forest cover is established in a smaller proportion, only in the margins of streams, in mosaics of gallery forest, bushes, stubble, and decid- uous forest. We performed five field trips to the Canime Creek, one in dry season (February 2011) and four in rainy season (April 2010, August 2012, April 2013, and June 2014), for a total of seven days for each trip. For frogs sighting, we used the Visual Encounter Survey (VES) (Crump and Scott, 1994). Daily, two observers conducted random walks between 09:00-12:00 h and 15:00-16:00 h, for a total sampling effort of 109 h × observer. We recorded the substrate just under the specimen at the time of sighting. We collected some voucher specimens, sacrificed on immersion to chlorobutanol solution (10%). All collected vouchers were deposited in the Amphibian Collection of the Centro de Colecciones Biológicas de la Universidad del Mag- dalena, at Santa Marta, Colombia (CBUMAG:ANF). To con- firm the species level determination, the collected voucher was directly compared to the type series of A. ignotus deposited in the Amphibian Collection of the Instituto de Ciencias Naturales at Universidad Nacional de Colombia (ICN 55434, holotype; 55427-433, 55435-439, paratypes) Relative abundance was calculated as the number of indi- viduals in each sample relative to capture effort, expressed in individuals/hours × observer (RA = Ind/h × obs) (Lips, 1999). We performed a homogeneity Chi square test (χ2) in order to reject the null hypothesis that there is no differential use of the microhabitat by A. ignotus. The expected values were calculated equitably from each substrate category detected in field, assum- ing that all substrates had the same probability to be occupied. Free-swimming tadpoles From December 2015, we collected tadpoles in La Veguita Creek (10°23’12.57”N, 73°03’19.03”W, 633 m a.s.l.), La Vegui- ta Village, Municipality of Manaure Balcón del Cesar, Cesar department, Colombia. This creek is surrounded for a second- ary tropical dry forest relict, on a slope of approximately 20°, with a more or less staggered configuration of small waterfalls and pools, and abundant boulders. La Veguita Creek is the repository of waste waters of the municipality, showing grayish color. We found the tadpoles on the marginal puddles of the stream. These puddles had a low depth (< 20 cm) and recorded water temperature was 28 °C. The puddle bottom was very sedi- ment, but column was transparent. Tadpoles were sacrificed in an immersion in the formaldehyde solution (10%). We assigned the tadpoles to Allobates ignotus because the metamorphic indi- viduals observed in the same site have adult features (dorsolat- eral, oblique, and ventrolateral stripes, and color pattern) of A. ignotus. In addition, the only one dendrobatoid frog detected in the study area was this species. Description was based on a set of 11 specimens (CBUMAG:ANF 936), collected by J.D. Jiménez-Bolaño, J.A. Rincón, and A.C. Montes-Correa. No other species of Allobates (or any other dendrobatoid frog) were found in La Veguita Creek. Here, we provided a description of the free-swimming tadpoles at stages 25-29 (according to Gosner, 1960). No other developmental phases were available (i.e., neither back-riding tadpoles nor metamorphic tadpoles). Measurements (in mm) and proportions are provided for all tadpoles. Larval terminol- ogy were determined according to Altig and McDiarmid (1999). To describe the attached type of the vent tube to the ventral fin, we follow the Figure 3.5 of Altig and McDiarmid (1999). Description and measurement of the number of marginal papil- lae on each lip followed Anganoy-Criollo (2013) and disposi- tion of the row of marginal papillae followed Sánchez (2013). Values of the gap of the second tooth row of other species of Allobates were obtained from the measurements or illustra- 54 Hernán Darío Granda-Rodríguez et alii tions of the oral disc of free-swimming tadpoles provided in each description. Measurements were taken with a manual cali- per (0.01 mm) and with an ocular micrometer of a Carl Zeiss Stemi 2000 stereomicroscope (0.001 mm). For morphologi- cal characters the following abbreviations were used: (AL gap) anterior lip gap, (A-2 gap) medial gap in second anterior tooth row, (BH) body height, (BL) body length, (BW) body width, (END) eye-nostril distance, (IND) internostril distance, (IOD) interorbital distance, (LJW) lower jaw sheath width, (ODW) oral disc width, (P-AL) papillae on the right side of anterior lip, (P-PL) papillae on posterior lip in a space of 1 mm, (SH) spira- cle height at its base, (SL) spiracle length, (SND) snout-nostril distance, (SOH) spiracle opening height, (SSD) snout-spiracle distance, (TAL) tail length, (TMW) tail muscle width at its base, (TMH) tail muscle height at its base, (TL) total length, (UJW) upper jaw sheath width, (VT) vent tube length. Advertisement call From December 2015, we collected three recordings of advertisement calls of three males in La Veguita Creek, vil- lage of La Veguita, municipality of Manaure Balcón del Cesar, department of Cesar, Colombia. Digital recordings were made at a sampling rate of 44 kHz and 16 bit resolution with a SONY LCD-PX 240 Voice Recorder with built-in microphone. We collected 281 sec of total recording. One of the males was pre- served as a voucher specimen. Calls were recorded at 16:20 pm, with a temperature of 25.4 °C under shade. Calls were analyzed in PRAAT 6.0.13 for Windows (Boersma and Weenink, 2007). Each sound file of each individual contained four to six calls, which were analyzed individually. The acoustic parameters measured were: call duration (sec), number of notes per call, note duration (sec), internote interval (sec), and rate of notes per sec (notes/sec); as well as the spectral properties, such as dominant frequency and frequency range, with respective mean and standard deviation (SD) (Littlejohn, 2001; Köhler et al., 2017). Graphics was produced with the R package Seewave (Sueur et al., 2008). RESULTS Geographic distribution Based on museum specimens and field observations, we recorded six new localities with confirmed presence of Allobates ignotus (Fig. 2); five localities in Cesar Depart- ment (three sectors in the Canime Creek, 194-250 m a.s.l., La Jagua de Ibirico; La Frontera’s way, 1236 m a.s.l., Agustin Codazzi; La Veguita Creek, 570 m a.s.l., Manaure Balcón del Cesar) and one locality in La Guajira Depart- ment (Nicaragua Creek , 905 m a.s.l., La Jagua del Pilar). The new localities are between 7-96 km in straight line from the type locality, between 194-1236 m a.s.l. The life zone of these localities is tropical dry forest and the tran- sition with the low montane wet forest. The new records widen the occurrence of the species to 605 km2, and rep- resent the first record of A. ignotus in tropical dry forest and La Guajira Department. Habitat uses In the three sections of the Canime Creek, La Jagua de Ibirico lowlands, we recorded 187 individuals of Allo- bates ignotus, all restricted to low flow and shallow creeks, with abundant gallery forest of closed canopy. Allobates ignotus were not recorded in the deciduous forest, bushes, stubble, or pastures adjacent to the riparian forest. In the gallery forest interior, we determined four cat- egories of microhabitat (substrate) occupied by Allobates ignotus: leaf-litter, rocky substrate (boulders-likes), fallen trunks, and lower branches (<50 cm to floor). Allobates ignotus uses differentially the available substrata (χ2 = 51.65, df = 3, P < 0.001, Table 1): 46.64% were record- ed active on the leaf-litter; a slightly lower percentage Fig. 2. Geographic distribution of Allobates ignotus in Colombia. Blue (light grey in the printed version) symbols for known localities and red (dark grey in the printed version) symbols for new records. Development by Lorena Benitez-Cubillos. Table 1. Frequency and homogeneity Chi square test (χ2) of micro- habitat uses by Allobates ignotus in Canime Creek Basin, Cesar department, Colombia. Substrata N χ2 df P Leaf-litter 76 18.301 Rocks 65 7.124 Fallen trunks 30 6.001 Lower branches 16 20.226 Total 187 51.652 3 <0.001 55Natural history and conservation of Allobates ignotus (34.76%) occurred on rocky substrates; fallen trunks and lower branches were used to a lesser extent (16.04% and 8.56%, respectively). Free-swimming tadpoles All measurements and proportions of free living tad- poles are given in Table 2. Tadpoles have an ovoid body in dorsal view, wider at mid-body; depressed in lateral view (Fig. 3A-C), barely higher at level of intestines (BH/ BW = 0.67-0.80; BW/BL = 0.57-0.71), shorter than half of the total length (BL/TL = 0.27-0.39). Snout round- ed in dorsal and lateral view (Fig. 3A, C). The nostrils are rounded or slightly ovoid, positioned dorsally, and opening directed dorsolaterally; located about midway between the eye and the tip of snout; nostril 25-36% of eye diameter; inner margin with a thick fleshy ring and very low fleshy projection dorsally, sometimes no vis- ible. The internarial distance is equivalent to 53-75% of interorbital distance. Interorbital distance 43-55% of body width. Eyes dorsal, directed dorsolaterally. Eye diameter larger than nostril-eye distance (except a tadpole in stage 27, with eye about equal to the nostril-eye distance) and 39-51% of interorbital distance. Spiracle sinistral, slightly longer than high (0.9-1.2 times) with cylindrical shape (i.e., the spiracle height is the same through of spiracle length). Spiracle directed posterodorsally is located below longitudinal midline and on vertical midline of body. Spiracular opening ending 63-77% of body length (from tip of snout), and is barely separated from body. Vent tube short, 8-15% of body length, attached to right side of ventral fin in B-type; opening of vent tube with poste- rior direction and smooth edge. Inner intestines were visible through translucent skin; the longitudinal axis of intestines is sinistral to body; guts in 5-10 coils in ventral view (Fig. 3B). Tail length of 61-73% of total length. At body-tail junction, tail muscle height from 45-66% of body height. Myotomes higher than wide at body-tail junction, except in a tad- Table 2. Measurements (in mm) of the free-swimming tadpoles of Allobates ignotus. The range (mean ± standard deviation) for linear meas- urements are reported; only the range for proportions. For abbreviations, see “Tadpoles” section in Materials and Methods. Measurement Stage 25 (n = 2) Stage 26 (n = 4) Stage 27 (n = 3) Stage 28-29 (n = 2) TL 8.7-16 (12.4 ± 5.2) 9.6-14.9 (12 ± 2.2) 13.9-18.2 (16.2 ± 2.2) 17.8-21 (19.4 ± 2.3) BL 3.2-4.3 (3.8 ± 0.8) 3.6-4.9 (4.3 ± 0.5) 5-6.7 (5.9 ± 0.9) 7-7.6 (7.3 ± 0.4) BW 2-2.9 (2.5 ± 0.6) 2.4-3.3 (2.9 ± 0.4) 3.4-4.2 (3.9 ± 0.4) 4.5-4.6 (4.6 ± 0.1) BH 1.6-2.2 (1.9 ± 0.4) 1.6-2.6 (2.2 ± 0.4) 2.4-3 (2.8 ± 0.3) 3.2-3.3 (3.3 ± 0.1) TAL 5.5-11.7 (8.6 ± 4.4) 6-10 (7.7 ± 1.7) 8.9-11.5 (10.3 ± 1.3) 10.8-13.4 (12.1 ± 1.8) Eye 0.56 0.6-0.7 (0.6 ± 0.1) 0.7-0.8 (0.7 ± 0.1) 0.8-0.9 (0.8 ± 0.1) Nostril 0.2 0.2 0.2 0.2 IOD 1.1-1.4 (1.3 ± 0.2) 1.3-1.6 (1.4 ± 0.1) 1.7-1.9 (1.8 ± 0.1) 2 IND 0.7-0.9 (0.8 ± 0.1) 0.8-1 (0.9 ± 0.1) 0.9-1.3 (1.2 ± 0.2) 1.4-1.5 (1.4 ± 0.1) SND 0.2-0.4 (0.3 ± 0.1) 0.4-0.7 (0.5 ± 0.2) 0.4-0.6 (0.5 ± 0.1) 0.9-1 (0.9 ± 0.04) END 0.4-0.5 (0.4 ± 0.03) 0.4-0.5 (0.4 ± 0.1) 0.5-0.7 (0.6 ± 0.1) 0.7-0.8 (0.7 ± 0.1) SL 0.6-1 (0.8 ± 0.3) 0.7-1.2 (0.9 ± 0.2) 0.8-1 (0.9 ± 0.1) 1.2-1.3 (1.3 ± 0.1) SH 0.5-1 (0.8 ± 0.3) 0.6-1.1 (0.9 ± 0.2) 0.8-1.1 (1 ± 0.1) 1.4-1.5 (1.4 ± 0.1) SOH 0.1-0.3 (0.2 ± 0.1) 0.1-0.2 (0.2 ± 0.04) 0.2-0.3 (0.3 ± 0.02) 0.4 SSD 2-3.3 (2.7 ± 0.9) 2.7-3.4 (3.1 ± 0.3) 3.8-4.8 (4.3 ± 0.5) 5.3-5.5 (5.4 ± 0.1) VT 0.4 0.4-0.5 (0.4 ± 0.03) 0.6-0.9 (0.8 ± 0.1) 1.0 TMW 0.8-0.9 (0.9 ± 0.1) 0.8-1.2 (1 ± 0.2) 1.1-1.7 (1.4 ± 0.3) 1.8 TMH 1-1.3 (1.2 ± 0.2) 1-1.3 (1.1 ± 0.2) 1.5-1.8 (1.7 ± 0.2) 2-2.1 (2.1 ± 0.1) ODW 1.4-2.2 (1.8 ± 0.2) 1-1.6 (1.4 ± 0.3) 1.5-2 (1.8 ± 0.3) 2.1-2.4 (2.3 ± 0.2) AL gap 0.7-1 (0.8 ± 0.2) 0.8-1 (1 ± 0.1) 1.1-1.2 (1.2 ± 0.1) 1.4-1.6 (1.5 ± 0.2) UJW 0.5-0.6 (0.5 ± 0.1) 0.5-0.8 (0.6 ± 0.1) 0.7-0.9 (0.8 ± 0.1) 0.9-1.1 (1.0 ± 0.1) LJW 0.3 0.3-0.5 (0.4 ± 0.1) 0.6-0.7 (0.6 ± 0.1) 0.7-0.8 (0.7 ± 0.1) A-2 gap 0.3 0.3 0.3-0.4 (0.3 ± 0.03) 0.4 P-AL 5 5-7 7-9 10-15 P-PL 23-25 13-19 14-17 16-18 56 Hernán Darío Granda-Rodríguez et alii pole in stage 26 that have myotomes as high as wide. The myotomes are straitened gradually towards the tip of the tail. At mid-tail, dorsal fin barely higher than ventral fin and myotomes higher than dorsal fin. Dorsal fin originat- ing at body-tail junction; dorsal fin lower on first ante- rior third of the tail, posteriorly of same height through tail length, sometimes higher at mid-tail. Ventral fin of same height through all of tail length. Tip of tail rounded. Maximum tail height less, equal to, or higher than body height, 88-116% of body height. Lateral line system symmetrical on both sides of body with supraorbital, anterior pit, infraorbital, oral, temporal infraorbital, supratemporal, dorsal, middle and ventral trunk lateral lines. Lateral lines were visible from stage 27 (except for one tadpole in stage 26 with supraor- bital lateral line). In one tadpoles in stage 27 only the supraorbital, infraorbital lateral, dorsal and middle trunk line are present, and in other tadpoles (stages 27-29) the lines already mentioned are observable. The stitches in all lateral lines are equidistant between each of them and predominantly rounded (see additional changes below). Pale white spot(s) (cumuli of neuromasts), and pale line between eye-nostril are absent. Oral disc positioned and ventrally directed, later- ally emarginated, and surrounded by marginal papillae, except on anterior lip; sub-marginal papillae absent (Fig. 3D, E). Oral disc width 42-52% of body width. Ante- rior lip gap 60-88% of oral disc width. Marginal papil- lae arranged in rows varying by lip and stage from 1-2 rows (see below). Papillae abundant on anterior and posterior lip, 5-7 papillae (stages 25-27), 15 (stage 28), and 10 (stage 29) on right side of anterior lip, and 23-25 papillae on entire posterior lip (stage 25, due that poste- rior lips is less than 1 mm wide) and from 13-19 papil- lae/mm on posterior lip (stages 26-29). Papillae low and narrow on anterior and posterior lip, barely elongated, papilla width (at base) third quarter or two third than papilla height. Tips of papillae rounded to sub-acuminate. Labial tooth row formulae (LTRF) 2(2)/3; second upper row (A-2) with moderate gap, from 17-28% of oral disc width. Length of first upper row (A-1) equal to second upper row (A-2). First lower row (P-1) equal to second and third lower row (P-2, P-3) (n = 4); P-1 equal to P-2 and P-3 slightly shorter than P-1 and P-2 (n = 4); or P-1 equal than P-3 and these slightly shorter than P-2 (n = 3). Upper rows sub-equal (commonly) or slightly short- er (rarely) than first and second lower rows. Jaw sheath keratinized and black or dark brown. Upper jaw sheath (UJS) low in height, with long and thin lateral processes; UJS height 11-18% of UJS width. UJS width 37-50% of oral disc width. Anterior edge of UJS well defined (com- monly) or indistinct (rarely), forming a convex shape; posterior free edge of UJS barely sinuous, with low ser- rations; mid-UJS with very low notch (Fig. 3E), 20-22% of UJS height. Lateral processes without serrations. Tips of serrations rounded on middle of UJS and acuminate to sub-acuminate towards ends. Shelf on concealed UJS absent. Lower jaw sheath (LJS) thin, V-shaped with prox- imal ends slightly open. Only distal half of LJS pigmented black, LJS height (black pigmented area) 50-100% of UJS height. Serrations on LJS low and with rounded tips. In preservative, the skin of all body of the free-swim- ming tadpoles is translucent; body dorsally and laterally with abundant dark brown (stages 25-27) or less brown dots (stages 28-29), distributed uniformly on body, without darkening the body, forming a dark brown region around of nostril. Ventrally with dark brown dots (stages 25-27) or brown dots (stages 28-29), distributed uniformly from pos- terior oral disc to middle intestines, without darkening the venter. Under magnification, the dark brown dots of venter are asterisk-like (stages 25-27) or rounded (stages 28-29) dots (Fig. 3B). Intestines yellowish brown, visible in lateral Fig. 3. Tadpole in stage 27 of Allobates ignotus, in dorsal (A), ven- tral (B), and lateral (C) view. Oral disc of a tadpoles in stage 26 (D) and in stage 29 (E). See the upper jaw sheath shape (D) and the lower tooth rows (E). Scale bar equal to 1.0 mm (A, B, C) and 0.5 mm (D, E). Photographs by M. Anganoy-Criollo. 57Natural history and conservation of Allobates ignotus and ventral views and partially in dorsal view. Myotomes white and fins white translucent with brown dots covering the dorsal fin, myotomes and dorsal portion of the ven- tral fins, distributes uniformly (stages 25-27) and forming some weak reticulate pattern (stages 28-29). Nostril white; spiracle and tube vent translucent and oral disc white with a few brown dots on anterior lip. The variation in free-swimming tadpoles (stages 25-29) (Table 3) is as it follows: in dorsal view, one tad- pole in s25 has the body slightly more rounded than the rest of tadpoles. The marginal papillae row number var- ies between stages 25 and 29. In the anterior lip, the mar- ginal papillae are disposed in one row from stages 25 to 29. In the posterolateral side of posterior lip, the marginal papillae are arranged in one row in stage 25-26, one bise- riated row in stage 27 and two row in stage 28-29. On the posterior region of posterior lip, the marginal papillae are in one row in stages 25-27 and 29, in one biseriated row in one tadpole in stage 25 and 27, and in two rows in stage 28. The lateral line system also varies between these tad- poles. The supraorbital lateral, dorsal and middle trunk lines are incomplete. The supraorbital lateral line of one tadpole in stage 26 and other in stage 27 extends from oral disc to nostril; whereas, the dorsal and middle trunk lines of the stage 27 ranges from body to mid-tail. The stitches are rounded in supraorbital, anterior pit, infraor- bital, temporal infraorbital, supratemporal lateral and dorsal trunk lines, but are slightly elongate and paral- lel to the line in the oral lateral and middle trunk lines, although in middle trunk line the stitches are ovoid in the stage 29. The stitches are oval in ventral trunk line. Advertisement call The advertisement call of Allobates ignotus is com- posed by a series of pulsed notes (x = 32; SD = 27.44; range = 6-116 notes; n = 13 calls), with a mean duration of 8.168 sec (SD = 6.36; range = 1.35-27.53 sec; n = 13 calls). The mean duration of the note was 0.046 sec (SD = 0.009; range = 0.026-0.085 sec; n = 456 notes), emitted at a rate of 3.94 notes per sec (SD = 0.61; range= 2.32-4.46), in an internote interval of 0.211 sec (SD = 0.110; range = 0.043-1.790 sec; n = 443) (Fig. 4b). The call frequency was between 4600-14000 Hz, and the dominant frequen- cy was 4942.82 Hz (SD = 204.3; range = 5312-6662Hz; n = 13). Pulse has three distinct harmonics, 7490-7993 Hz, 9784-10065 Hz, and 12834-13132 Hz, respectively (Fig. 4a). Our observations in La Veguita Creek indicate that A. ignotus may be a philopatric species regarding to vocalization site, since an uncollected male vocalized for four hours approximately on the same rock. When this frog perceived disturbances in the vegetation caused by our capturing attempts, it escaped quickly, but returned in a few minutes to continue vocalizing in the same place. This situation was repeated about six times. Relative abundance The surveys performed from 2010 to 2014 showed an abrupt decrease in relative abundance of Allobates ignotus in Canime Creek Basin in the last year (Fig. 5). In April 2010, we recorded 0.46 ind/h × obs (n = 25, rare) of this frog; while in February 2011 observed 0.55 ind/h × obs (n = 30, common); in August 2012, we obtained 0.92 ind/h × obs (n = 50, abundant); and April 2013 we calcu- Table 3. Variation of the rows of marginal papillae of the oral disc of tadpoles of Allobates ignotus through of the available stages of development. Whether there was variation, this is reported as range. Conventions: (A) papillae on anterior lip, (B) papillae on postero-lateral side of posterior lip, and (C) papillae on middle of posterior lip; (1) one row, (2) one biseriated row, (3) two rows of papillae, (n) number of tadpoles. Stages refer to Gosner (1960). Stages (n) A B C 25 (2) 1 1 1-2 26 (4) 1 1 1 27 (3) 1 2 1-2 28-29 (2) 1 3 1-3 Fig. 4. Spectrogram (a) and oscillogram (b) of the advertisement call of Allobates ignotus of La Veguita Creek, La Veguita Village, Manuaure Balcón del Cesar, Cesar department, Colombia. Air tem- perature 25.4 °C (CBUMAG:ANF:00933). Total duration = 1.70 sec, 512 bands resolution. 58 Hernán Darío Granda-Rodríguez et alii lated 1.47 ind/h × obs (n = 80, very abundant). Neverthe- less, in June 2014, we found 0.04 ind/h × obs (n = 2, very rare). Although we did not find tadpoles or amplectant couples in any of the fieldtrips to Canime Creek Basin, we observed and listened males vocalizing in all field expeditions in this basin, except to June 2014. DISCUSSION Geographic distribution and habitat use Allobates ignotus was previously known in only three localities in the low wet forest of the western foothills of the Serranía del Perijá, with an extent of occurrence of 364 km2 and 180 m altitudinal range (400 to 580 m a.s.l., Anganoy-Criollo, 2012; IUCN SSC Amphibian Specialist Group, 2017). With the records reported in the present study, we extended the distribution range of this frog by 241 km2, with a total extent of occurrence at 605 km2 and a altitudinal range from 194 m to 1236 m. This records are the first for the Tropical Dry Forest Life Zone and for the department of La Guajira. The lowest localities where A. ignotus was found fall within the altitudinal range (0-249 m a.s.l.) with the greatest amphibian diversity in Colombia (284 species according to Acosta-Galvis, 2012). Amphibians from dry environments require physi- ological, ecological, and behavioral strategies for dehy- dration tolerance (Urbina-Cardona et al., 2014). Habi- tat selection is an essential aspect for amphibians of dry ecosystems by the occupation of humid microhabitats in dehydrating environments (Bentley, 1966). Little is known about habitat use and mechanism to avoid the drying of dendrobatoids frog from tropical dry forests. Nevertheless, some species such as Dendrobates truncatus (Dendrobatidae) have relatively well described ecological and behavioral strategies that allow them to withstand drought conditions in Caribbean tropical dry forest. Strategies like displacement to humid refuges or per- manently staying in them allow taking advantage of the temperature and humidity provided by some microhabi- tats available (Cuentas et al., 2002). We do not know the strategies used by A. ignotus for dehydration tolerance, but this frog is active all year round (we found vocaliz- ing males in dry and rainy season). It is possible that the occupation of rocky substrate and leaf litter by A. igno- tus corresponds to some mechanism for temperature and humidity capture (Feber, 1982; Navas, 1996). However, this hypothesis needs additional studies that evaluate the physical and structural variables of the habitat that influ- ence the distribution and density of the frogs (Giaretta et al., 1999; van Sluys et al., 2007). Free-swimming tadpoles Prior to Grant et al. (2006), most species of Allo- bates were grouped in the Colostethus sensu lato. Colo- ma (1995) described the larval external morphology of the species that were allocated in Colostethus sensu lato. Afterwards, Anganoy-Criollo (2013) provided larval features that are shared between the species previously included in Colostethus sensu lato and now split in Allo- bates, Aromobates, Mannophryne, Rheobates (Aromo- batidae), Colostethus and Hyloxalus (Dendrobatidae). The general morphology of the tadpoles of A. ignotus resembles well the one described in Coloma (1995) and in Anganoy-Criollo (2013). However, both these works report that A-2 gap is short or narrow, though explic- it measurements of A-2 gap are not provided in these descriptions. On the opposite, the A-2 gap of Allobates ignotus appears nor short or narrow. Anganoy-Criollo (2013) tells about a narrow or short A-2 gap of Hylox- alus subpunctatus (Dendrobatidae) being 4-15% of OD width, while the A-2 gap of A. ignotus is 17-28% of OD width, therefore markedly wider. Previous descriptions of tadpoles of other Allobates also showed “wide” A-2 gap; for example, in A. brunneus (30% of ODW; Lima et al., 2009), A. caeruleodactylus (35% of ODW; Caldwell et al., 2002), A. femoralis (21-35% of ODW; Lescure, 1976; Silverstone, 1976 ), A granti (52% of ODW; Kok et al., 2006), A. grillisimilis (30% of ODW; Simões et al., 2013), A. hodli (23% of ODW; Simões et al., 2010), A. kingsbur- gyi (38% of ODW; Castillo-Trenn, 2004), A. magnussoni (22% of ODW; Lima et al., 2014), A. niputidea (although not explicitly reported in the text of the description, but evident from the drawing of the tadpole; Grant et al., 2007), A. paleovarzensis (35% of ODW; Lima et al., 2010), A. subfolionidificans (23% of ODW; Lima et al., 2007), A. sumtuosus (25% of ODW; Simões and Lima, 2012), A. Fig. 5. Temporal variation of relative abundance of Allobates ignotus in Canime Creek Basin, Cesar department, Colombia. 59Natural history and conservation of Allobates ignotus talamancae (35 or 31% of ODW; Breder, 1946; Savage, 2002), and A. tapajos (25% of ODW; Lima et al., 2015). In Dendrobatoidea, this wide A-2 gap was also found in some species of dendrobatid genus Andinobates (Sil- verstone, 1975; Myers and Daly, 1980; Ruiz-Carranza and Ramírez-Pinilla, 1992), Ameerega (Haddad and Martins, 1994; Twomey and Brown, 2008; Poelman et al., 2010), some Phyllobates (Siverstone, 1976; Donnelly et al., 1990; Savage, 2002), and Ranitomeya (Brown et al., 2008; von May et al., 2008; Twomey and Brown, 2009; Perez-Peña et al., 2010). The observed variability of the A-2 gap among the above genera suggests it is a potentially mean- ingful trait from both a taxonomic and phylogenetic per- spective; therefore, an adequate characterization of A-2 gap width variability in the superfamily Dendrobatoidea would be worthwhile. The systematic value of the larval morphology has been used in the phylogenetic relationships history of poison frogs (e.g., Savage, 1968; Silverstone, 1975, 1976; Myers, 1987; Grant et al., 2006; Sánchez, 2013) showing that there is correspondence between larval features and dendrobatoids taxonomy (e.g., reduction in labial tooth row, oral disc without emargination, umbelliform oral disc, notch on UJS). At this moment, despite that the “wide” A-2 gap is present in several poison frog genera, the “wide” A-2 gap of the tadpoles of A. ignotus is shared with other Allobates species, which supports the assig- nation of A. ignotus within the genus Allobates, where Anganoy-Criollo (2012) provisionally located this spe- cies based on adult characters. Molecular and phenotypi- cal characters known for the genus Allobates (Grant et al., 2006, 2017) suggest that the “wide” A-2 gap may be a derived and shared character for this genus; however a phylogenetic analysis is needed to test this hypothesis. Advertisement call Comparisons of advertisement calls within others species of the genus Allobates are limited by the reduced number of calls described. Calls of most of the Colom- bian Allobates are unknown. The advertisement call of A. ignotus is composed by a sequence of pulsed notes (6-116 notes/call) with ascending modulation frequency, like to other aromobatid frogs as A. talamancae (5-15 notes/call, Lechelt et al., 2014), A. humilis (7-16 notes/call, La Marca et al., 2002), A. myersi (5-10 notes/call, Simões and Lima, 2011), A. paleovarsensis (3-21 notes/call, Lima et al., 2010), and Aromobates saltuensis (2 notes/call, Barrio-Amorós and Santos, 2012), from which it differs by the greater number of notes per call. Although in the genus Allobates the advertisements calls are highly variable, some species converge their calls in the long trains of notes, such as A. crombiei (25-59 notes/call, Lima et al., 2012), A. goianus (2-41 notes/call, Carvalho et al., 2016), and A magnussoni (not specified in the call description, Lima et al., 2014). Some Allobates can produce their advertisement calls at a dominant frequency similar to A. ignotus (4942 Hz), such as Allobates algorei (5065 Hz, Barrio-Amorós and Santos, 2009), A. femoralis (2853 Hz), A. goianus (4946- 5230.79 Hz, Bastos et al., 2011; Carvalho et al., 2016), A. humilis (4200 Hz, La Marca et al., 2002), A. myersi (3400 Hz, Simões and Lima, 2011), A. sumptuosus (6500 Hz, Kok and Ernst, 2007), A. aff. brunneus (5450 Hz, Lötters et al., 2003); however, these species occupy distant geo- graphic areas and are easily morphologically distinguisha- ble from A. ignotus. On the other hand, only two aromo- batid frogs geographically close to A. ignotus have adver- tisement calls described, A. talamancae and Rheobates palmatus. The advertisement call of A. talamancae dif- fers from that of A. ignotus by the slightly higher domi- nant frequency (5036 Hz, Lechelt et al., 2014). Despite the advertisement call of R. palmatus consists of a long train of notes, this differs from the advertisement call of A. ignotus by the lower dominant frequency (2083-2700 Hz, Lüddecke, 1999; Bernal et al., 2006). For other den- drobatoid with trans-Andean distribution or geographi- cally close to A. ignotus, such as A. niputidea (Grant et al., 2007), A. wayuu (Acosta-Galvis et al., 1999), Aromo- bates aff. saltuensis (Anganoy-Criollo, 2012, see speci- mens examined), Aromobates totuko (Rojas-Runjaic et al., 2011), Colostethus inguinalis (Grant, 2004), and “Coloste- thus” ruthveni complex (Granda-Rodríguez et al., 2014); there are no descriptions of their advertisement calls. Relative abundance and conservation status Habitat degradation and fragmentation are the prin- cipal factors in the amphibian decline and extinction, since they generate drastic changes on the amphibian community structure (Mazerolle, 2003). Some dendro- batoid frogs are very sensitive and have declined their populations due to forest fragmentation and degradation, e.g., A. juanii in the eastern foothills of the Cordillera Oriental of Colombia (Cáceres-Andrade and Urbina-Car- dona, 2009) and Anomaloglossus stepheni (Aromobatidae) in the central Amazon (Funk and Mills, 2003). On the other hand, dendrobatoid frogs like Epipedobates bouleng- eri (Dendrobatidae) can tolerate structural changes on their habitat, although environment variables like micro- habitat temperature are strongly associated with the rela- tive abundance of this frog, conditioning their establish- ment (Urbina-Cardona and Londoño-Murcia, 2003). In the three localities reported herein, Allobates igno- tus showed fluctuations in their relative abundance dur- 60 Hernán Darío Granda-Rodríguez et alii ing the study period, and we found the lowest abundance in the last year. These areas are immersed in a zone of hydrocarbons exploitation activity. Fragmentation and degradation of riparian forest of Canime Creek by min- ing activity, plus the stream flow reduction by severe droughts (recent and predicted for future, see Nakae- gawa and Vergara, 2010), can be both short and long term threats, due to the decrease of habitat availability and quality. In addition, the influence of pollution with chemical and solid mining wastes on populations of these frogs should be assessed, since these factors are known to affect the development and survival of amphibians (Rowe et al., 1996). All these potential threats can act synergisti- cally and cause population declines in A. ignotus. Although Allobates ignotus was previously catego- rized as endangered species [EN B1ab(iii), IUCN SSC Amphibian Specialist Group. 2017], this decision was based on the limited data provided by Anganoy-Criollo (2012). Therefore, we re-evaluate the conservation status of the species with the data provided herein. Following the IUCN criteria (IUCN, 2012), we suggest categorizing Allobates ignotus as Vulnerable [VU B2ab (iii)]; given to the decrease of the relative abundance in the three locali- ties of the Canime Creek Basin, the extent of occurrence lesser than 2000 km2 (605 km2), and loss of habitat qual- ity by degradation of creeks and forests where this species lives. ACKNOWLEDGEMENTS Some specimens were collected under the project “Estudios técnicos, sociales, económicos y ambientales, con su respectiva cartografía temática, correspondi- ente al entorno local, del complejo de páramo Perijá en jurisdicción de la Corporación Autónoma Regional del Cesar (CORPOCESAR) y la Corporación Autónoma Regional de la Guajira (CORPOGUAJIRA)”. This project is excluded of permits according to the Decree 1376 at 2013, in the frame of a scientific and technologic coop- eration agreement between the Instituto de Investiga- ciones Biológicas Alexander von Humboldt (IAvH) and Fundación Prosierra. We thanks to Adolfo del Portillo Mozo, Efrain Rada Vargas, and Miguel Arévalo Páez for donating us the specimens collected in La Frontera’s way and Manaure Balcón del Cesar. Thanks to Ruben Fon- talvo, José Rincón, and Carlos Ardila Rodríguez for his great help on the field trip to Manaure Balcón del Cesar. Likewise, to Alina Gámez, Ángel Santana and family, Joán Zambrano, and the Direction of the Escuela Nor- mal Superior María Inmaculada for helping us in our stay in Manaure Balcón del Cesar. We are grateful to Lorena Benítez-Cubillos for construction of the map and Gra- ham Louis for lenguage revision. Speciall mentios goes to Patricio Hernáez, Martín Caicedo, Andrea Cotes, Carlos Mario López Orózco, Santiago González, María V. León, Rosana Londoño, and Sigmer Quiroga for their invalu- able contributions on the manuscript. REFERENCES Acosta-Galvis, A.R. (2012): Anfibios de los enclaves secos en la ecorregión de La Tatacoa y su área de influen- cia, alto Magdalena, Colombia. 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(2008): The tadpole of the bamboo-breeding poison frog Ranitomeya biolat (Anura: Dendrobati- dae): Zootaxa 1857: 66-68. Zimmermann, H., Zimmermann, E. (1988): Etho-taxon- omie und zoogeographische artengruppenbildung bei pfeilgiftfröschen (Anura: Dendrobatidae). Salamandra 24: 125-160. APPENDIX Examined specimens Allobates ignotus. - Colombia: Cesar: La Jagua de Ibirico: Zumbador River (580 m a.s.l.), Nueva Granada Village, ICN 55434 (holotype), ICN 55435-436 (para- types); El Indio Creek (520 m a.s.l.), La Victoria de San Isidro Village, ICN 55427-433; Canime Creek 1 (250 m a.s.l.), CBUMAG:ANF:00254-256; Canime Creek 2 (200 m a.s.l.), CBUMAG:ANF:0069; Canime Creek 3 (194 m a.s.l.), CBUMAG:ANF:00270-272. Becerril: El Veranero Creek (400 m a.s.l.), El 11 Village, ICN 55437-439 (para- types). Agustín Codazzi: La Frontera’s way (1236 m a.s.l.), La Frontera Village, CBUMAG:ANF:00937. Manaure Bal- cón del Cesar: La Veguita Creek (633 m a.s.l.), La Veguita Village, CBUMAG:ANF:00933, 00935-936. La Guajira: La Jauga del Pilar, Nicaragua Creek, El Plan Village, CBUMAG:ANF: 00934. 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