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Andinobates victimatus sp. nov. Figures 2 and 4 A–K. Table 1. Holotype. An adult female (ANDES-A 3686) collected by Daniel Mejía-Vargas in Chichiridó, Dabeiba, Antioquia (7.02°, -76.38°), located in the Llorona Canyon, at 280 meters above sea level (m.a.s.l.). Paratypes. Five adults (4 males and 1 female; ANDES-A 3684, 3685, 3687–3689), collected at the type locality, along with the holotype. Etymology. From the Latin victimatus (the victimized); noun in apposition. The Urabá region, where this species occurs, has historically been flailed by Colombia’s longstanding armed conflicts, perhaps more than most other regions of the country, leaving a virtually countless trail of innocent victims. We name this species in honor and remembrance of all these victims. Definition and diagnosis. Andinobates victimatus sp. nov. is a minute (SVL mean = 14.23 ± SD = 0.67 mm; Table 1) dendrobatid frog, characterized by having homogeneous scarlet red dorsal coloration, which extends throughout the limbs. Fingers and toe tips are light metallic gray to metallic gray. The venter and flanks are also scarlet red, with patches degrading into vermilion and finally black, forming irregular blotches (Fig. 4 A–F). The skin is slightly granular, with granularity increasing towards the urostyle and hind limbs. This species is assignable to Andinobates based on the following traits: small adult size (SVL Andinobates, we place A. victimatus sp. nov in the A. fulguritus group (Brown & Twomey et al. 2011), also based on phylogenetic analyses, juvenile color pattern (Fig. 4 H–K; see Coloration in life), and calls of the “short buzz” type (Brown & Twomey et al. 2011; Fig. 5). The new species can be easily distinguished from most other known members of Andinobates by its distinctive solid red dorsal coloration (Fig. 4). The only exception is the recently described A. geminisae (Batista et al. 2014), which has uniform orange dorsal coloration, but lacks light gray/tan pigmentation on finger and toe tips, has uniform ventral coloration, and produces calls of the “tonal buzz” type, with distinctive clicks at the beginning and end of the call, as opposed to the “short buzz” calls of A. victimatus. Moreover, calls of A. victimatus sp. nov. are shorter in duration (0.42– 0.58 s vs. 1.52– 1.6 s) and higher in peak frequency (5.65–5.86 kHz vs. 4.40–4.50 kHz; Fig. 6) than those of A. geminisae. Finally, A. geminisae is, on average, smaller than A. victimatus sp. nov. (SVL 12.68 ± 0.87 mm vs. 14.23 ± 0.67 mm; t 15 = 3.78, p = 0.002, N = 6 A. vic, 11 A. gem; Fig. 6). However, the SVL ranges of both species overlap slightly (A. geminisae: 11.63–13.63 mm, A. victimatus sp. nov.: 13.22–14.98 mm). Some individuals of A. opisthomelas, A. virolinensis, and A. cassidyhornae can have nearly homogenous red dorsal coloration (Amézquita et al. 2013; Ruiz-Carranza & Ramírez-Pinilla 1992; Silverstone 1975). However, they can be distinguished from A. victimatus sp. nov. by their ventral pattern which consists of red blotches on a black background in A. cassidyhornae, light gray spots/marbling in A. virolinensis and some populations of A. opisthomelas, and dark brown with irregular suffusions of red from the flanks in other populations of A. opisthomelas. Moreover, the three species are, on average, larger than A. victimatus (A. cassidyhornae: 19.031 ± 0.31 mm, t 11 = 17.04, p A. vic, 7 A. cas; A. virolinensis: 16.86 ± 0.91 mm, t 58 = 6.85, p A. vic, 54 A. vir; A. opisthomelas: 16.80± 1.24 mm, t 53 = 5.14, p A. vic, 49 A. opi; data from original descriptions and Silverstone 1975), although, again, with slight range overlap with A. virolinensis (13.22– 14.98 mm vs. 14.6–18.9 mm) and A. opisthomelas (13.22–14.98 mm vs. 14.5–19.5 mm). In addition to coloration, A. victimatus sp. nov. can be distinguished from other lowland species of Andinobates (i.e. A. minutus and A. fulguritus groups) in the following ways (Fig. 6): The calls of A. victimatus sp. nov. are shorter than those of A. claudiae (0.42– 0.58 s vs. 0.85– 1.03 s), and have a higher peak frequency than those of A. fulguritus (5.65–5.86 kHz vs. 4.83–5.16 kHz); A. victimatus sp. nov. is smaller in body size than A. altobueyensis (13.22–14.98 mm vs. 16.30–17.00 mm). Description of the holotype. An adult female, 13.22 mm in snout-vent length. Snout dorsally subtruncate, laterally rounded and slightly protrusive; canthus rostralis rounded; loreal region vertical and slightly concave. The head is wider than it is long, and the same width as the body (HW/GBW = 1.002); head width 29.5% of SVL; head length 25.0% of SVL; nares 1.24 times closer to tip of snout than to eyes; pupil rounded and horizontally elliptical; eye length 52.6% of head length. Tympanic ring visible, oval shaped, of moderate size (HDT 41.4% of EL); posterior and dorsal margins hidden under skin. Maxillary and premaxillary teeth absent; median lingual process absent; tongue about two times longer than wide; anterior margin of tongue not indented; posterior third of tongue not attached to floor of mouth. Hands of moderate size (HaL 26.6% of SVL); adpressed finger lengths are I In life, the dorsum was homogeneous scarlet red, with faintly visible dorsolateral stripes; flanks were mostly uniform vermilion, with a few small, dark specks towards the ventral area. The venter was also vermilion with lesssaturated blotches transitioning from vermilion to black towards the chest and throat. Fingers and toes were light metallic gray, and palms and soles were black. The skin was slightly granular, especially in the posterior venter, cloaca, and dorsal surface of hind limbs. In ethanol (70%), the dorsum and dorsal surfaces of limbs are metallic gray; hands, feet, and articulations are tan to light brown. The venter and flanks are also metallic gray, with some scattered dark brown specks; a large dark brown blotch is present on the throat. The skin is mostly smooth, except for the ventral surface of the hind limbs and cloaca, where weak granularity is still visible. Coloration in life (Figs. 4, 4 A–K). The dorsum and dorsal surfaces of limbs of all individuals in the type series are homogenous scarlet red. In some collected and uncollected individuals, a faint, complete dorsolateral stripe (type A in Grant et al. 2006) of a more saturated shade of red is visible (Fig 4 A-D). The scarlet red becomes less saturated in the flanks and venter, transitioning between scarlet-red, vermilion, and black in gradually changing blotches, especially in the throat, chest, and the dorsal surface of the thighs (forming a seat patch; Fig 4 E–F). The pupil and iris are both black, and nearly indistinguishable. The nares are surrounded by thin black margins, and the anterior edge of the tympanic ring is also black, with the first quarter of the tympanum degrading from black to vermilion. Fingers and toes are light metallic gray to metallic gray, and palms and soles black to dark brown. Juveniles reared in the laboratory (Fig. 4 H–K) developed tan/orange dorsolateral stripes as metamorphs (around stage 40), and later acquired a faint incomplete dorsal median stripe that extended from the tip of the snout to the posterior-dorsal region, and a labial stripe extending from just below the snout to the insertion of the arm, both reminiscent of the color pattern of some A. fulguritus. As time went by, the stripes became a more saturated red, and the dark brown background gave way to homogenous scarlet red pigmentation. Adult coloration was attained around 3.5 months after leaving the water. These individuals were reared at a lower temperature (~18–22 °C) than their natural habitat (~22–28 °C), so this process is probably quicker in nature. Coloration in preservative (70% ethanol). In preservative, the red color turns metallic gray. Fingers, toes, and tubercles become light tan/brown color, and the pupil and cornea become light gray to white. The heterogeneity in ventral coloration becomes more defined in preservative, as the black blotches become dark brown, or stay black, while the scarlet red/vermilion background becomes metallic gray. Vocalization (Figs. 5–6). Similar to other members of the A. fulguritus group, A. victimatus sp. nov. produces calls of the “short buzz” type, consisting of a series of pulses modulated in amplitude but not frequency (Brown & Twomey et al. 2011; Myers & Daly 1976). Calls range from 0.42– 0.58 s in length (mean = 0.49 ± SD = 0.06 s), and contain 99–190 pulses per call (142 ± 42.6). The peak frequency ranges between 5.65–5.86 kHz (5.75 ± 0.08 kHz), and the frequency interquartile bandwidth between 0.76–1.03 kHz (0.88 ± 0.11 kHz). Natural history. A. victimatus sp. nov. is a terrestrial species, which inhabits the understory of lowland forests in the Urabá region of Northwestern Colombia (Fig. 7). Males call to defend territories and attract mates, and were typically observed doing so from the leaf litter, or perched on low branches ( Egg clutches are presumably laid within male territories, and tadpoles are transported by males, one at a time, to bromeliad tanks. Tadpoles were observed in bromeliads up to 4m above the ground (although we did not survey plants above this height). All of the observed tadpoles were in tanks of Guzmania musaica bromeliads (Fig. 7 D), which leads us to speculate that A. victimatus sp. nov. prefers this bromeliad over other similarly-sized species (e.g. other Guzmania or Tillandsia), for yet unknown reasons. At the type locality, A. victimatus sp. nov. is sympatric with Oophaga histrionica (Anura: Dendrobatidae), which deposits larvae in similar phytotelmata. However, we did not find any O. histrionica tadpoles in G. musaica tanks; only in smaller bromeliad species. Whether this indicates some instance of resource partitioning, competition, or another kind of ecological interaction between the two species remains to be studied. Distribution and conservation status. This species occurs in the Urabá region of Northwestern Colombia, inhabiting tropical rain forests at ~ 200– 600 m.a.s.l. (Figs. 1, 7) in the foothills of the Western Andes, west of the Paramillo Massif, on the Atrato River drainage. The type locality is in the Llorona Canyon in Dabeiba, Antioquia, Colombia, just southwest of the Massif. We have become aware of reports of similar frogs from other biologists and conservationists working in the Urabá region (Carlos Bran-Castrillón, pers. com.), which would extend the new species’ distribution north into the municipalities of Mutatá and Carepa, almost to the Gulf of Urabá, on the Caribbean coast (Fig. 1). In addition, two specimens at the Natural History Museum of the Universidad de Antioquia (MHUA-A 3382 and 3864), currently assigned to A. opisthomelas, are from the municipality of Tierralta, Córdoba, at ~ 590 m. a.s.l (Fig. 1). It is possible that these two specimens actually correspond to A. victimatus sp. nov., since A. opisthomelas does not normally occur below 1100 m.a.s.l (Silverstone, 1975), which would mean that this species is distributed around the northern end of the Paramillo Massif, into the Sinú River drainage. As we have not examined specimens, alive or preserved, from any of these localities, we consider them tentative reports. Andinobates victimatus sp. nov. has, to the best of our knowledge, not been reported south or west of the type locality. However, given the continuous rain forests and gentle topography of the Northeastern Chocó, it is likely that this species does extend into these areas. Nonetheless, Silverstone did not find it when collecting A. fulguritus in the Arquía river, ~ 85 km south of the type locality, in the 1960s, and it has not been reported in the vicinities of Quibdó (~ 130 km south) or the San Juan River drainage (> 180 km south), both well sampled in terms of Dendrobatid frogs. Therefore, we suspect that this species’ range does not extend too far south of the type locality. Further exploration and inspection of museum specimens are needed to get a better grasp of this species’ geographic distribution. In spite of the current uncertainty regarding the distribution of A. victimatus sp. nov., it seems clear that its extent of occurrence is less than 5,000 km 2. It is currently known from one locality with certainty, plus five tentative additional ones, and the forests in this species’ range have been, and will probably continue to be under strong deforestation pressures due to mining (both legal and illegal), cattle farming, and large-scale agriculture (e.g. banana and plantain plantations), which reduce the amount and quality of available habitat, and increase its fragmentation. Therefore, we tentatively designate A. victimatus sp. nov. as Endangered (EN: B1a, biii; IUCN, 2001). However, it is possible that the range of A. victimatus sp. nov. does exceed 5,000 km 2, in which case this species should be listed as Vulnerable (VU; B1a, biii). Most of the range of A. victimatus sp. nov. surrounds the Nudo de Paramillo National Natural Park, a 460,000 ha nature reserve, managed by the Colombian government, which is closed to the public. However, we are unaware of any reports of A. victimatus sp. nov within the park, which encompasses most of the Paramillo Massif above 700 m. a.s.l. This national park, nonetheless, protects ~250,000 ha in the 700–1000 m.a.s.l elevation belt, where A. victimatus could exist at the higher end of its elevational range. Considerably more research on this species’ distribution, habitat requirements, and population ecology is needed to confidently assign it to a category of threat. Voucher Species Locality Study GenBank Accession 16S COI Cytb ……continued on the next page Voucher Species Locality Study GenBank Accession 16S COI Cytb GECOH 1216 Andinobates opisthomelas I la đel SƟl, Guatapé, AntiƟquia, Amézquita et al. JQ936634 KY885051 JQ936620 CƟlƟmbia 2013 GECOH 297 Andnobates bombetes Bosque YƟtƟcƟ Natural Re erve, YƟtƟcƟ, Thi tuđy KY885045 KY885067 KY885092 Valle đel Cauca, CƟlƟmbia GECOH 305 Andinobates virolinensis CƟ tilla đe fara, VirƟlín, Santanđer, ƂrƟwn et al. 2011; JN635873 KY885068 JQ936626 CƟlƟmbia Amézquita et al. 2013 GECOH 474 Andinobates tolimensis Falan, TƟlima, CƟlƟmbia ƂrƟwn et al. 2011; JN635857 KY885069 JQ936632 Amézquita et al. 2013 GECOH 1523 Andinobates cassidyhornae La Me enia, Jarđín, AntiƟquia, CƟlƟmbia Amézquita et al. JQ936636 KY885054 JQ936622 2013 MVUP 2428 Andinobates geminisae 6 Km ea t Ɵf El Carmen (RíƟ Ƃelén), Ƃati ta et al. 2014 KM212167 KM212166 - Heađwater Ɵf RíƟ CanƟ, DƟnƟ Ɵ, CƟlón, Panamá USNM-FS Andinobates claudiae SƟuth enđ Ɵf I la PƟpa, 1km E Ɵf FrƟ t et al. 2004; DQ283042 DQ502748 DQ502453 59980 SunwƟƟđ Channel, ƂƟca đel TƟrƟ, Grant et al. 2006 Panamá Paratype; ** HƟlƟtype

Andinobates cassidyhornae sp. nov. Dendrobates opisthomelas Silverstone 1975. Quebrada Arriba, bus stop 10 km by road from town of Andes, mountains near road (LACM 71962 - 70). Andinobates opisthomelas Brown & Twomey et al. 2011. pp 33 Plate 3, Figure O. Guatapé Antioquia, Colombia, because of a mistake in the final manuscript. The correct locality according to one of the authors of that paper (D.M.-V.), who actually took the photograph, is Carmen de Atrato, Chocó, Colombia. Holotype: An adult female (Figure 2) deposited in the amphibian collection of the Universidad de los Andes, Bogotá, Colombia, Andes-A 1095 (Field number LUMA 1001) is one of a series collected on March 30 th 2012 at the Mesenia-Paramillo Nature Reserve, Municipality of Andes, Department of Antioquia, Colombia, by R. Medina, G. Suárez & L. Mazariegos. Paratypes: Four adult females, and two adult males (Andes-A 1088–1091, and 1093–1095) collected by R. Medina, G. Suárez & L. Mazariegos. Locality data is the same as the type locality (see below). Type locality. Mesenia-Paramillo Nature Reserve, Vereda La Mesenia, about 12 km south of the municipality of Jardín but politically within the Municipality of Andes, (both in Departamento de Antioquia, Colombia), ca 5 ° 31´N, 75 ° 53´W at 2000 m elevation. Because of the heavy smuggling on dendrobatid frogs we refrain from providing more accurate coordinates. Etymology. This specific epithet cassidyhornae is a patronym in honor of Cassidy Horn, for her passionate interest in poison frogs and her generous contributions to the conservation of cloud forests in Colombia. Definition and diagnosis. A small-sized dendrobatid frog that we assign to the Andinobates bombetes species group (Brown & Twomey et al. 2011) based on the phylogenetic affinity (see Molecular and Phylogenetic Analyses) and on the following morphological characters: adult snout-vent length (SVL) et al. 2011). Andinobates cassidyhornae sp. nov. has an SVL of 19.03 ± 0.31 mm (mean±SD, N = 12 frogs), a bright red dorsum with the color extending onto the upper front and hind limbs; lower forearms and hind limbs are dark brown. Ventral coloration is black with bright red irregularly sized and spaced ovoid or ‘comma’ shaped blotches or spots. It can be externally distinguished from other species in the bombetes group by the distinctive color pattern: in A. cassidyhornae dorsum is bright red, and venter is black with well-defined bright red blotches or spots (Figure 4) vs. (1) in A. opisthomelas dorsum is red often with a posterior suffusion to brown, and venter is black with numerous white spots or reticulation (white-venter form) or venter is chocolate brown sometimes with red suffusion from the flanks (brown-venter form); (2) in A. virolinensis venter is whitish or bluish with black reticulation; (3) in A. bombetes the anterior half of dorsum exhibits bright red, yellow or rarely orange longitudinal and broad dorsolateral stripes; (4) in A. tolimensis the head is yellow, fading to brown towards the dorsum; (5) in A. dorisswansonae the dorsum is black or brown with red blotches, and the venter entirely black or black with few white or yellowish blotches; and (6) in A. daleswansoni the head is entirely red, and the body dull gold or brown. Measurements of the holotype (mm). The holotype measurements correspond to an adult female with a SVL of 18.99 mm; TL of 8.10; HaL of 4.58; HL of 4.85; HW of 6.29; GBW of 7.54; IOD of 2.31; HDT of 1.23; ED of 2.15; TSCN of 1.40; NED of 1.52; IND of 2.36; MTD of 0.75; W 3 FD of 0.78; W 3 F of 0.46; W 3 TD of 0.83; W 3 T of 0.47; W 4 TD of 0.87; and W 4 T of 0.49. The corresponding measurements of all specimens collected are shown in Table 2. Measurements Andes- Andes- Andes- Andes- Andes- Andes- Andes- Median SD A 1093 A 1088 A 1089 A 1090 A 1094 A 1091 A 1095 SVL 19.36 19.34 19.17 18.92 18.45 18.99 18.99 18.99 0.31 TL 8.59 8.52 8.23 8.51 8.50 8.66 8.10 8.51 0.20 HaL 4.59 4.67 4.67 4.60 5.14 4.91 4.58 4.67 0.21 HL 6.06 5.59 5.17 4.75 5.09 5.00 4.85 5.09 0.46 HW 5.38 5.99 5.94 5.75 6.20 5.90 6.29 5.94 0.30 GBW 8.50 8.85 8.13 7.11 7.56 7.20 7.54 7.56 0.66 IOD 2.38 2.30 2.12 1.85 2.26 2.14 2.31 2.26 0.18 HTD 1.41 1.17 0.98 0.96 1.05 0.97 1.23 1.05 0.17 ED 2.49 2.31 2.12 2.45 2.47 2.21 2.15 2.31 0.16 TSCN 1.78 1.34 1.26 1.22 1.54 1.35 1.40 1.35 0.19 NED 1.71 1.68 1.54 1.57 1.78 1.68 1.52 1.68 0.10 IND 2.47 2.33 2.32 2.42 2.84 2.56 2.36 2.42 0.18 MTD 0.84 0.72 0.67 0.60 0.89 0.91 0.75 0.75 0.12 W 3 FD 0.94 0.99 0.85 0.95 1.12 1.10 0.78 0.95 0.12 W 3 F 0.49 0.44 0.48 0.53 0.65 0.52 0.46 0.49 0.07 Description of the holotype: The head is slightly wider than longer, and is narrower than the body. Snout subovoid in dorsal view and truncated in lateral view. Canthus rostralis subovoid, loreal region flat and vertical. Nares situated much closer to the end of the snout than to the eyes, ovoid in shape and directed posterolaterally. Eyes large and prominent with a diameter of 11.3 % of SVL. The pupil is rounded and horizontally elliptical. Tympana and tympanic rings are in the posterolateral regions, ovoid and wider dorsoventrally, measuring 57.2 % of the diameter of the eyes. Supratympanic fold absent. Rounded choanae, not visible in ventral view, as they are completely covered by the maxillary arch. Vomerine, maxillary and premaxillary teeth are absent. Tongue is elongated, almost two times longer than wide; the posterior margin of the tongue is not indented and its posterior third is not adhered to floor of mouth. Hand relatively large (Figure 3), with a length equal to 24.1 % of the snout-vent length. The relative length of the fingers in increasing order of size is: IV The relative length of the toes, in increasing size order is I Coloration of holotype in life: Iris very dark brown; almost indistinguishable from black pupil. Nares encircled by black; margins of upper and lower jaws are very dark brown; tympana are black. Dorsum primarily bright scarlet red, sharply defined along the margins with a few irregularly scattered black speckles and small irregular black markings; in other specimens, black marks may reflect attacks by predators and should therefore not be considered an element of the dorsal coloration. Flanks are red laterally and black ventrolaterally with no gradient merging of the two colors as is seen in A. opisthomelas. Venter ground color black with irregular sharply contrasting bright scarlet red irregularly shaped blotches or spots. Upper forelimbs bright scarlet red and tinged brown where the upper and lower colorations meet, with lower forearm dark brown below the elbow; wrists and hands brown with tips of toes and fingers beige. Thigh mostly red, irregularly speckled with brown; shanks mostly brown tinged with red irregularly (Figure 2). Color in preservative (ethanol 70 %): The dark brown and black colors turn dull black to dark olive in preservative. Discs and tubercules on hands and feet, pupil and cornea become grey or nearly white with time. The predominantly bright scarlet red dorsal coloration and ventral red blotches and spots turn metallic olive; pattern remains clearly distinguishable in preservative (ethanol 70 %). Natural history: The natural history of this species is poorly known. We found individuals in areas covered with a thick layer of leaf litter and where abundant refuges were available. Males call regularly from the leaf litter or hidden amidst tree roots, throughout the day but prominently between 10 h– 14 h and after periods of rain. Most calling males were observed accompanied by a female. We also encountered several courting pairs during a visit in March and April 2012. Males carry 1–3 tadpoles (Figure 5) on their dorsum. Males of other species of the bombetes group are known to release their tadpoles in bromeliad water tanks. Some tadpoles of the new species were found in water within the inflorescence husks of Wettinia palms. Whether this species displays biparental care is unknown. Molecular and phylogenetic analyses. The final alignment consisted of 1119 bp (700 bp unambiguously aligned from the Cytb gene and 419 bp from the 16 S gene). The chosen partitioning strategy was as follows: 16 S unpartitioned under the GTR+ Γ model, and Cytb partitioned by codon position with K 80 +I, HKY+I and GTR+ Γ for the first, second, and third positions, respectively. Both ML and Bayesian phylogenies placed A. cassidyhornae sp. nov. as an independent, well-supported clade, separate from other species, within the bombetes species group. The two sequenced populations, the white- and the brown-venter forms, of A. opisthomelas formed a monophyletic albeit poorly supported group (Figure 6). The topology obtained is roughly consistent with the results of Brown & Twomey et al. (2011), but changes the relationships between A. virolinensis, A. tolimensis and A. bombetes, placing the first two as reciprocally monophyletic clades, and A. bombetes as the sister group of the two. However, the deeper phylogenetic relationships between the species in this group still require further study, since the obtained topology is highly polytomic and not well supported at this level. Within A. cassidyhornae sp. nov., individuals from the Mesenia-Paramillo Natural Reserve, Antioquia, cluster together in a monophyletic group, whereas individuals from other localities do not show evidence of genetic structuring. The SH tests rejected, at a very high level of statistical significance, two out of the three null topologies (H0) that placed A. cassidyhornae sp. nov. within a clade with A. opisthomelas (P A. virolinensis (P A. bombetes (P = 0.05134). Additionally, pairwise genetic distances (K 2 P) between A. cassidyhornae sp. nov. and other species within the bombetes species group ranged between 0.0320–0.0677, well within the range of interspecific distances (Figure 7) observed for the group (0.0216–0.1053), and about tenfold the intraspecific distances observed (0–0.0091). Altogether, the reconstructed phylogenetic tree, the topology (SH) tests, and pairwise genetic distances offer strong support for A. cassidyhornae sp. nov. being a distinct species within the bombetes species group. Bioacoustic analyses. All species’ calls within the Andinobates bombetes species group consist of long and atonal series of pulses sounding like a ‘buzz’ or rattle, which is often longer than 1 sec. The advertisement call of A. cassidyhornae sp. nov. (Figure 8) follows roughly the same pattern. Calls consist of 234.3 ± 20.3 (mean±SD) pulses, last 1.94 ± 0.26 s, and are often uttered as series of calls separated by regular silent intervals of 10.1 ± 2.1 s. The rise time is 50.0± 13.2 % of the call duration. The peak call frequency averages 4.32 ± 0.14 kHz and the frequency bandwidth 0.81 ± 0.40 kHz. Combining the calls of A. cassidyhornae sp. nov., A. opisthomelas (two localities), A. bombetes, and A. virolinensis, we found that call duration (linear regression, R 2 = 0.31, F= 13.9, P= 0.0008, N= 16 males), inter-call interval (R 2 = 0.17, F= 6.5, P= 0.0162), and the number of pulses per call (R 2 = 0.12, F= 4.3, P= 0.0472) decreased at higher temperatures. After removing the temperature effect by calculating regression residuals, the calls of A. cassidyhornae sp. nov. were clearly separated from the calls of the other species in a two-dimensional discriminant space (Figure 9, above, discriminant analysis, Wilks’ Lambda approxim. F= 13.9, PA. cassidyhornae sp. nov. from the call of any other species because the former were lower in peak frequency (F to enter= 39.6, standardized discriminant coefficient= 1.10, PA. cassidyhornae sp. nov. from the call of A. bombetes because the former were longer in duration (F to enter= 6.9, standardized discriminant coefficient= 0.97, P= 0.0013). Regarding pairwise species comparisons by univariate (Figure 9, below) tests, the advertisement call of A. cassidyhornae sp. nov. was lower in peak frequency than the call of A. opisthomelas (Tukey-Kramer HSD test, - 0.97 kHz, PA. virolinensis (- 0.71, PA. bombetes (- 0.53, PA. bombetes. Finally, its frequency bandwidth was wider compared to A. virolinensis (+ 0.40 kHz, P= 0.0220) and A. opisthomelas (+ 0.35 kHz, P= 0.0246). The strong among-species variation in call peak frequency could be partly attributed to concomitant variation in body size. Larger frogs usually produce calls at lower frequency values (e.g. Erdtmann & Amézquita 2009 for dendrobatid frogs) due to allometric constraints in larynx size. Indeed, increasing peak frequency is roughly related to decreasing body size in our study species: A. cassidyhornae sp. nov. is the largest species (mean±SD, 18.73 ± 0.22 mm, N = 5 recorded males, this study), followed by A. bombetes (17.76 ± 0.55, N = 28 males, Myers & Daly 1980), A. opisthomelas (16.80 ± 1.24, N = 26 males, Silverstone 1975), and A. virolinensis (16.72 ± 0.54, N = 127 males, Valderrama-Vernaza et al. 2009). We did not correct for body size effects on peak frequency, because we did not have all information on body size of recorded individuals. In any case, the difference in call frequency alone probably has important evolutionary implications. Across many frog species, ear sensitivity appears to match the peak frequency of the advertisement call (Capranica & Moffat 1983, see Amézquita et al. 2006, 2011 for examples on dendrobatid frogs). Thus, among-lineages differences in call frequency would imply a frequency mismatch between senders and receivers in the mate recognition signal, which could have promoted reproductive isolation between any pair of the Andinobates species we studied here. Distribution, habitat and ecology. At the type locality, the Mesenia-Paramillo Natural Reserve, Andinobates cassidyhornae sp. nov. was found in two fragments of heavily disturbed cloud forests. The approximately 1.52 hectare site is located on a steep montane slope with a gradient of 65 % (Figure 10). This mountain forest fragment is entirely surrounded by cattle grazing grasslands and agriculture crops. The forest fragment there has a dense, sometimes broken canopy with a complex stratification and emergent trees up to 20 m in height. The dominant canopy trees are Lauraceae (Nectandra acutifolia, Nectandra laurel, Aniba coto, Aiouea dubia, Aniba perutiles and Ocotea sp.), Oak (Quercus humboldtii) and Cedar (Cedrela montana). Shrubs and small trees in the families Ericaceae, Gesneriaceae, Melastomataceae, Piperaceae and Rubiaceae dominate the understory. Cyathea sp. tree ferns are common and epiphytes are dense on most trees dominated by the families Araceae, Bromeliaceae, Dryopteridaceae and Orchidaceae. Bromeliads are predominantly of the genus Guzmania. The forest floor is covered with abundant leaf litter and decomposing wood. Remains of inflorescences of Wettinia kalbreyerii palms are common, and serve as terrestrial water reservoirs for extended periods of time. Near daily (primarily evening and early morning) cloud cover of fog, mist and precipitation provide a very cool and humid mesic environment. No additional water sources, i.e. streams or springs were found there. The average annual rainfall is 2500 mm. The average annual regional temperature is 15.0° C. There are two conspicuous wet seasons beginning in March and lasting until May, and another extending from October through December. The species is also known from other three localities visited by one of the co-autors (D. M.-V., Figure 1) that are much less known than the type localitity. They, however, look roughly similar in topography and frogs’ microhabitat. Conservation status. Almost all the known localities for species in the A. bombetes group are within the 1200–2100 m elevational belt. In Colombia, the forests within this range have been severely degraded by intensive agriculture, remarkably coffee plantations, which is a first order national product. Andinobates cassidyhornae sp. nov. was found at four localities within 1800–2059 m elevation (Figure 1). The minimum area of the elevational range encompassing these localities, equivalent to the area achieved by the sum of the occupied grid squares (Figure 2 C in IUCN 2001), is between 200–300 km 2. However, to the best of or experience, the species distribution is sparsely patched including just few of the apparently suitable hills, probably occupying a minor fraction of the available habitat. In addition, most of the suitable forest below 2000 m elevation was cleared since many years ago at the type locality, which further limits the potential distribution of the new species. Unfortunately, most localities included in this study are exposed to severe degradation by intensive agriculture. Contamination of watersheds by pesticides, herbicides, and soi

Amézquita, Adolfo, Márquez, Roberto, Medina, Ricardo, Mejía-Vargas, Daniel, Kahn, Ted R., Suárez, Gustavo, Mazariegos, Luis (2013): A new species of Andean poison frog, Andinobates (Anura: Dendrobatidae), from the northwestern Andes of Colombia. Zootaxa 3620 (1): 163-178, DOI: http://dx.doi.org/10.11646/zootaxa.3620.1.8

Ranitomeya vanzolinii species group Figs. 3, 4, 9, 23 – 29 Tables 1, 4 – 6 A monophyletic assemblage of six species: Ranitomeya vanzolinii Myers 1982; R. sirensis Aichinger 1991 with its junior synonyms R. biolat Morales 1992 and R. lamasi Morales 1992; R. imitator Schulte 1986 with its junior synonyms R. imitator intermedius Schulte 1999 and R. imitator yurimaguensis Schulte 1999; R. flavovittata Schulte 1999; R. yavaricola Perez-Peña, Chavez, Twomey & Brown 2010 and R. cyanovittata Perez-Peña, Chavez, Twomey & Brown 2010. Definition and diagnosis. Medium to large adult SVL (17 – 21 mm); black dorsum; highly variable morphology; LTRF 2(2)/3(1), oral disc emarginated; large intestine unpigmented; larvae whitish to gray; eggs cream; territoriality present in males; promiscuous (known in R. sirensis) or monogamous mating system (known in R. vanzolinii and R. imitator), male parental care (observed in R. sirensis) or biparental (observed in R. vanzolinii and R. imitator); females produce between 2 – 4 eggs per mating. The advertisement call of species in this group consists of a loud trill, highly tonal, with notes 0.4 – 1.1 sec in length, repeated at 2 – 14 notes per minute (Fig. 28)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 61, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Myers, C., W. (1982) Spotted poison frogs: Description of Three new Dendrobates from Western Amazonia, and resurrection of a lost species from \" Chiriqui \". American Museum Novitates, 2721, 23.","Aichinger, M. (1991) A new species of poison-dart frog (Anura: Dendrobatidae) from the Serrania de Sira, Peru. Herpetologica, 47, 1 - 5.","Morales, V. (1992) Dos especies nuevas de Dendrobates (Anura: Dendrobatidae) para Peru. Caribbean Journal of Science, 28, 191 - 199.","Schulte, R. (1986) Eine neue Dendrobates - Art aus Ostperu (Amphibia: Salentia: Dendrobatidae). Sauria, 8, 11 - 20.","Schulte, R. (1999) Pfeilgiftfrosche \" Artenteil - Peru \". INBICO, Wailblingen, Germany, 294 pp.","Perez-Pena, P., Chavez, G., Twomey, E. & Brown, J. L. (2010) Two new species of Ranitomeya (Anura: Dendrobatidae) from eastern Amazonian Peru. Zootaxa, 2439, 1 - 23."]}

Ranitomeya variabilis species group Figs. 3, 4, 9, 14, 32 – 37 Tables 1, 4 – 6 A monophyletic assemblage of two species: Ranitomeya variabilis Zimmermann & Zimmermann 1988 and R. amazonica Schulte 1999. Definition and diagnosis. Medium to large adult size (SVL 14 – 21 mm); two prevalent dorsal patterns: large ovoid black spots on greenish dorsum or complete yellow to red dorsolateral stripes, middorsal stripe and oblique lateral stripes (note that occasionally these species possess ventrolateral stripes, or an intermediate between oblique lateral and ventrolateral stripes, suggesting this trait is a continuous character); large intestine of larvae entirely pigmented; LTRF 2 (2)/3(1), oral disc emarginated; larvae gray, tadpoles cannibalistic on embryos and other tadpoles; eggs gray; neither males nor females territorial; promiscuous mating system with male parental care; females produce between 2 – 6 eggs per mating. Advertisement calls consist of short, regularly spaced buzz calls, notes 0.16 – 0.44 sec in length, repeated at 24 – 70 notes per minute (Fig. 14)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 78, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Zimmermann, H., & Zimmermann, E. (1988) Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfroschen (Anura: Dendrobatidae). Salamandra, 24, 125 - 160.","Schulte, R. (1999) Pfeilgiftfrosche \" Artenteil - Peru \". INBICO, Wailblingen, Germany, 294 pp."]}

Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel, Summers, Kyle (2011): A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083. Zootaxa 3083 (1): 1-120, DOI: 10.11646/zootaxa.3083.1.1, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3083.1.1

Ranitomeya reticulata species group Figs. 3, 4, 9, 15 – 18 Tables 1, 4 – 6 A monophyletic assemblage of six species: Ranitomeya reticulata Boulenger 1884 “1883” with its junior synonym R. ignea Melin 1941; R. fantastica Boulenger, 1884 “1883”; R. ventrimaculata Shreve 1935; R. uakarii Brown, Schulte & Summers 2006; R. summersi Brown, Twomey, Pepper & Sanchez-Rodriguez 2008 and R. benedicta Brown, Twomey, Pepper, & Sanchez-Rodriguez 2008. Definition and diagnosis. Medium to large adult SVL (17 – 21 mm); black dorsum; typically orange to red dorsal patterning; LTRF 2 (2)/3, oral disc emarginate; large intestine entirely pigmented; eggs dark; promiscuous mating system, male parental care; grayish tadpoles and embryos, females produce between 2 – 5 eggs per mating. Vocalizations consist of a series of very short buzz-like notes (0.1 – 0.5 sec in length) given in rapid succession (104 – 200 notes per minute) (Fig. 18)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 48, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Boulenger, G. A. (1884 \" 1883 \") On a collection of frogs from Yurimaguas, Huallaga River, Northern Peru. Proceedings of the Zoological Society of London, 1883, 635 - 638, 1 colour pl, 1 B & W pl.","Melin, D. E. (1941) Contributions to the knowledge of the Amphibia of South America. Goteborgs Kungl. Vetenskaps-och Vitterhetssamhalles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1 - 71.","Shreve, B. (1935) On a new Teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional Papers of the Boston Society of Natural History, 8, 209 - 218.","Brown, J. L., Schulte, R. & Summers, K. (2006) A new species of Dendrobates (Anura: Dendrobatidae) from the Amazonian lowlands in Peru. Zootaxa, 45 - 58."]}

Andinobates minutus species group Figs. 3 – 5 (a – m), 8 Tables 4 –6 A monophyletic assemblage of two divergent species: Andinobates minutus (Shreve 1935) comb. nov. and A. claudiae (Jungfer, Lötters & Jörgens 2000) comb. nov. Definition and diagnosis. SVL 13–16 mm; dark brown or black dorsum; largely complete light colored dorsolateral stripes not extending to thigh and oblique lateral stripes present, typically incomplete and not extending to eye; large spots near body on upper surface of legs and forearms; limbs and venter black with pale marbling on venter; LTRF either 2(2)/3(1) (A. claudiae) or 2(2)/3 (A. minutus); larvae with complete papillae on posterior labium and wide gap in papillae on anterior labium; oral disc emarginated (Table 4); larvae gray; eggs dark (Table 6). Vocalizations short, tonal buzz-notes, notes less than 1 sec in length, repeated at 3 – 17 notes per minute (Fig. 8, Table 5).

Andinobates minutus species group Figs. 3 – 5 (a – m), 8 Tables 4 –6 A monophyletic assemblage of two divergent species: Andinobates minutus (Shreve 1935) comb. nov. and A. claudiae (Jungfer, Lötters & Jörgens 2000) comb. nov. Definition and diagnosis. SVL 13–16 mm; dark brown or black dorsum; largely complete light colored dorsolateral stripes not extending to thigh and oblique lateral stripes present, typically incomplete and not extending to eye; large spots near body on upper surface of legs and forearms; limbs and venter black with pale marbling on venter; LTRF either 2(2)/3(1) (A. claudiae) or 2(2)/3 (A. minutus); larvae with complete papillae on posterior labium and wide gap in papillae on anterior labium; oral disc emarginated (Table 4); larvae gray; eggs dark (Table 6). Vocalizations short, tonal buzz-notes, notes less than 1 sec in length, repeated at 3 – 17 notes per minute (Fig. 8, Table 5)., Published as part of Brown, Jason L., Twomey, Evan, Amézquita, Adolfo, Souza, Moisés Barbosa De, Caldwell, Jana- Lee P., Lötters, Stefan, May, Rudolf Von, Melo-Sampaio, Paulo Roberto, Mejía-Vargas, Daniel, Perez-Peña, Pedro, Pepper, Mark, Poelman, Erik H., Sanchez-Rodriguez, Manuel & Summers, Kyle, 2011, A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083, pp. 1-120 in Zootaxa 3083 (1) on page 30, DOI: 10.11646/zootaxa.3083.1.1, http://zenodo.org/record/5244214, {"references":["Shreve, B. (1935) On a new Teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional Papers of the Boston Society of Natural History, 8, 209 - 218.","Jungfer, K. - H., Lotters, L. & Jorgens, D. (2000) Der kleinste Pfeilgiftfrosch - eine neue Dendrobates - Art aus West-Panama. Herpetofauna, 22, 11 - 18."]}