A new species of Dendrobates (Anura: Dendrobatidae) from the Amazonian lowlands in Perú Author Brown, Jason L. Author Schulte, Rainer Author Summers, Kyle text Zootaxa 2006 1152 45 58 journal article 50621 10.5281/zenodo.172186 b6e040b8-f21a-4fec-93a0-c02c193ad8cb 1175­5326 172186 Dendrobates uakarii , sp. nov. Holotype . MHNSM 23246, (Field number MC0140), an adult female collected on 20 June 2004 by Mario Callegari upstream Quebrada Blanco in Tamshiyacu­Tahuayo Reserve, Departamento Loreto, Peru ( 4º 11’21.88” S , 73º6’15.66” W ), Elevation: 140 m . Paratypes . MHNSM 23247­23250, (Field numbers MC0141­MC0144), adults collected on 20 June 2004 by Mario Callegari upstream Quebrada Blanco in Tamshiyacu­ Tahuayo Reserve, Departamento Loreto, Peru ( 4º 11’21.88” S , 73º6’15.66” W ), Elevation: 140 m . Diagnosis A species of Dendrobates as characterized by the first finger, which is shorter than the second, the lack of webbing between the toes, absence of premaxillary and maxillary teeth ( Myers 1982 ). Dendrobates uakarii , sp. nov. is a species of small size ( 16.16 mm SVL in female holotype ); with smooth skin; paired­parallel dorsolateral lines: one yellow oblique lateral and one red dorsolateral (on each flank), with one vertebral line, which is the same color of the dorsolateral lines; black coloration between the dorsolateral lines and the vertebral line generally creates a “U” around the snout. This species can be distinguished from its sister taxa on the basis of 19 unique mitochondrial gene nucleotide site substitutions ( Fig. 4 ). The new species can be distinguished from all known species of Dendrobates occurring in the Amazonian lowlands by the presence of differently colored parallel dorsolateral lines (as described above). The new species is similar in appearance to D. duellmani and D. ventrimaculatus . Dendrobates uakarii , sp. nov. is distinguished from D. ventrimaculatus sensu strico and D. ventrimaculatu sensu lato ( Fig. 2 ) by: complete parallel dorsolateral stripes; red to orange red in life (vs. yellow to orange­red “ Y ” to single incomplete vertebral stripe on dorsum), presence of a single broad oblique lateral stripe (vs. often connecting, or fine single oblique lateral stripe) and presence of black “U”, which is formed from the vertebral and dorsolateral lines. Dendrobates uakarii , sp. nov. is distinguished from D. duellmani by: dorsolateral stripe red in life (vs. reddish pink), and broad yellow oblique lateral stripe present (vs. fine blue oblique stripe, matching the color of reticulation on limbs, Fig. 2 ). Description of Holotype Size small, SVL 16.16 mm ( Table 1 ). Widest part of head between eyelids. Body slightly wider than head. Greatest head­width 30.6% of SVL. Eyes very protuberant. Tongue small, oval. Teeth absent. TABLE 1. Measurements of D.uakarii .

SVL	MHNSM 23246 16.16	MHNSM 23247 15.47	MHNSM 23248 14.79	MHNSM 23249 15.70	MHNSM 23250 14.82	MHNSM** 23251 13.04	Average ± StDev*** 15.39 ± 0.59
FL	6.46	6.32	6.24	6.44	6.27	5.36	6.35 ±0.10
TL KK	7.10 12.70	7.11 13.19	7.02 12.87	7.05 13.62	6.71 12.80	5.82 11.35	7.00 ±0.17 13.04 ±0.37
FoL	6.08	6.13	5.96	6.01	5.77	4.75	5.99 ±0.14
HaL HL	4.11 5.49	3.82 5.43	3.63 5.24	3.83 4.94	3.61 4.89	3.08 4.40	3.80 ±0.20 5.20 ±0.27
±HW	4.95	5.27	5.08	5.17	4.92	4.52	5.08 ±0.15
BW TD*	5.88 0.58	5.52 0.49	5.04 0.62	5.84 0.62	4.88 0.53	4.61 0.44	5.43 ±0.46 0.57 ±0.06
ED*	1.47	1.60	1.64	1.51	1.60	1.42	1.56 ±0.07
IOD* L1F*	2.27 1.20	2.27 1.07	2.22 1.24	2.35 1.24	2.13 1.02	1.96 0.89	2.25 ±0.08 1.15 ±0.10
L2F*	1.69	1.47	1.64	1.69	1.47	0.93	1.59 ±0.11
DET* UEW*	0.58 1.38	0.58 1.29	0.58 1.33	0.62 1.24	0.53 1.38	0.49 0.98	0.58 ±0.03 1.32 ±0.06
SEX	Ψ	ɗ	ɗ	Ψ	ɗ	–

* Measurements taken with dissecting microscope **sub­adult, not used in description *** MHNSM 23251 not included FIGURE 1. Maximum likelihood phylogram derived from a Bayesian backbone constraint consensus tree constructed using only taxa for which 12S, 16S, cytochrome b and cytochrome oxidase I sequence data were available. Numbers indicate posterior probabilities from the Bayesian analysis. Species of the Ventrimaculatus group are denoted with s.s. (sensu stricto), s.l. (sensu lato) and sp. aff (species affinis). FIGURE 2. A photographic illustration comparing D. uakarii , sp. nov. to similar species. D. uakarii , sp. nov. (holotype) from Tamashiyacu­Tahauyo Communal Reserve ( A. Dorsum B. Venter). Near Rio Yarapa, south of Rio Tahuayo, populations have thinner dorsolateral and vertebral lines with brighter red stripes ( C. photo: Devon Graham). D. D. duellmani, Yasuni , Ecuador (photo: Petra Bartelds). E. D. ventrimaculatus sp. aff. Solimoes, Amazonas, Brazil (photo: Janalee P. Caldwell). F. D. ventrimaculatus sensu lato , Tamashiyacu­Tahauyo Communal Reserve, Peru. G. D. reticulatus, Iquitos , Peru (photo: Evan Twomey). Skin in preservative completely smooth on all surfaces of body. Snout sloping laterally; bluntly rounded dorsally; truncate ventrally. Nares situated and directed laterally to the tip of snout; both nares barely visible from front, well from below but not from above. Canthus rostralis rounded; loreal region sloping and flat or slightly concave. Interorbital distance 1.7 times wider than upper eyelid. Tympanum circular, half concealed posterodorsally (7 o’clock to 2 o’clock, when center of snout is at 3 o’clock), without tympanic annulus, its area less than 50% of ED. Hands relatively small, length being 25.4% of SVL. Relative length of appressed fingers 1<2<4<3; finger I slightly shorter than finger II. Discs conspicuously expanded on all fingers but finger I. In adults disc on finger III is 2 times wider than distal end of adjacent phalanx. A large, circular outer metacarpal tubercle on median base of palm; a smaller inner metacarpal tubercle on base of finger I; one prominent subarticular tubercle on fingers I, II, and IV, two on finger III. Hind limbs relatively short, with heel of appressed limb reaching the tympanum. Tibia 43.9% of SVL. Relative lengths of appressed toes 1<2<5<3<4; first toe short (but conspicuously present), barely reaching bottom of subarticular tubercle on base of second toe, with unexpanded disc; and toes II, III, and IV barely expanded (much smaller than finger discs), and toe IV expanded (disc 1.5 broader than adjacent phalanx). Moderatesized inner and small outer metatarsal tubercles, somewhat protuberant with rounded surfaces. One slightly protuberant subarticular tubercle on toes I, II and V, two on toe III, and three on toe IV. Hands and feet lacking supernumerary tubercles, lateral fringes and webbing. FIGURE 3. Call comparisons of closely related species. The data is listed as following: species (ambient temperature, location of individual, mean frequency). A. D. uakarii (26°C, Tahauyo, 3780 hz). B. D. ventrimaculatus sensu lato (24°C, Iquitos, 4900 hz) C. D. fantasticus (24°C, Iquitos, 3350 hz). D. D. reticulatus (29°C, Iquitos, 4130 hz). Color in life (based on photographs of a living specimen, Fig. 2 ): dorsum and upper flanks black, with paired dorsolateral lines: one yellow oblique lateral and one red dorsolateral (on each flank); and one vertebral line, similar in color to the dorsolateral lines. The paired dorsolateral stripes are bright reddish­orange, which continue on the exterior edges of the eyelids, the canthus rostralis, and the superior part of the snout. The vertebral line spans from the preorbital to the sacral region. The paired oblique lateral lines extend from the groin to the upper forelimbs around the upper lip. Limbs, hands, feet, and belly blue with a fine reticulum of black dots and larger spots. Throat yellow, with paired black spots below the mandible, and large black gular spot. Iris black. In 70% alcohol the color is almost identical to the living animal described above. The only differences are that the dorsolateral stripes and vertebral stripe changed to pinkishyellow, and oblique lateral stripes and throat changed to silver­white. TABLE 2. Species names, collection localities, and GenBank accession numbers (where applicable) for taxa included in the analysis.

Species	Location	12S	16S	COI	CytB
D. amazonicus	Iquitos, Loreto, Peru	AF482770	AF482785	AF482815	AF482800
D. duellmani	Yasuni, Ecuador	AY364566	AY263246	NA	NA

D. fantasticus	N. Sauce, San Martin, Peru	AF412444	AF412472 AF412416	AF412500
D. fantasticus	Cainarachi, San Martin, Peru	AF412447	AF412475 AF412419	AF412503
D. imitator	Huallaga, San Martin, Peru	AF412448	AF412476 AF412420	AF412504
D. reticulatus	Punte Itaya, Loreto, Peru	AF482772	AF482787 AF482817	AF482802
D. reticulatus	B. Achille, Loreto, Peru	AF482771	AF482786 AF482816	AF482801
D. reticulatus DARST	Peru	AY326029	AY326029 NA	NA
D. reticulatus KS	Peru	NA	AF412467 AF412411	AF412495
D. reticulatus SANTOS	Peru	AY364567	AY364567 NA	NA
D. reticulatus KS	Peru	AF412439	AY263245 NA	NA
D. sp.	Puente Itaya, Peru	AF482777	AF482792 AF482821	AF482807
D. sp. G.	Ecuador	AY364568	AY364568 NA	NA
D. uakarii	Tahuayo, Loreto, Peru	DQ371305	DQ371316 DQ371325	DQ371335
D. variabilis	Cainarachi, San Martin, Peru	AF412463	AF412491 AF412435	AF412519
D. ventrimaculatus s.l.	N. Bonilla, San Martin, Peru	AF412466	AF412494 AF412438	AF412522
D. ventrimaculatus s.l.	Near Rio Napo, Loreto, Peru	AF482781	AF482796 AF482825	AF482811
D. ventrimaculatus s.l.	Ituxi, Acre, Brazil	Noonan and Wray 2006
D. ventrimaculatus s.l.	French Guiana	DQ371302	DQ371313 DQ371325	DQ371332
D. ventrimaculatus sp. aff. D. ventrimaculatus s.l. 1 D. ventrimaculatus s.l. 2	Solimoes, ΑmƋzοnƋsʹ ΒrƋzⅰ| Solimoes, ΑmƋzοnƋsʹ ΒΓƋzi| Solimoes, ΑmƋzοnƋsʹ ΒΓƋzi|	DQ371307 DQ371308 DQ163089	DQ371318 DQ371327 DQ371319 DQ371328 DQ163079 DQ163063	DQ371337 DQ371338 DQ163074
D. ventrimaculatus sp. aff.	Porto Walter, Acre, Brazil	AF482782	AF482797 AF482826	AF482812
D. ventrimaculatus s.l. 1	Porto Walter, Acre, Brazil	DQ371301	DQ371312 DQ371322	DQ371331
D. ventrimaculatus s.l. 2	Porto Walter, Acre, Brazil	AF482783	AF482798 AF482827	AF482813
D. ventrimaculatus s.s. 1	Ecuador	AF482780	AF482795 AF482824	AF482810
D. ventrimaculatus s.s. 2	Ecuador	AF128620	AF128619 AF097502	AF120013

Etymology The red uakari Dendrobates . The specific epithet is a patronym for the red uakari ( Cacajao calvus ucayalii ), an endangered Amazonian primate. Both black ( C. melanocephalus ) and red uakari ( C. calvus ) are probably the primate species at greatest risk in South America (CITES). Because these monkeys are quite large and prefer flooded forest habitats, they are easily hunted from boats. The epithet was chosen because both the red uakari and D. uakarii , sp. nov. shared similar historical distributions and have bright red dermal pigmentation. FIGURE 4. Character transformation for Dendrobates uakarii , sp. nov. diagnostic nucleotide site substitutions using ACCTRAN optimization. Alignment available online at http://www. dendrobates .org/permanent/duakarii.zip Va r i a t i o n Within populations the black between the dorsolateral lines and vertebral line may form a “U” around the snout (77%, n=13). Other individuals have a connected dorsolateral line and vertebral line near the canthus rostralis (23%, n=13). The black gular spot is absent in some individuals (7%, n=13). Coloration and width of the dorsolateral and vertebral lines vary between populations. Near Rio Yarapa, south of Rio Tahuayo, populations have thinner dorsolateral and vertebral lines, are yellow near the sacrum (almost the same color as oblique lateral stripes), and change to brick­red near the occiput ( Fig 2 C). We amplified regions of the cytochrome b gene from two individuals from the Tamshiyacu­Tahuayo Reserve and uncovered only one haplotype; although the sample size is very small, this suggests that intrapopulation variation may be minimal. Distribution and ecology We found Dendrobates uakarii , sp. nov. throughout the Tamshiyacu­Tahuayo Community Reserve and further south to Pacaya Samiria Reserve, on the Rio Yarapa. Its distribution is likely between Rio Amazonas, Rio Javari, and Rio Ucayali (north of the Contamana­Sierra Divisor Arch) in western Brazil and eastern Peru . This putative distribution is supported by Christmann’s (2004) book, which contains pictures of a similar looking species 200 km east of Iquitos, south of Rio Napo, and further east to the tributaries on the south side Amazon, near the Brazilian border. In Tahuayo, it occurs sympatrically with two other species of Dendrobates : D. ventrimaculatus sensu lato and D. flavovittatus . D. uakarii , sp. nov. occurs in primary forests. It spends a majority of its time on the forest floor, however it occasionally ventures a few meters into the canopy. The call is a long series of soft buzz­like notes, similar to D. fantasticus , but with shorter pauses between each note, making the call sound nearly continuous ( Fig 3 ). Males have been observed to carry one to four tadpoles to large phytotelmata such as bromeliads.